A Natural History of the Ground-Beetles (Coleoptere: Carabidae) of American North of Mexico (Pensoft Series Faunistica, 27)

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A Natural History of the Ground-Beetles (Coleoptera: Carabidae) of America north of Mexico

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A. Larochelle & M.-C. Larivière

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Colliuris pensylvanica (Linné) Photo: Dr Henri Goulet, Agriculture Canada, Ottawa

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ISBN 954-642-165-0

A. Larochelle & M.-C. Larivière

Pensoft Series Faunistica No 27

A Natural History of the Ground-Beetles (Coleoptera: Carabidae) of America north of Mexico

A summary of the natural history of the ground-beetles (Coleoptera: Carabidae; over 2400 taxa) from North America north of Mexico. Under each taxon, the ecology, biology, dispersal power, collecting techniques, and selected references are given. The section on ecology comprises the following: altitudinal distribution, habitat, diel activity, and gregariousness. The section dealing with biology gives details on seasonality, mating, gravid females or oviposition, tenerals, overwintering, feeding, predation, parasitism, and defense-mechanism. The section on dispersal power, or the capability of dispersal, has been assessed when possible, using three main criteria: wing condition, flight data (including light-trapping observations), and other locomotory habits. The section on collecting techniques provides the best means of capture. The section references lists the most relevant papers, with appropriate key-words. An exhaustive bibliography dealing with the natural history of North American Carabidae is also provided. This work follows the Catalogue of Bousquet and Larochelle (1993) Catalogue of the Geadephaga (Coleoptera: Trachypachydae, Rhysodidae, Carabidae including Cicindelini) of America north of Mexico, which provided nomenclatural and distributional information on North American ground-beetles. An indispensable tool for anybody interested in ground-beetles, as well as to libraries due to the incredible amount of information summarised in this book.

A natural history of Carabidae (Coleoptera) of America north of Mexico A NATURAL HISTORY OF THE GROUND-BEETLES (COLEOPTERA: CARABIDAE) OF AMERICA NORTH OF MEXICO André Larochelle and Marie-Claude Larivière

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A natural history of Carabidae (Coleoptera) of America north of Mexico

A Natural History of the Ground-Beetles (Coleoptera: Carabidae) of America north of Mexico André Larochelle & Marie-Claude Larivière

Sofia-Moscow 2003

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A NATURAL HISTORY OF THE GROUND-BEETLES (COLEOPTERA: CARABIDAE) OF AMERICA NORTH OF MEXICO © André Larochelle1 and Marie-Claude Larivière2 New Zealand Arthropod Collection Landcare Research Private Bag 92170 Auckland, New Zealand. E-mail: [email protected], [email protected]

Pensoft Series Faunistica No 27 ISSN 1312-0174

First published 2003 ISBN 954-642-165-0

© PENSOFT Publishers All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form by any means, electronic, mechanical, photocopying, recording or othrewise, without the prior written permission of the copyright owner.

Pensoft Publishers, Acad. G. Bonchev Str., Bl.6, 1113 Sofia, Bulgaria Fax: +359-2-70-45-08, e-mail: [email protected], www.pensoft.net Printed in Bulgaria, January 2003

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Foreword

“Two roads diverged in a wood, and I – I took the one less traveled by, And that has made all the difference”. Robert Frost, The Road Not Taken (1915)

I take great pleasure in dedicating this work to the memory of Carl H. Lindroth (1905-1979), my late mentor and friend, a naturalist interested in understanding carabid life in all its manifestations. I was twenty-two years old and a fervent beetle collector, devoid of systematic knowledge, exploring the natural world day in, day out, when I was invited by the famous odonatologist Adrien Robert from the Université de Montréal to specialize in carabid beetles (then abundant in my insect collection) and to contact Carl H. Lindroth, a Swedish scientist, who had started to revise the fauna of Canada and Alaska. Without hesitation, I wrote him a letter, confessing my desire to specialize and requesting his invaluable help. After a short time, I received an encouraging letter and two of his major works entitled: Die Fennoskandischen Carabidae: Eine tiergeographische Studie (1945 and 1949) and The ground-beetles (Carabidae, excl. Cicindelinae) of Canada and Alaska (Part 2, 1961). Those works became to me daily tools. I was immediately conquered by Lindroth’s publications which would eventually include five other sections on Canada and Alaska. I would also be very much inspired fifteen years later by Thiele’s Carabid beetles in their environments (1977). The kind attention of a world authority such as Carl H. Lindroth changed the course of my life. I was destined to become a carabid naturalist. Through the years I became increasingly impressed by Carl’s friendly manner, generosity, self-discipline, creativity, and modesty. He was a single-minded man who led an unfragmented life, devoting job and hobby to reaching his goals. Carl was held in profound respect by the scientific community. He devoted his career (1952-1979) to the study of the geographic distribution of carabids and to the taxonomic revision of the fauna of Canada and the adjacent territories. As a naturalist, he practised a multidisciplinary methodology, attaching as much importance to field surveys, direct observations in nature, observations and experiments in the laboratory, and study of entomological museums and collections, as to the world literature. His user-friendly identification keys and attrac-

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tive presentation of natural history are still being used on a regular basis by both professional and amateur entomologists. Good overviews of his life and scientific contribution can be found in Darlington (1971a, 1971b), Steffan and Thiele (1980), and Ball (1981). My encounter with Carl H. Lindroth encouraged me to collect carabids every day, in all habitats, localities and weather conditions, and to record and publish my own observations, however small. By profession a secondary school teacher, I also decided that the study of carabid natural history would become the very fabric of my life. From that period until 1992 (when I moved to New Zealand), I explored North American nature, visited countless museums, collections and carabidologists, observed carabids in the laboratory, reviewed the North American literature, and published several papers on natural history and distribution. The results of my 40-year research on the geographic distribution of North American carabids were published in Bousquet and Larochelle’s 1993 Catalogue (Memoirs of the Entomological Society of Canada, No. 167). This natural history was likewise initiated 40 years ago by myself and subsequently carried on in collaboration with my wife MarieClaude. It is intended as a continuation of the 1993 Catalogue. In future too, Lindroth’s work will be my guide. I very likely face a challenge similar to his when he dealt with the fauna of Canada and Alaska. I must now revise a largely undescribed fauna in order to obtain the necessary bases for the study of its natural history and biogeography. André Larochelle Auckland, N.Z.

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Introduction

Carabids, or ground-beetles, are a highly adaptable group which has managed to occupy most terrestrial habitats, from tropical rainforests to semi-arid deserts, on nearly all continents and most islands since their origin in the early Mesozoic. They are rich in species, abundant in individuals, often attractive in coloration, which makes them popular objects of study in professional and amateur circles. As a family they exhibit a relatively high degree of morphological uniformity which makes these insects suitable to study the ecophysiological adaptations required to cope with environmental demands. Being sensitive to their environment, they demonstrate a flexible set of responses to both abiotic and biotic factors. As bioindicators, carabids have the added advantage to be increasingly better known taxonomically. Most carabids are omnivorous feeders, mostly predators (Larochelle, 1990). The majority of species are nocturnal and live at the surface of the ground. Some species live in the soil, in caves, or on the vegetation. In general, ground-beetles show good dispersal capabilities. Carabids are relatively easy to sample by reliable quantitative methods, especially pitfall trapping. These and other factors have made ground-beetles favourites with biologists investigating evolutionary and ecological hypotheses. As a result they are commonly used to: (a) assess the biodiversity of ecosystems, (b) indicate the impact of landscape changes, (c) evaluate environmental health, (d) predict the effect of climate changes, (e) classify habitats for nature protection, (f) characterise soil-nutrient status in forestry, and (g) control pest invertebrates, e.g., lepidopterous caterpillars (Lövei and Sunderland, 1996; Brandmayr et al. 2000; Holland, 2002). Carabids may become more commonly used in the future in biological and integrated pest management programs, e.g., as natural control agents of noxious invertebrates, especially soil insect pests, or of weeds, especially their seeds (Rivard, 1966; Tomlin, 1975b; Lund and Turpin, 1977b; Allen, 1979; Morrill, 1992). The Carabidae of America north of Mexico comprise over 2400 species and subspecies (Ball and Bousquet, 2001) and are among the best represented groups of insects in entomological museums and collections. Despite such high interest in this group, no modern effort has been made to summarize their natural history. A few papers have already summarized natural history information for parts of the fauna: Lindroth (1955a and 1963a for NF; 1961-1969 for Canada and Alaska) and Erwin (1981 for the Plummers Island, MD). Most known information, however, remains scattered through the literature. The present publication is an attempt to bring together all

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knowledge available from the literature, the authors’ own observations, personal communications from colleagues, and data from collections and museums. In North America, tiger beetles are generally regarded as a family distinct from the Carabidae. Recent phylogenetic evidence (Pearson and Vogler, 2001) strongly supports this view which the authors follow here. Consequently, Cicindelidae are not covered in this work. A Natural History of the tiger beetles of America north of Mexico has been published separately in the specialized journal Cicindela (Larochelle and Larivière, 2001b). This Natural History is intended as a reference tool for a wide audience, e.g., carabidologists, other entomologists, ecologists, conservation managers, applied biologists, as well as the general naturalist public.

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Methods

From 1962 to 2002, data were gathered from a comprehensive review of the literature, visits of entomological collections, consultations with colleagues, observations in the laboratory, and field surveys. This information was subsequently summarized on data sheets compiling the species profiles for ecology, biology, and dispersal power, following the method used by Larochelle and Larivière (2001a). In order to eliminate spurious records, an effort was made to summarize available information by using the smallest common denominator representing the essentials of each species’ requirements. In this Natural History, information is presented in a manner similar to that of previous authors (e.g., Lindroth, 1945, 1961-1969; Koch, 1989), in a catalogue format with taxa listed alphabetically by genera and species to provide quicker access to data. The valid names of taxa follow Bousquet and Larochelle (1993) unless there have been subsequent changes which are indicated in the text with appropriate references. Subgeneric names have been added as a taxonomic aid, but are not necessarily listed alphabetically. Unless otherwise stated, data refer to the adult stage. When information relates to particular geographic entities, these are expressed as two-letter abbreviations as in Table 1. The section on ecology comprises the following: altitudinal distribution, habitat, diel activity, and gregariousness. Altitudinal distribution, or distribution related to altitude or elevation, is expressed as lowlands, mountains or uplands, subalpine and alpine zones. The section dealing with biology gives details on seasonality, mating, gravid females or oviposition, tenerality, overwintering, feeding, predation, parasitism, and defensemechanism. Seasonality, or the period when an animal is active, is expressed as months of the year. The section on dispersal power, or the capability of dispersal, has been assessed using three main criteria: wing condition, flight data (including light-trapping observations), and other locomotory habits. The wing condition was evaluated for each species based on the literature, and personal observations in the field and in the laboratory. The locomotory habits of species has been recorded under three categories: fast runner, moderate runner, and slow runner; each species has been assigned to one of these categories using field observations and examination of leg characteristics (Evans, 1977 and 1986; Forsythe, 1981 and 1983). In general, fast runners have longer, thinner legs and occur in open, bare habitats, e.g., Nebria species. Moderate runners have moderately long, moderately large legs, and live generally in leaf litter or on low vegetation. Slow runners have shorter, more

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inflated legs armed with tooth-like excrescences, and dig burrows, e.g., Broscini and Clivinini. For flight and climbing data, three categories were used: occasional (only a few specimens and instances observed), regular (a moderate number of specimens and instances), and frequent (numerous specimens and instances). Indirect evidence of flight was also provided in the case of specimens found in shore-drift material. The section on collecting techniques enumerates, in order of decreasing importance, the best methods for collecting each species. Although most carabids can be secured by hand collecting, special techniques are often required to obtain large populations or quantitative samples for scientific research. For example, these include: pitfall trapping; treading vegetation into water; raking or sifting leaf litter; turning embedded logs, stones, moss carpets, reindeer moss (Cladonia) or plant rosettes (Verbascum); digging at the base of plants; raking loose gravel; pouring water over ground; treading soil with the feet; sweeping or beating the vegetation; lifting the loose bark of logs and fallen trees; dismantling logs; using malaise traps, rotary traps, interception traps, window traps, or tanglefoot screens; collecting with a headlamp at night; fogging the canopy; pyrethrum spraying; smoking stumps; sugaring trees; sifting fermented sawdust or garden compost; inspecting soil crevices, the tunnels of small rodents or the walls of beaver houses; and turning drift material along the seashore, lake shores or river banks. Pitfall trapping is, however, the most commonly used method for evaluating the presence, locomotory activity, and community composition of carabids. The section references under each taxon enumerates the most relevant papers, with appropriate key-words. Table I. Two-letter abbreviations of geographic entities covered in this paper. AB AK AL AR AZ BC CA CO CT DC DE FL GA GL IA ID IL IN KS KY LA LB

Alberta Alaska Alabama Arkansas Arizona British Columbia California Colorado Connecticut District of Columbia Delaware Florida Georgia Greenland Iowa Idaho Illinois Indiana Kansas Kentucky Louisiana Labrador

MA MB MD ME MI MN MO MS MT NB NC ND NE NF NH NJ NM NS NT NV NY OH

Massachusetts Manitoba Maryland Maine Michigan Minnesota Missouri Mississipi Montana New Brunswick North Carolina North Dakota Nebraska Newfoundland New Hampshire New Jersey New Mexico Nova Scotia Northwest Territories Nevada New York Ohio

OK ON OR PA PE PM QU RI SC SD SK TN TX UT VA VT WA WI WV WY YT

Oklahoma Ontario Oregon Pennsylvania Prince Edward Island St. Pierre and Miquelon Québec Rhode Island South Carolina South Dakota Saskatchewan Tennessee Texas Utah Virginia Vermont Washington Wisconsin West Virginia Wyoming Yukon Territory

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Acknowledgments

We are indebted to the following people who have contributed precious personal observations on natural history: G.E. Ball (University of Alberta, Edmonton), T.C. Barr Jr (University of Kentucky, Lexington), R.T. Bell (University of Vermont, Burlington), Y. Bousquet (Agriculture and Agri-Food Canada, Ottawa, Ontario), R.L. Davidson (Carnegie Museum, Pittsburgh, Pennsylvania), S. Elias (University of Colorado, Boulder), F. Génier (Musée de la Nature, Hull, Québec), H. Goulet (Agriculture and Agri-Food Canada, Ottawa, Ontario), R. C. Graves (University of Michigan, Bowling Green), D. Hildebrandt and M. Plonczynski (Jackson, Mississipi); H.J. Lee, Jr (Fairview Park, Ohio), F. Liard (Joliette, Québec), J.K. Liebherr (Cornell University, Ithaca, New York), and P. Paquin (Université de Montréal, Montréal, Québec). Thanks are extended to present and past library staff from the MacDonald Campus of McGill University (Ste-Anne-de-Bellevue, Québec), Agriculture and AgriFood Canada (Ottawa, Ontario), and HortResearch (Auckland, New Zealand) for their valuable help with the literature search. We are also grateful to Y. Bousquet for allowing us to consult his carabidological library and L. LeSage (Agriculture and Agri-Food Canada, Ottawa, Ontario) for providing copies of papers unavailable in New Zealand. We are very grateful to A. Weir and M.B. Hughes (State University of New York, Syracuse) who have made available extensive unpublished records from the late J. Balazuc on Laboulbeniales. We also wish to thank the following colleagues for their helpful comments and suggestions for improving the manuscript: Y. Bousquet (Agriculture and Agri-Food Canada, Ottawa, Ontario) and R.B.J. Hoare (Landcare Research, Auckland, New Zealand). We express our gratitude to H. Goulet (Agriculture and Agri-Food Canada, Ottawa, Ontario) for the frontispiece photo. Finally, we are most obliged to Landcare Research’s science management, particularly R.E. Beever, P.K. Buchanan, T.K. Crosby, O.R.W. Sutherland, and for their encouragements and for allocating resources towards completion of this work. Most of this research was supported by the authors’ own resources. It has required almost 40 years of intensive work by the first author and part-time contribution by the second author. Through the course of this study, we have been repeatedly impressed by the generosity and the cooperative mind of our fellow entomologists. Most of them have wel-

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comed our efforts and have not seen us as intruders into the right owner’s territory. We think highly of their open-mindedness and we sincerely thank all of them.

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Species accounts

Abaris splendidula (LeConte, 1863) Ecology. Lowlands and mountains. Open forests (tropical and subtropical; e.g., acacia). Open ground; moderately moist, sandy-clayish soil. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: January, July, November. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Abax parallelepipedus (Piller and Mitterpacher, 1783) (Abax ater (Villers)) Ecology. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: June. Brood care exhibited by female. Adults found overwintering under logs and in leaf litter (Europe). Adult food, in the field (Europe): lepidopterous caterpillars, springtails, flies, mites, earthworms, slugs, and small dead animals. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. References. Lindroth, 1945 (natural history); Drift, 1951 (biology); Greenslade, 1965 (biology); Löser, 1972 (biology); Loreau, 1985 (biology; as ater Villers); Lindroth, 1986 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology). Acupalpus (Philodes) alternans (LeConte, 1853) Ecology. Lowlands. Stream edges. Open ground; moist or wet, sandy or muddy soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: March, MayAugust. Dispersal power. Wing-condition unknown. Moderate runner. Collecting technique. Turning stones. Reference. Lindroth, 1968 (ecology). Acupalpus (Acupalpus) canadensis Casey, 1924 Ecology. Lowlands. Borders of eutrophic marshes; swamps, marshy borders of ponds, lakes, and slow rivers; wet meadows. Open or shaded ground; poorly drained, soft, wet, clayish or muddy soil, rich in organic debris, covered with dense vegetation. Nocturnal; sheltering during the day in debris. Biology. Seasonality: January, March-September, November-December. Adults found overwintering on higher and drier ground, at the edge of woods and on the top of sand dunes; in leaf litter and rotten logs. Dis-

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persal power. Macropterous. Frequent flier (at dusk; to artificial lights, in captivity). Moderate runner. Occasional climber (on plants). Favored by spring floods along large rivers. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping; sweeping plants. References. Lindroth, 1963a and 1968 (natural history). Acupalpus (Acupalpus) carus (LeConte, 1863) Ecology. Lowlands. Borders of marshes (e.g., Salix, Typha); swamps, swamp-forests, margins of irrigation ditches, marshy borders of ponds and lakes. Shaded ground; wet clay soil, rich in organic debris, covered with dense vegetation. Mostly nocturnal; usually sheltering during the day under dead leaves. Biology. Seasonality: January-December. Tenerals: April, June, August. Adults found overwintering on higher and drier ground, at the edge of woods and swamps, in forests and hedges; under dead leaves, stones, fallen trees, in moss and clumps of grass. Parasites: Laboulbeniales (Fungi). Dispersal power. Wing-dimorphic: macropterous (capable of flight) or brachypterous (incapable of flight). Frequent flier (at dusk; to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Favored by spring floods along large rivers. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping; sweeping plants. References. Blatchley, 1910 (ecology); Lindroth, 1963a and 1968 (natural history); Kirk, 1975c (dispersal power). Acupalpus (Philodes) flavilimbus (LeConte, 1868) Ecology. Lowlands and mountains. Biology. Seasonality: May. Dispersal power. Wingcondition unknown. Moderate runner. Acupalpus (Acupalpus) hydropicus (LeConte, 1863) Ecology. Lowlands. Deciduous forests, cranberry bogs, and roadside ditches. Wet soil. Nocturnal; sheltering during the day under dead leaves and stones. Biology. Seasonality: February-July, October. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; turning stones. References. Lindroth, 1968 (dispersal power); Erwin, 1981 (natural history). Acupalpus (Tachistodes) indistinctus Dejean, 1831 Ecology. Lowlands. Borders of lakes, ponds, slow rivers, and brooks. Open ground; poorly drained, wet, clayish or muddy soil covered with some vegetation. Mostly nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: February-August. Tenerals: July-August. Adults found overwintering (SC). Dispersal power. Macropterous. Frequent flier (in the spring sunshine; at dusk; to artificial lights at night). Slow runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping. References. Lindroth, 1968 (natural history); Kirk, 1969 (biology).

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Acupalpus (Philodes) longulus Dejean, 1829 Ecology. Lowlands. Macrohabitat unknown. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: January-March, May-August. Adults found overwintering in wood debris and under dead leaves. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Turning wood debris; raking the leaf litter. Reference. Kirk, 1969 (biology). Acupalpus (Philodes) lucens (Casey, 1914) Natural History. Unknown. Acupalpus (Acupalpus) meridianus (Linné, 1761) Ecology. Lowlands. Vacant lots and gardens. Open ground; moderately moist, mostly clay soil covered with weeds. Nocturnal; sheltering during the day in grass and weed stacks. Biology. Seasonality: May-June. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Occasional climber (on plants). Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Pitfall trapping; inspecting grass and weed stacks; sweeping plants. References. Lindroth, 1945 and 1968 (natural history); Sharova, 1981 (ecology, biology); Lindroth, 1986 (natural history); Koch, 1989 (ecology); Pollock, 1991 (ecology, dispersal power); Luff, 1993 (biology). Acupalpus (Acupalpus) nanellus Casey, 1914 Ecology. Lowlands. Borders of eutrophic ponds and marshes; swamps, marshy lake shores, Sphagnum-bogs, and wet meadows. Shaded ground; soft, wet soil, rich in organic debris, covered with dense tall vegetation (Typha). Nocturnal; sheltering during the day in leaf litter, under dead grass, and at the base of plant tufts (Juncus). Biology. Seasonality: March-August. Tenerals: April, July. Adults found overwintering on higher and drier ground, in mixed woods and on lake shores; in leaf litter. Dispersal power. Wing-dimorphic: macropterous (mostly) or submacropterous (incapable of flight). Occasional in lake shore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Reference. Lindroth, 1968 (natural history). Acupalpus (Tachistodes) partiarius (Say, 1823) Ecology. Lowlands. Borders of ponds, lakes, rivers, and marshes; pastures, cultivated fields (e.g., corn), and vacant lots. Open ground; moderately moist, muddy soil. Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering during the day in burrows dug in the soil (mostly), in leaf litter, and under pieces of wood. Biology. Seasonality: February-September, November. Adults found overwintering on higher and drier ground, in the vicinity of water, in woods and vacant lots; under pieces of wood and dead

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leaves. Predators: robins. Dispersal power. Macropterous. Frequent flier (in the spring sunshine; at dusk; to artificial lights at night). Regular in lakeshore drift material, which indicates flight ability. Slow runner. Occasional climber (on plants). Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping; sweeping plants; baiting traps with molasses and fermented sugar; turning drift material. References. Lindroth, 1968 (dispersal power); Kirk, 1970 (ecology) and 1975c (dispersal power). Acupalpus (Tachistodes) pauperculus Dejean, 1829 Ecology. Lowlands, mountains, subalpine and alpine zones. Borders of pools and ponds (even temporary ones), lakes, brooks, rivers, cattle water holes, eutrophic marshes, swamps, cranberry bogs, roadsides, gravel pits, meadows, pastures, and vacant lots. Open ground; wet, sandy or sandy-clayish soil covered with dense vegetation (e.g., Glyceria, Carex). Mostly nocturnal; active in the spring sunshine; sheltering during the day in burrows dug in the soil (mostly), under dead leaves, mats of dead vegetation, pieces of wood, and stones, and at the base of plant tufts (Juncus). Gregarious. Biology. Seasonality: January, MarchDecember. Tenerals: April-May; August-December (mostly). Adults found overwintering on higher and drier ground, in fields, on the top of dunes, and at the edge of woods; under dead leaves, depressed grass, stones, and in rotten logs. Dispersal power. Macropterous. Frequent flier (in the spring sunshine; at dusk; to artificial lights at night). Frequent in lakeshore drift material, which indicates flight ability. Slow runner. Occasional climber (on plants). Favored by spring floods along large rivers. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping; sweeping plants; turning drift material. References. Lindroth, 1968 (natural history); Kirk, 1970 and 1975c (dispersal power); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Acupalpus (Acupalpus) pumilus Lindroth, 1968 Ecology. Lowlands. Marshy borders of ponds, lakes, and slow rivers; Sphagnum-bogs and moist meadows. Shaded ground; soft, wet, clayish or muddy soil, rich in organic debris, covered with dense vegetation (e.g., Typha, Juncus, Carex, grass, moss). Nocturnal; sheltering during the day in leaf litter and under heaps of grass. Biology. Seasonality: AprilDecember. Tenerals: May-June, August. Adults found overwintering on higher and drier ground, at the edge of woods, in woods, and on the top of dunes; under dead leaves and stones, and in rotten logs. Dispersal power. Macropterous. Frequent flier (at dusk; to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Favored by spring floods along large rivers. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping; sweeping plants. Reference. Lindroth, 1968 (natural history). Acupalpus (Anthracus) punctulatus Hatch, 1953 Ecology. Pond edges. Moist, muddy soil covered with dense vegetation. Nocturnal; sheltering during the day under plant mats. Biology and Dispersal power. Unknown. Collecting technique. Turning plant mats.

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Acupalpus (Philodes) rectangulus Chaudoir, 1868 Ecology. Lowlands. Edges of small bodies of standing water; eutrophic marshes (e.g., Salix, Typha), forest marshes; edges of slow streams. Somewhat shaded ground; wet clayish soil, rich in dead leaves and dead grass. Nocturnal; sheltering during the day mostly among layers of dead leaves and dead grass, also in clumps of grass. Biology. Seasonality: January-December. Tenerals: July (WV). Adults found overwintering on higher and drier ground, at the edge of woods; in leaf litter and under stones. Dispersal power. Macropterous. Frequent flier (at dusk; to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping. References. Lindroth, 1968 (natural history); Kirk, 1975c (dispersal power). Acupalpus (Anthracus) tener (LeConte, 1857) Natural History. Unknown. Acupalpus (Tachistodes) testaceus Dejean, 1829 Ecology. Lowlands. Borders of ponds, lakes, puddles, rivers, brooks, and marshes; vacant lots. Open ground; wet or moist, muddy or muddy-sandy soil. Nocturnal; active in the spring sunshine; usually sheltering during the day under dead leaves and in the soil. Biology. Seasonality: January-November. Tenerals: July (AL). Adults found overwintering on higher and drier ground, in vacant lots; under dead leaves, in the soil at the base of grass tufts, under heaps of hay, and in wood debris. Predators: starlings. Dispersal power. Macropterous. Frequent flier (in the spring sunshine; at dusk; to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Slow runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping; sifting leaf litter; using an aspirator; digging in the mud. References. Lindroth, 1968 (dispersal power); Kirk, 1969 (ecology) and 1970 (ecology, dispersal power); Erwin, 1981 (natural history). Agonum (Circinalidia) aeruginosum Dejean, 1828 Ecology. Lowlands. Borders of eutrophic open marshes and forest marshes; swamps, borders of slow rivers and brooks; low deciduous forests, Sphagnum-bogs, and wet meadows. Ground often shaded by Salix-bushes; soft, wet, muddy soil covered with dense vegetation (e.g., Carex, Scirpus, Solanum). Nocturnal; sheltering during the day in cracks in the soil and at the base of plants. Biology. Seasonality: March-October. Copulating pairs: June (QU). Gravids: early July (QU). Tenerals: August (NH). Adults found overwintering on higher and drier ground; under logs and pieces of wood. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (on plants, shrubs, and trees). Collecting techniques. Treading the vegetation into the water; pouring water over the ground; light trapping;

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sweeping or beating the vegetation. References. Blatchley, 1910 (ecology, biology); Lindroth, 1966 (natural history); Kirk, 1969 (ecology, dispersal power); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Agonum (Agonum) affine Kirby, 1837 Ecology. Lowlands, mountains, subalpine and alpine zones. Borders of eutrophic marshes and ponds; swamps, mossy bogs, and marshy borders of brooks; drainage channels. Open ground; soft wet, muddy or peaty soil covered with dense vegetation (e.g., Carex, Menyanthes, carpets of brown moss (Scorpidium, Climacium)). Nocturnal; sheltering during the day in moss (mostly) and in leaf litter. Biology. Seasonality: March-October. Tenerals: August (NF). Adult and larval food, in captivity: mealworms. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Collecting techniques. Treading the vegetation into the water; sweeping plants. References. Lindroth, 1955a (as carbo LeConte), 1963a, and 1966 (natural history). Agonum (Agonum) albicrus Dejean, 1828 Ecology. Lowlands. Margins of cypress-swamps; flood-plain forests; low deciduous forests bordering lakes. Shaded ground; wet, clayish or muddy soil, rich in organic debris (e.g., leaf litter), bare or covered with some vegetation. Nocturnal; sheltering during the day in leaf litter, in humus, under logs, and among the roots of rotten tree-stumps. Biology. Seasonality: February-August. Tenerals: May (NC), July (VA). Adults found overwintering on higher and drier ground, in cypress-swamps; in standing rotten tree-stumps. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pitfall trapping; night collecting with a headlamp. References. Blatchley, 1910 (ecology); Lindroth, 1966 (dispersal power); Kirk, 1969 (dispersal power); Allen and Thompson, 1977 (ecology); Krinsky and Oliver, 1988 (ecology); Will and Liebherr, 1997 (ecology). Agonum (Europhilus) anchomenoides Randall, 1838 Ecology. Lowlands and mountains. Margins of eutrophic marshes, ponds, lakes, rivers and brooks; gravel pits near ponds; wet meadows. Open ground; moist, clayish or muddy soil, sometimes mixed with sand, bare or covered with sparse vegetation (e.g., Typha, Equisetum, Juncus). Nocturnal; sheltering during the day under dead leaves, pieces of wood, depressed grass, and stones. Biology. Seasonality: January-December. Copulating pairs: late June (VT). Gravids: early (QU) and late (VT) June. Tenerals: July-October. Adults found overwintering on higher and drier ground, in pastures, hawthorn hedges, edges of woods, and woods; in leaf litter, under stones, and in the soil. Able to survive winter inundations. Predators: frogs and toads. Defense-mechanism: body colorpattern matching the soil. Dispersal power. Macropterous. Frequent flier (at dusk; to

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artificial lights at night; into rotary traps). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Collecting techniques. Treading the soil with the feet; pouring water over the ground; light trapping; using rotary traps; sweeping plants. References. Lindroth, 1963a and 1966 (natural history); Kirk, 1975c (dispersal power). Agonum (Agonum) anthracinum Dejean, 1831 Ecology. Mexico: forests (e.g., Pinus, Quercus); neighboring pastures and grasslands. Nocturnal; sheltering during the day in leaf litter. Biology. Unknown. Dispersal power.Wing-dimorphic: macropterous (probably capable of flight) or brachypterous (incapable of flight). Moderate runner. Collecting technique. Raking the leaf litter. Agonum (Circinalidia) basale LeConte, 1848 Ecology. Forests and muddy habitats. Nocturnal; sheltering during the day. Biology. Seasonality: April-December. Adults found overwintering under pieces of wood and among dead leaves. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Blatchley, 1910 (biology). Agonum (Agonum) belleri (Hatch, 1933) Ecology. Lowlands. Sphagnum-bogs (sometimes situated near lakes and ponds). Open ground; soft, wet soil covered with Sphagnum-moss and Drosera. Mostly diurnal; active in the sunshine on moss. Biology. Seasonality: July-August. Tenerals: early August (WA). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Treading the vegetation into the water. References. Hatch, 1933 (ecology); Lindroth, 1966 (natural history); Kavanaugh, 1992 (natural history). Agonum (Platynomicrus) bicolor (Dejean, 1828) Ecology. River banks. At some distance from water. Shaded ground; moderately dry, clay soil covered with some vegetation (e.g., Equisetum). Nocturnal; sheltering during the day. Biology. Seasonality: June-July, September-October. Tenerals: late June (NT). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Lindroth, 1966 (natural history). Agonum (Agonum) brevicolle Dejean, 1828 Ecology. Borders of eutrophic marshes and bogs. Open ground; soft, wet soil, rich in organic debris, covered with some vegetation (e.g., Carex, Typha). Nocturnal; sheltering during the day under vegetal debris. Biology. Seasonality: May-August. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Treading the vegetation into the water. References. Lindroth, 1966 (natural history); Kavanaugh, 1992 (natural history).

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Agonum (Europhilus) canadense Goulet, 1969 Ecology. Lowlands. Eutrophic marshes, swamps, and Sphagnum-bogs. Open ground; soft, wet soil covered with some vegetation (e.g., Typha, Carex, Sphagnum). Nocturnal; sheltering during the day in leaf litter and moss. Biology. Seasonality: January-December, except February. Adults found overwintering on higher and drier ground, at the edge of woods and in woods; under dead leaves, the loose bark of logs, under stones, and in the soil. Dispersal power. Macropterous. Frequent flier (at dusk; to artificial lights at night). Moderate runner. Occasional climber (on plants and shrubs). Collecting techniques. Treading the vegetation into the water; light trapping; sweeping plants; beating trees. References. Goulet, 1969 (ecology); Lindroth, 1969a (ecology). Agonum collare (Say, 1830) Ecology. Lowlands. Edges of eutrophic marshes. Soft, wet, muddy soil, rich in organic debris, covered with dense vegetation. Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: March, May-July. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Occasional climber. Collecting techniques. Pitfall trapping; baiting traps with malt; using tanglefoot screens; sweeping plants. Agonum (Europhilus) consimile (Gyllenhal, 1810) Ecology. Lowlands and mountains. Borders of lakes, ponds, and pools; Sphagnum-bogs. Close to water. Open ground; soft, wet soil covered with dense vegetation (e.g., tall Carex) and moss carpets (rarely Sphagnum). Often associated with warmer water. Nocturnal; sheltering during the day in moss and vegetal debris. Biology. Seasonality: MayAugust. Tenerals: August (NF). Dispersal power. Macropterous. Occasional flier (to artificial lights at night; into rotary traps). Moderate runner. Collecting techniques. Treading the vegetation carpet into the water; using rotary traps. References. Lindroth, 1945, 1955a, 1955b, 1963a, 1966 and 1986 (natural history). Agonum (Agonum) corvus (LeConte, 1860) Ecology. Lowlands, mountains, subalpine and alpine zones. Margins of sloughs, marshes, and pools. Open ground; wet clay soil covered with some vegetation (e.g., Eleocharis, Carex, Juncus). Nocturnal; sheltering during the day under vegetal debris and stones. Biology. Seasonality: April-July, September, November. Tenerals: July (BC). Adult food, in the field: coccinellids. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Pitfall trapping. References. Lindroth, 1966 (natural history); Kirk, 1975c (dispersal power). Agonum (Olisares) crenistriatum (LeConte, 1863) Ecology. Lowlands and mountains. Meadows, abandoned fields, pastures, cultivated fields (e.g., corn, alfalfa), vacant lots, sandhills, sand pits, gravel pits, and open forests. Open

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ground; dry, sandy (mostly) or gravelly soil covered with sparse vegetation (e.g., grass, weeds). Nocturnal; sheltering during the day under stones (mostly), logs, and dead leaves. Biology. Seasonality: March-December. Gravids: late April (QU). Tenerals: June (QU) and July (OH). Adults found overwintering on hills; under stones and at the base of treestumps. Adult food: corn (in the field); immature stages of ants and beetles, adult carabids, earthworms, and pieces of meat and fruit (in captivity). Predators: toads, crows, and starlings. Dispersal power. Macropterous. Occasional in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pitfall trapping; turning stones. References. Lindroth, 1966 (natural history); Larochelle, 1974a (ecology). Agonum (Circinalidia) crenulatum (LeConte, 1854) Ecology. Lowlands. Macrohabitat unknown. Occurring on wet sand. Biology. Seasonality: April-July. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Occasional climber (on trees). Collecting techniques. Using tanglefoot screens; beating trees. Agonum (Agonum) cupreum Dejean, 1831 Ecology. Lowlands, mountains, subalpine and alpine zones. Grasslands, meadows, cultivated fields (e.g., oat, wheat, barley, bean, fescue), pastures, and abandoned fields. Open ground; dry, more or less sandy soil covered with sparse vegetation (e.g., grass). Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under stones, logs, and dry cow dung. Biology. Seasonality: April-November. Copulating pairs: November (AB). Gravids: late August (AB). Tenerals: July-September. Adults found overwintering on higher and drier ground, at the edge of woods and in woods; under pieces of wood and under the loose bark of tree-stumps. Adult food: flies and coccinellids (in the field); lepidopterous eggs and dipterous pupae (in captivity). Predators: crows. Parasites: hymenopterans. Dispersal power. Wing-polymorphic: macropterous (capable of flight) to brachypterous (incapable of flight). Occasional flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning debris. References. Lindroth, 1966 and 1971a (natural history); Frank, 1971b (biology); Larochelle, 1977g (dispersal power); Levesque and Levesque, 1987 (biology). Agonum (Agonum) cupripenne (Say, 1823) Ecology. Lowlands, mountains, subalpine and alpine zones. Grasslands, pastures, meadows, abandoned fields, cultivated fields (e.g., wheat, rye, corn, oat, barley, alfalfa, cabbage, potato, asparagus, soybean), field margins, ski fields, hills, roadsides, gravel pits, sand pits, orchards (e.g., apple), and open forests. Open ground; rather dry, often gravelly or sandy soil covered with thin vegetation. Mostly diurnal; active in the sunshine; sheltering on cloudy and cool days under stones and dead leaves, at the base of plants, in clumps of grass, under heaps of hay, under pieces of wood, and in the soil. Gregarious (in winter). Biology. Seasonality: January-December. Copulating pairs: May-June

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(QU); coitus lasting 5-7 minutes. Gravids: late May and early June (QU). Tenerals: JulySeptember. Adults found overwintering on higher and drier ground, on hills and hillocks, at the edge of woods, and in woods; under well-embedded stones (mostly), logs, pieces of wood, dead leaves, the loose bark of logs, in the soil, and in rotten tree-stumps. Adult food: flies, coccinellids, orthopterans (in the field); mealworms, lepidopterous caterpillars, dead beetles, earthworms, pieces of meat, and corn kernels (in captivity). Larval food: mealworms (in captivity). Predators: frogs, toads, crows, grouses, and starlings. Dispersal power. Macropterous. Frequent flier (in the sunshine; at dusk; to artificial lights at night). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Strong burrower. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning well-embedded stones; raking the leaf litter; light trapping; sweeping plants; turning drift material. References. Blatchley, 1910 (biology); Lindroth, 1966 (natural history); Kirk, 1975c (ecology, dispersal power); Levesque and Levesque, 1986 (biology); Holliday, 1991 (biology); Will et al. 1995 (dispersal power). Agonum (Stictanchus) cyanope (Bates, 1882) Ecology. Mountains. Lake and brook margins; wet pastures; mesic forest canyons (e.g., Pinus, Quercus) and clearings. Open ground; moist or wet soil. Nocturnal. Biology. Seasonality: March-December. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Pitfall trapping. Reference. Liebherr, 1986b (ecology, dispersal power). Agonum (Agonum) cyclifer (Bates, 1884) Ecology. Mountains. Banks of brooks and arroyos. Open ground; moist, muddy or clayish soil. Nocturnal; sheltering during the day. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pitfall trapping. Agonum (Europhilus) darlingtoni Lindroth, 1954 Ecology. Lowlands. Sphagnum-bogs: along lakes, ponds, pools, and marshes. Open ground; soft, very wet soil covered with rich carpets of moss (Sphagnum). Nocturnal; sheltering during the day among moss. Gregarious. Biology. Seasonality: April-October. Tenerals: September. Dispersal power. Macropterous. Occasional in seashore drift material, which indicates flight ability. Moderate runner. Collecting technique. Treading moss carpets into the water. References. Lindroth, 1955b and 1966 (natural history). Agonum (Agonum) deceptivum (LeConte, 1879) Ecology. Lowlands. Vicinity of brackish water (NS). Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: April-May, August. Dispersal power. Macropterous, probably capable of flight. Moderate runner.

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Agonum (Stictanchus) decorum (Say, 1823) Ecology. Lowlands and mountains. Marshy borders of lakes, ponds, reservoirs, slow rivers and brooks; desert springs. Often associated with small bodies of water. Shaded ground; wet clay soil covered with dense vegetation (e.g., grass) or a layer of decaying vegetation. Mostly crepuscular (at dusk) and nocturnal; sometimes active in the sunshine on bushes; usually sheltering during the day under stones and vegetal debris. Biology. Seasonality: January-November. Gravids: mid-June and early July (QU). Tenerals: July-August. Adults found overwintering on higher and drier ground, at the foot of hills and in woods; under stones, logs, and in leaf litter. Adult food: animal matter (in the field); mealworms (in captivity). Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants and bushes). Collecting techniques. Pitfall trapping; turning debris; light trapping; sweeping plants. References. Blatchley, 1910 (ecology, biology); Lindroth, 1966 (natural history); Kirk, 1969 (dispersal power) and 1975c (ecology, dispersal power); Liebherr, 1983a (ecology) and 1986b (ecology, dispersal power). Agonum (Stictanchus) elongatulum (Dejean, 1828) Ecology. Lowlands. Borders of marshes along lakes and rivers. Open marshy expanses between hammocks. Open ground; wet soil covered with some vegetation. Nocturnal; sheltering during the day under vegetal debris. Biology. Seasonality: March-April, June, December. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting technique. Light trapping. References. Liebherr, 1986b and 1988b (ecology, dispersal power). Agonum (Stereagonum) errans (Say, 1823) Ecology. Lowlands and mountains. Borders of lakes, ponds, marshes, rivers, brooks, and cattle water holes; peat bogs, damp pastures, cultivated fields, gravel pits, open forests, and forest ravines. Open ground; wet soil usually consisting of sand-mixed clay, often saline (Prairies), bare or covered with sparse vegetation. Nocturnal; sheltering during the day under stones, logs, branches, boards, other debris, and in the soil at the base of plants. Biology. Seasonality: January, April-October. Copulating pairs: May (SD) and June (QU). Gravids: late June (QU). Tenerals: July-August. Adults found overwintering on higher and drier ground, in woods. Dispersal power. Macropterous. Occasional flier (at dusk). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pitfall trapping; turning debris; sweeping. References. Blatchley, 1910 (ecology; as subcordatus (LeConte)); Lindroth, 1966 (natural history); Kirk, 1975c (ecology, biology). Agonum (Europhilus) exaratum (Mannerheim, 1853) Ecology. Lowlands. Marshy margins of pools and ponds. Open ground; soft, wet soil covered with some vegetation (e.g., Carex, Eriophorum, Sphagnum). Nocturnal; shelter-

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ing during the day in moss. Biology. Seasonality: May-August. Tenerals: May (YU), August (AK). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Treading moss carpets into the water. Reference. Lindroth, 1966 (natural history). Agonum (Agonum) excavatum Dejean, 1828 Ecology. Lowlands and mountains. Margins of lakes, ponds, marshes, rivers, and brooks; Sphagnum-bogs and wet meadows. Open ground; soft, wet, muddy, clayish or sandy soil, bare or covered with sparse vegetation (e.g., grass). Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: March-October. Copulating pairs: MayJune (QU). Tenerals: July-August. Adults found overwintering on higher and drier ground; in the soil and under well-embedded stones. Predators: frogs. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Occasional climber (on plants). Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet; sweeping plants. References. Blatchley, 1910 (ecology); Lindroth, 1966 (natural history); Erwin, 1981 (natural history). Agonum (Stictanchus) extensicolle (Say, 1823) Ecology. Lowlands and mountains. Margins of rivers, brooks, lakes, and ponds; gravel pits near bodies of water. Caves (occasionally). Close to water. Open or shaded ground; wet soil containing clay or interstitial sand and mud, with abundant stones, bare or covered with sparse vegetation. Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering during the day under stones (mostly), logs, vegetal debris, and the loose bark of logs and tree-stumps. Larval habitat: edge of a brook, in the soil (NY). Biology. Seasonality: February-November. Copulating pairs: May-June (NY, QU). Gravids: early May to mid-July (NY, QU). Tenerals: mid-June to September. Adults found overwintering on higher and drier ground, at the base of hills; under stones, logs, and pieces of wood. Adult able to survive winter inundations. Adult food, in captivity: wireworms, mealworms, dead beetles, and pieces of meat. Larval food, in captivity: mealworms and a scarabaeid larva. Predators: frogs, toads, and quails. Parasites: podapolipid mites and Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult may emit a repugnant Brachinus-like smoke from the pygidial glands. Dispersal power. Wingdimorphic: macropterous (mostly; capable of flight) or brachypterous (incapable of flight). Occasional flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Favored by spring floods along large rivers. Collecting techniques. Turning debris; light trapping. References. Schaupp, 1881b (biology); Lindroth, 1966 (natural history); Erwin, 1981 (natural history); Liebherr, 1986b and 1988b (ecology, dispersal power); Will et al. 1995 (dispersal power). Agonum (Stictanchus) extimum Liebherr, 1986 Ecology. Lowlands. Stream banks. Muddy soil. Nocturnal; sheltering during the day in cracks in the soil. Biology. Seasonality: March, June-October. Dispersal power. Mac-

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ropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting technique. Light trapping. Reference. Liebherr, 1986b (ecology, dispersal power). Agonum (Agonum) fallianum (Leng, 1918) Ecology. Subalpine and alpine zones. Meadows. Open ground; wet soil. Diurnal and nocturnal. Biology. Seasonality: June, September, December. Copulating pairs: late June (CA). Dispersal power. Wing-condition unknown. Moderate runner. Collecting technique. Pitfall trapping. Agonum (Stereagonum) ferreum Haldeman, 1843 Ecology. Lowlands. Low forests bordering ponds, marshes, slow rivers and brooks. Shaded ground; soft, wet, sandy-clayish soil, rich in organic debris (e.g., dead leaves. Nocturnal; sheltering during the day in leaf litter, under logs, stones, and other debris. Biology. Seasonality: January-November. Tenerals: July (MA, WV). Adults found overwintering on higher and drier ground, in woods; among vegetal debris and in tree hollows. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Occasional climber (on trees). Collecting techniques. Turning debris; pitfall trapping; washing the leaf litter in water; beating trees. References. Blatchley, 1910 (ecology); Lindroth, 1966 (natural history); Kirk, 1969 (ecology, biology); Erwin, 1981 (natural history). Agonum (Platynomicrus) ferruginosum (Dejean, 1828) Ecology. Lowlands, mountains, and subalpine zone. Margins of eutrophic marshes, bogs, and small ponds. Open or partially shaded ground; soil, when flooded, with dense emergent vegetation (e.g., mostly Typha; also Carex), clumps of dead floating vegetation or emergent substrate; soil, when not flooded, soft, wet clayish or muddy, with similar vegetation. Mostly nocturnal; usually sheltering during the day in floating Typha-mats. Biology. Seasonality: April-October. Tenerals: April (rarely), late May to September. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (in the sunshine). Moderate runner. Regular climber (on plants). Collecting techniques. Treading the vegetation into the water; sweeping plants. References. Lindroth, 1966 (natural history); Kavanaugh, 1992 (natural history); Landry, 1994 (ecology, biology). Agonum (Agonum) fidele Casey, 1920 Ecology. Lowlands, mountains, and subalpine zone. Swamps; margins of eutrophic marshes, ponds, lakes, rivers, and brooks; wet forests, beaver houses, roadside ditches. Ground shaded by bushes (e.g., Salix, Alnus); wet, muddy, clayish or peaty soil covered with sparse vegetation and thick leaf litter. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in leaf litter, under depressed grass, in moss, and under stones. Biology. Seasonality: January-December. Gravids: early July (QU).

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Tenerals: July-September. Adults found overwintering on higher and drier ground, in woods; in leaf litter. Predators: toads. Dispersal power. Macropterous. Occasional flier (in the sunshine; to artificial lights at night). Frequent in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber. Collecting techniques. Raking the leaf litter; night collecting with a headlamp; turning drift material. References. Lindroth, 1966 (natural history); Larochelle, 1974a (ecology); Erwin, 1981 (natural history). Agonum (Agonum) fossigerum Dejean, 1828 Ecology. Stream margins. Greenhouses (occasionally). Moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: April-August. Dispersal power. Wing-dimorphic: macropterous (mostly; probably capable of flight) or brachypterous (incapable of flight). Moderate runner. Collecting technique. Pitfall trapping. Agonum (Europhilus) galvestonicum (Casey, 1920) Ecology. Lowlands. Shores of lakes and ponds. Open ground; inundated or wet, soft, muddy soil covered with dense tall vegetation (e.g., Typha). Mostly nocturnal; usually sheltering during the day in sheaths of standing Typha-plants (mostly) and in clumps of broomsedges. Biology. Seasonality: April-August. Tenerals: August (ON). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Frequent climber (on plants). Collecting techniques. Treading the Typha-vegetation into the water; sweeping the vegetation. References. Lindroth, 1966 (natural history); Kirk, 1969 (ecology); Davidson, 1988a (ecology). Agonum (Europhilus) gratiosum (Mannerheim, 1853) Ecology. Lowlands and mountains. Borders of eutrophic marshes, ponds, lakes; swamps, flood-plain forests; marshy borders of rivers and brooks; roadside ditches and drainage channels; Sphagnum-bogs, gravel pits, wet meadows, low forests, beaver houses and dams. Open ground; mostly firm, often peaty, moderately moist soil covered with some vegetation (e.g., Carex). Nocturnal; sheltering during the day under dead leaves, heaps of hay, depressed grass, stones, or in moss. Gregarious (in winter). Biology. Seasonality: January-December. Copulating pairs: July (QU). Gravids: early July (QU). Tenerals: April, mid-June, August-November. Adults found overwintering on higher and drier ground, on the top of sand dunes, in hawthorn hedges, at the edge of woods, and in woodland swamps; under stones, dead leaves, logs, the loose bark of tree-stumps and fallen trees, in rotten logs and tree-stumps, and in the soil. Adult food, in captivity: mealworms and meat. Predators: frogs and toads. Also eaten by insectivorous plants (Sarracenia). Phoretic mites found on adult body. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (on plants and trees). Favored by spring floods along large rivers. Collecting techniques. Raking the leaf litter; treading the vegetation into

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the water; sweeping or beating the vegetation. References. Lindroth, 1955a, 1955b and 1963a (natural history); Kirk, 1975c (dispersal power); Holliday, 1991 (biology). Agonum (Agonum) harrisii LeConte, 1848 Ecology. Lowlands and mountains. Margins of eutrophic marshes, ponds and lakes; swamps; marshy borders of slow rivers and brooks; flood-plain forests, drainage channels, beaver houses and dams, and Sphagnum-bogs. Open or half-shaded ground; soft wet soil, rich in organic debris, covered with dense vegetation (e.g., Carex, Typha). Nocturnal; sheltering during the day under dead leaves and stones. Gregarious (in winter). Biology. Seasonality: March-December. Copulating pairs: early June (QU). Gravids: early May (QU). Tenerals: July-August; mostly after mid-August. Adults found overwintering on higher and drier ground, on the top of sand dunes, at the edge of woods, and in woods; in the soil (as deep as 10 cm), under stones, dead leaves, and the loose bark of fallen trees, and in rotten logs. Predators: frogs, toads. Phoretic mites found on adult body. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Treading the vegetation into the water; light trapping; using malaise traps. Reference. Lindroth, 1966 (natural history). Agonum (Agonum) imitans (Notman, 1919) Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Liebherr, 1991b (dispersal power). Agonum (Circinalidia) limbatum Motschulsky, 1845 Ecology. Lowlands, mountains, subalpine and alpine zones. Edges of ponds, reservoirs, pools, and irrigation ditches; alpine meadows. Open ground; moist soil. Nocturnal; sheltering during the day under grass mats and the loose bark of trees. Biology. Seasonality: May-July. Adult food, in the field: seeds, vegetal tissue, aphids, and lepidopterous caterpillars. Predators: Grosbeaks and swallows. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pitfall trapping. Agonum (Europhilus) lutulentum (LeConte, 1854) Ecology. Lowlands and mountains. Margins of eutrophic marshes (mostly) and swamps; marshy borders of lakes, ponds, pools, and slow rivers; roadside ditches, drainage channels, cranberry bogs, wet meadows and pastures, flood-plain forests, low forests, and orchards (e.g., apple). Often associated with quite small bodies of water. Open ground; soil flooded, with dense emergent vegetation (e.g., Typha, tall Carex). Nocturnal; sheltering during the day in floating heaps of dead plants and leaves. Gregarious (in winter). Biology. Seasonality: January-December. Tenerals: May-September; mostly August-September. Adults found overwintering on higher and drier ground, on the top of hills and sand dunes, in hawthorn hedges, at the edge of woods, and in woods; among dead

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André Larochelle & Marie-Claude Larivière

leaves, under stones, logs, pieces of wood, depressed grass, in the soil (as deep as 20 cm), in rotten logs, and under the loose bark of fallen trees. Adult food, in captivity: grass seeds and a piece of apple. Phoretic mites found on adult body. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; into rotary traps). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Moderate climber (on plants and trees). Collecting techniques. Treading the Typha-vegetation into the water; sweeping or beating the vegetation; light trapping; using rotary traps. References. Blatchley, 1910 (biology, dispersal power); Lindroth, 1955b (ecology) and 1966 (natural history); Kirk, 1975c (dispersal power); Landry, 1994 (ecology, biology). Agonum (Agonum) melanarium Dejean, 1828 Ecology. Lowlands, mountains, subalpine and alpine zones. Margins of eutrophic marshes; swamps; marshy borders of ponds, lakes, slow rivers, brooks, and irrigation channels; flood-plain forests; low deciduous forests; alluvial places, beaver houses and dams, and wet pastures. Occurring near water. Open ground or shaded by trees, shrubs and bushes; soft, wet soil, usually consisting of clay mixed with organic debris, covered with some vegetation (e.g., Typha, Carex). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under dead leaves and stones (mostly), and under pieces of wood. Gregarious (in winter). Biology. Seasonality: January-December. Copulating pairs: June (QU). Gravids: early May to late July (NY, QU). Tenerals: MayNovember; mostly after mid-July. Adults found overwintering on higher and drier ground, on the top of sand dunes, at the foot of hills, in hawthorn hedges, at the edge of woods, and in woods; in the soil (as deep as 30 cm), under stones and vegetal debris, in rotten logs and tree-stumps, and under the loose bark of tree-stumps. Able to survive winter inundations. Adult food, in captivity: earthworms, chrysomelid larvae, and meat. Predators: frogs, toads, and crows. Parasites: Laboulbeniales (Fungi). Phoretic mites found on adult body. Dispersal power. Macropterous. Frequent flier (in the sunshine; at dusk; to artificial lights at night). Moderate runner. Occasional climber (on plants). Favored by spring floods along large rivers. Collecting techniques. Pitfall trapping; turning debris; light trapping; sweeping plants. References. Blatchley, 1910 (biology); Lindroth, 1966 (natural history); Kirk, 1975c (ecology, dispersal power); Epstein and Kulman, 1990 (ecology, biology); Will et al. 1995 (dispersal power). Agonum (Agonum) metallescens (LeConte, 1854) Ecology. Lowlands, mountains, and subalpine zone. Borders of eutrophic marshes and ponds; swamps and bogs; marshy borders of slow rivers and brooks; wet pastures. Open ground; soft wet soil, usually consisting of clay mixed with organic debris, covered with some vegetation (e.g., Carex, Equisetum, depressed grass). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under dead leaves and in moss. Biology. Seasonality: April-September, December. Gravids: early June to late July (QU). Tenerals: July-September. Predators: frogs. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants and shrubs).

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Collecting techniques. Treading the vegetation into the water; light trapping; sweeping plants; beating trees. References. Lindroth, 1955a, 1963a, and 1966 (natural history); Kavanaugh, 1992 (natural history). Agonum (Agonum) moerens Dejean, 1828 Ecology. Lowlands. Flood-plain forests, swamp-forests, marshy borders of pools in open deciduous forests; dystrophic marshes. Open or shaded ground; soft, wet soil, rich in organic debris, covered with some vegetation (e.g., Typha, Solanum, Iris, Alisma, moss). Nocturnal; sheltering during the day under pieces of wood and in leaf litter, often at the base of rotten tree-stumps. Biology. Seasonality: May-October. Tenerals: July (LA, VA, WI). Dispersal power. Macropterous. Occasional flier (to artificial lights at night; into malaise traps). Moderate runner. Occasional climber (on trees). Collecting techniques. Pitfall trapping; turning debris; beating trees; using malaise traps; pyrethrum spraying. References. Lindroth, 1966 (natural history); Krinsky and Oliver, 1988 (ecology, dispersal power). Agonum (Agonum) muelleri (Herbst, 1784) Ecology. Lowlands, mountains, subalpine and alpine zones. Meadows, abandoned fields, pastures, cultivated fields (e.g., corn, cabbage, bean, raspberry), field margins, ski fields, roadsides, gravel pits, sand pits, gardens, lawns, parks, warfs, forest edges; upper zone of lake shores, river and brook banks. Open ground; moderately dry, often clayish soil covered with some vegetation (e.g., grass, weeds). Mostly diurnal; quite active in the sunshine; plastic in its diel rhythm (Europe); sheltering on cloudy or cool days under stones and dead leaves. Gregarious (in winter). Biology. Seasonality: February-November. Copulating pairs: June-July (ME, QU). Gravids (QU): late June to early July (mostly); also early August (once). Tenerals: July-August. Adults found overwintering on higher and drier ground, in gardens, at the edge of fields and woods, in woods; under stones, vegetal debris, and under the loose bark of tree-stumps and fallen trees. Adult food: wireworms (in the field); chrysomelid larvae (in captivity). Predators: toads (regularly). Dispersal power. Macropterous. Frequent flier (in the sunshine; to artificial lights at night). Frequent in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones;turning drift material; sweeping plants. References. Lindroth, 1945, 1955a, 1963a, and 1966 (natural history); Greenslade, 1965 (ecology, biology); Tyler and Ellis, 1979 (biology); Sharova, 1981 (ecology, biology); Lindroth, 1986 (natural history); Levesque and Levesque, 1987 (biology); Koch, 1989 (ecology); Levesque and Levesque, 1994 (biology). Agonum (Agonum) muiri Liebherr, 1984 Ecology. Mountains. Shaded areas above a lake, with mossy soil watered by snowfield runoff. Nocturnal; sheltering during the day. Biology. Seasonality: June-September. Dispersal power. Wing-polymorphic: macropterous (probably capable of flight) to brac-

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André Larochelle & Marie-Claude Larivière

hypterous (incapable of flight). Moderate runner. Reference. Liebherr, 1984b (ecology, dispersal power). Agonum mutatum (Gemminger and Harold, 1868) Ecology. Lowlands, mountains, subalpine and alpine zones. Peat-bogs, swamps, and marshes: often along pools, ponds, and lakes. Close to water. Open ground; soft, often floating, very wet soil covered mostly with moss carpets (Sphagnum), also Oxycoccus, Myrica, Carex). Nocturnal; sheltering during the day in the soaking wet moss (Sphagnum). Biology. Seasonality: March-November. Copulating pairs: July (QU). Gravids: early July (QU). Tenerals: July-August. Phoretic mites found on adult body. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Collecting techniques. Treading the Sphagnum-carpets into the water; using malaise traps; using tanglefoot screens; sweeping plants. References. Lindroth, 1955a, 1963a, and 1966 (natural history); Larochelle, 1974a (ecology). Agonum (Platynomicrus) nigriceps LeConte, 1848 Ecology. Lowlands and mountains. Margins of eutrophic marshes, ponds, and lakes; swamps and lagoons; marshy borders of slow rivers and brooks. Close to open water. Mostly associated with sedge tussocks. Open ground; soil flooded, with dense emergent vegetation (mostly e.g., Carex rostrata Stokes; also Typha). Nocturnal; active during the night on plants. Biology. Seasonality: April-November. Gravids: April to mid-August, mostly before mid-July (AB, QU). Eggs are laid on Typha- and Carex-plants. Tenerals: June-September. Adult and larval food, in captivity: mealworms and shrimps. Defense-mechanism: body color-pattern and shape matching background foliage and coocccurring spiders (Pachygnatha tristriata Koch; Tibellus oblongus (Walckenauer)). Dispersal power. Wing-dimorphic: macropterous (probably capable of flight) or brachypterous (incapable of flight). Occasional flier (in the sunshine). Moderate runner. Frequent climber (on plants). Collecting techniques. Treading the Typha- or Carex-vegetation into the water; sweeping plants. References. Lindroth, 1955a, 1963a, and 1966 (natural history); Larochelle, 1976i (ecology); Landry, 1994 (ecology, biology). Comments. The beetle was considered a a rarity until the discovery of its aquatic habitat and the treading technique necessary to collect it. Agonum (Circinalidia) nutans (Say, 1823) Ecology. Lowlands. Alvar grasslands (mostly near small bodies of standing water), pastures, tall-grass prairies, meadows, abandoned fields, edges of forests, and open forests. Open ground; dry or moderately moist, clayish or limestone soil covered with dense vegetation (e.g., mostly grass, also other herbaceous plants). Mostly nocturnal; usually sheltering during the day under stones and debris; often found on trees. Biology. Seasonality: January, March-December. Tenerals: early July to September. Adults

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found overwintering on higher and drier ground, in pastures and at the edge of woods; under stones, fallen grass, and pieces of wood. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Regular in lakeshore drift material, which indicates flight ability. Moderate runner. Frequent climber (on trees). Collecting techniques. Pitfall trapping; turning stones; beating trees; turning drift material. References. Blatchley, 1910 (biology); Leng, 1915 (ecology); Lindroth, 1966 (natural history); Kirk, 1975c (dispersal power); Bouchard et al. 1998 (ecology, biology); Tinerella and Rider, 2001 (ecology). Agonum (Agonum) octopunctatum (Fabricius, 1798) Ecology. Lowlands and mountains. Borders of rivers, ponds, brooks, lakes, pools, and small bodies of water in sand pits; flood-plain forests; moist or wet meadows, cultivated fields (e.g., corn, soybean, alfalfa, potato, tobacco). Usually close to water. Open ground; moist, sandy or sandy-clayish soil, bare or covered with sparse vegetation. Mostly nocturnal; usually sheltering during the day in leaf litter, under plants and stones, and in cracks in the soil. Biology. Seasonality: January-November. Copulating pairs: June-July (QU). Adults found overwintering on higher and drier ground, on hills, at the edge of woods, and in woods; under stones and vegetal debris. Adult food: lepidopterous caterpillars (in the field); aphids and mealworms (in captivity). Larval food, in captivity: mealworms. Predators: frogs and toads. Dispersal power. Macropterous. Frequent flier (in the rain; at dusk; to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on trees). Moderately favored by human activities. Collecting techniques. Pitfall trapping; light trapping; using tanglefoot screens; beating trees. References. Blatchley, 1910 (ecology, biology); Lindroth, 1966 (natural history); Kirk, 1969 (natural history) and 1970 (dispersal power); Erwin, 1981 (natural history). Agonum (Agonum) pacificum Casey, 1920 Ecology. Lowlands and mountains. Brook margins and arroyo banks. Open ground; moist soil covered with some vegetation. Nocturnal; sheltering during the day. Biology. Seasonality: April-June (Mexico). Dispersal power. Wing-condition unknown. Moderate runner. Agonum (Olisares) pallipes (Fabricius, 1787) Ecology. Lowlands. Pastures, cultivated fields (e.g., tobacco), old fields, vacant lots, orchards (e.g., peach, plum), and open forests. Open ground; dry, sandy (mostly) or gravelly soil covered with sparse vegetation (e.g., grass). Mostly nocturnal; usually sheltering during the day under debris and on trees. Biology. Seasonality: January-December. Tenerals: March (AL). Adults found overwintering in pastures and vacant lots; under pieces of wood, dead leaves, heaps of hay, and in the soil at the base of plants. Adult food, in the field: weed seeds, plant material, aphids, and lepidopterous caterpillars. Predators: robins. Dispersal power. Macropterous, probably capable of flight. Moder-

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André Larochelle & Marie-Claude Larivière

ate runner. Occasional climber (on trees). Collecting techniques. Pitfall trapping; beating trees. Agonum (Europhilus) palustre Goulet, 1969 Ecology. Lowlands and mountains. Deciduous woodland swamps, flood-plain forests, eutrophic marshes (Salix), marshy borders of slow rivers, brooks, and drainage channels. Ground shaded by trees and shrubs; soft, wet, clayish or muddy soil, rich in organic debris, covered with moderate or dense vegetation (e.g., Typha, Solanum, grass). Nocturnal; sheltering during the day under dead leaves, depressed grass, and stones. Biology. Seasonality: February-December. Tenerals: March; July-September (mostly). Adults found overwintering on higher and drier ground, at the edge of irrigation canals, on the top of sand dunes, at the edge of woods, and in woods; under dead leaves, stones, and the loose bark of fallen trees, in rotten logs and fallen rotten branches. Predators: toads. Phoretic mites found on adult body. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Occasional in seashore drift material, which indicates flight ability. Occasional climber (on plants). Favored by spring floods along large rivers. Collecting techniques. Treading the vegetation into the water; light trapping; sweeping plants. References. Goulet, 1969 (ecology, biology); Lindroth, 1969a (ecology). Agonum (Stictanchus) parextimum Liebherr, 1986 Ecology. Lowlands. Stream margins. Moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: February, July-October. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Reference. Liebherr, 1986b (ecology, dispersal power). Agonum (Platynomicrus) piceolum (LeConte, 1879) Ecology. Lowlands, mountains, and subalpine zone. Banks of river and brooks; lake shores. At a considerable distance from water. Ground somewhat shaded by trees; moderately moist, clayish, sandy or gravelly soil, bare or covered with sparse vegetation (e.g., grass) Nocturnal; sheltering during the day under dead leaves and stones. Biology. Seasonality: May-October, December. Gravids: late June (QU). Tenerals: June-July (NH). Predators: toads. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Lindroth, 1963a (as bicolor (Dejean)) and 1966 (natural history). Agonum (Europhilus) picicornoides Lindroth, 1966 Ecology. Lowlands, mountains, subalpine and alpine zones. Lake shores, beaver ponds, eutrophic marshes, and edges of slow brooks. Ground shaded by bushes (e.g., Salix, Alnus); wet, clayish or clayish-sandy soil covered with sparse vegetation. Nocturnal; sheltering during the day under dead leaves and stones. Biology. Seasonality: May-

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September. Gravids: early June (QU). Tenerals: July-September. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Occasional climber (on plants). Collecting technique. Raking the leaf litter. References. Lindroth, 1966 (natural history); Larochelle, 1974a (ecology). Agonum (Agonum) placidum (Say, 1823) Ecology. Lowlands, mountains, subalpine and alpine zones. Grasslands, relict prairies, rangelands, cultivated fields and their borders (e.g., corn, barley, oat, soybean, alfalfa, tobacco, wheat, asparagus, rye, cabbage, flax, potato, carrot, raspberry), abandoned fields, vacant lots, pastures, meadows, golf courses, gardens, roadsides, fencerows, sand pits, gravel pits, orchards (e.g., apple), forest clearings, and open forests. Open ground; very dry, usually sandy soil covered with weeds. Mostly nocturnal; usually sheltering during the day under stones, clods of earth, dead leaves and plants, heaps of hay, and at the base of plants. Gregarious (in winter). Biology. Seasonality: January-December. Copulating pairs: mid-June (QU). Gravids: June-August (AB, ON, QU). Tenerals: late June to September. Adults found overwintering on higher and drier ground, at the edge of fields, at the foot of hills, on the top of sand dunes, in hawthorn hedges, at the edge of woods, in woods, and woodland swamps; under stones, pieces of wood, logs, and dead leaves, in the soil (as deep as 20 cm), under the loose bark of tree-stumps, and also in rotten logs and tree-stumps. Adult food: dipterous larvae, lepidopterous caterpillars, and vegetal matter (in the field); lepidopterous eggs, slugs, and meat (in captivity). Predators: toads (frequently), frogs, woodpeckers, and starlings. Parasites: nematodes. Phoretic mites found on adult body. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Regular climber (on plants and trees). Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; sweeping or beating the vegetation; light trapping; turning drift material. References. Blatchley, 1910 (ecology, biology); Lindroth, 1955a, 1963a, and 1966 (natural history); Rivard, 1964c (biology); Frank, 1971b (biology); Kirk, 1975c (ecology, dispersal power); Erwin, 1981 (natural history); Levesque and Levesque, 1994 (biology); Will et al. 1995 (dispersal power). Agonum (Agonum) propinquum (Gemminger and Harold, 1868) Ecology. Lowlands and mountains. Borders of pools (often rather small ones), eutrophic marshes, ponds, and lakes; swamps, marshy edges of slow rivers and drainage channels; flood-plain forests and wet pastures. Open ground; rather firm, wet soil, mixed with organic debris, covered with moderately dense vegetation (e.g., Carex, Eleocharis, moss carpet (e.g., Drepanocladus)). Nocturnal; sheltering during the day under dead leaves and pieces of wood. Gregarious (in winter). Biology. Seasonality: January-December. Copulating pairs: May-July. Gravids: early to late July (QU). Tenerals: JulyNovember. Adults found overwintering on higher and drier ground, in hawthorn hedges, at the edge of marshes and woods, in woodland swamps; under stones, dead leaves, the loose bark of fallen trees, in rotten logs and tree-stumps, and in the soil (as deep as

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André Larochelle & Marie-Claude Larivière

15 cm). Adult food, in captivity: meat. Predators: toads. Phoretic mites found on adult body. Dispersal power. Macropterous. Occasional flier (to artificial lights in captivity). Moderate runner. Collecting technique. Raking the leaf litter. References. Lindroth, 1955a, 1963a (as fraterculum (LeConte)), and 1966 (natural history). Agonum (Olisares) punctiforme (Say, 1823) Ecology. Lowlands and mountains. Pastures, meadows, old fields, vacant lots, lawns, cultivated fields (e.g., corn, soybean, alfalfa, tobacco, cotton), roadsides, vicinity of small streams in forests, orchards (e.g., apple), bushy areas, forest edges, open forests (e.g., deciduous), pine stands, and flood-plain forests. Open (mostly) or shaded ground; moist soil covered with sparse vegetation or dead leaves. Mostly nocturnal; usually sheltering during the day under stones, dead leaves and plants, logs, pieces of wood, on bushes and trees, and in clumps of broomsedges. Larval habitat: in the soil. Biology. Seasonality: January-December. Tenerals: April, June-July; mostly July. Adults found overwintering in pastures and vacant lots; under pieces of wood, dead leaves, in the soil at the base of plants, and in clumps of broomsedges. Adult food: chrysomelids and other insects (in the field); lepidopterous caterpillars (in captivity). Larval food, in captivity: mealworms. Predators: toads, bluebirds, crows, woodpeckers, and spiders. Defensemechanism: when alarmed, the adult may emit a Brachinus-like smoke from the pygidial glands. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on bushes and trees). Strongly favored by human activities. Collecting techniques. Pitfall trapping; light trapping; beating bushes and trees. References. Leng, 1915 (ecology, dispersal power); Lindroth, 1966 (natural history); Kirk, 1969 (biology, dispersal power) and 1970 (ecology); Thompson and Allen, 1974 (ecology, biology); Allen and Thompson, 1977 (ecology); Riddick and Mills, 1995 (biology); Anderson et al. 1995 (ecology, biology); Will et al. 1995 (dispersal power); Chen and Willson, 1996 (biology). Agonum (Deratanchus) quadrimaculatum (G.H. Horn, 1885) Ecology. Lowlands. Flood-plain forests and low forests: along slopes, moist places, vernal pools, ponds, and rivers. Swamps and marshes. Ground shaded by trees; moist soil. Nocturnal; sheltering during the day under pieces of wood. Biology. Seasonality: January-July. Adults found overwintering under the loose bark of fallen trees. Dispersal power. Wing-dimorphic: macropterous (probably capable of flight) or brachypterous (incapable of flight). Moderate runner. Collecting techniques. Turning pieces of wood; pitfall trapping. References. Blatchley, 1906 (ecology) and 1910 (natural history); Lindroth, 1966 (dispersal power). Agonum (Agonum) quinquepunctatum Motschulsky, 1844 Ecology. Lowlands and mountains. Peat bogs (sometimes along rivers). Open ground; moist, peaty soil. Nocturnal; sheltering during the day. Biology. Seasonality: June-Au-

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gust. Tenerals: June (YU), August (AB). Dispersal power. Wing-polymorphic: macropterous (probably capable of flight) to brachypterous (incapable of flight). Moderate runner. Collecting technique. Treading the soil with the feet. Reference. Lindroth, 1966 (natural history). Agonum (Europhilus) retractum LeConte, 1848 Ecology. Lowlands and mountains. Deciduous (mostly), mixed and coniferous forests, including roads and edges. Vicinity of human habitations (urban and rural sites). Not associated with water bodies. Ground shaded by bushes and deciduous trees; moderately moist soil, with thick leaf litter. Nocturnal; sheltering during the day in leaf litter and humus, under logs, under the loose bark of fallen trees, and in rotten logs. Biology. Seasonality: February-December. Copulating pairs: June-July (QU). Gravids: mid to late July (QU). Tenerals: April-May, August-November. Adults found overwintering on higher and drier ground, in hawthorn hedges, at the edge of woods, in woods and woodland swamps; under dead leaves, stones, pieces of wood, clumps of grass, and in rotten tree-stumps. Adult food: soft-bodied arthropods (in the field); mealworms, shrimps, and meat (in captivity). Larval food: springtails, tipulid larvae, lepidopterous caterpillars, and spiders (in the field); mealworms (in captivity). Predators: toads and shrews. Defense-mechanism: when seized, the adult may emit a strong repugnant smell that clings to the fingers. Dispersal power. Wing-polymorphic: brachypterous (mostly; incapable of flight) to macropterous (probably capable of flight). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on shrubs, in malaise traps). Collecting techniques. Pitfall trapping; raking the leaf litter; using malaise traps. References. Darlington, 1936 (dispersal power); Lindroth, 1955a, 1963a, and 1966 (natural history); Freitag and Poulter, 1970 (biology); Carter, 1976 (dispersal power); Bousquet and Pilon, 1977 (biology); Levesque et al. 1979 (biology); Reeves et al. 1983 (biology); Levesque and Levesque, 1986 (biology); Niemelä and Spence, 1991 (ecology). Agonum (Olisares) rigidulum (Casey, 1920) Ecology. Lowlands. Moist places and vicinity of river banks in flood-plain forests. Moist soil composed of sand mixed with clay, covered with depressed vegetation. Nocturnal; sheltering during the day under depressed plants, stones, and pieces of wood. Biology. Seasonality: April-June, October. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting technique. Pitfall trapping. Reference. Erwin, 1981 (natural history). Agonum (Olisares) rufipes Dejean, 1828 Ecology. Lowlands. Vicinity of ponds, lakes, brooks, rivers, and Sphagnum-bogs; moist places in forests; open forests. Moist soil composed of sand-mixed clay or of sand and gravel. Mostly nocturnal; usually sheltering during the day. Biology. Seasonality: January-October, December. Adults found overwintering under pieces of wood. Predators: frogs. Dispersal power. Macropterous. Occasional flier (during the day). Occa-

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sional in seashore drift material, which indicates flight ability. Moderate runner. Collecting technique. Pitfall trapping. References. Lindroth, 1966 (natural history); Kirk, 1975c (ecology). Agonum (Europhilus) simile Kirby, 1837 Ecology. Margins of cold ponds and pools. Open ground; wet soil covered with dense vegetation (e.g., Carex, Comarum, Menyanthes, Myrica) or a continuous moss carpet (rarely Sphagnum). Biology. Seasonality: April, June-September. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Treading the vegetation into the water. Reference. Lindroth, 1966 (natural history). Agonum (Europhilus) sordens Kirby, 1837 Ecology. Lowlands, mountains, and subalpine zone. Borders of eutrophic marshes; swamps, marshy borders of slow rivers and small brooks; flood-plain forests. Frequently at the border of little rills (in spring). Not restricted to the vicinity of open water. Ground open (mostly) or shaded by bushes (e.g., Salix, Alnus); rather firm, moist, often sandy soil with organic debris on surface, covered with moderately dense vegetation (e.g., Carex, grass). Mostly nocturnal; sheltering during the day under dead leaves and depressed grass. Biology. Seasonality: January-December. Copulating pairs: May (ON), July (QU). Gravids: late June (QU). Tenerals: August (QU, VT). Adults found overwintering on higher and drier ground, on the top of sand dunes, in pastures, at the edge of woods, in woods and woodland swamps; in leaf litter, under stones, depressed grass, the loose bark of fallen trees and tree-stumps, fallen pieces of bark, in rotten logs and tree-stumps, and in the soil (as deep as 10 cm). Predators: toads. Phoretic mites found on adult body. Dispersal power. Macropterous. Frequent flier (in the sunshine; at dusk; to artificial lights at night; into rotary traps). Moderate runner. Frequent climber (on plants and shrubs). Favored by spring floods along large rivers. Collecting techniques. Treading the vegetation into the water; sweeping or beating the vegetation; pitfall trapping; using rotary traps. References. Lindroth, 1955a, 1955b, 1963a, and 1966 (natural history); Erwin, 1981 (natural history). Agonum (Circinalidia) striatopunctatum Dejean, 1828 Ecology. Lowlands. Flood-plain forests, low deciduous forests, swamp-forests, bogs, borders of ponds, lakes, and rivers; meadows, pastures, and cultivated fields (e.g., corn, peanut). Open ground; wet or moist, clayish or muddy soil covered with some vegetation (e.g., grass) or dead leaves. Mostly nocturnal; usually sheltering during the day under dead leaves, and pieces of wood; sometimes active on cloudy days on plants. Biology. Seasonality: March-December. Adults found overwintering at the edge of fields and in woods; in leaf litter and under logs. Adult and larval food, in captivity: mealworms. Predators: ducks. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Collecting techniques. Pitfall

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trapping; using tanglefoot screens; sweeping plants. References. Leng, 1915 (ecology, dispersal power); Lindroth, 1966 (natural history); Kirk, 1969 (ecology, dispersal power); Allen and Thompson, 1977 (ecology); Erwin, 1981 (natural history). Agonum (Agonum) subsericeum (LeConte, 1863), see Agonum (Agonum) suturale Say, 1830. Reference. Liebherr, 1994 (taxonomy). Agonum (Stereagonum) sulcipenne (G.H. Horn, 1881) Ecology. Lowlands and mountains. Flood-plain forests and swamp-forests (e.g., cypress, gum). Shaded ground; wet, muddy soil. Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: February-July. Tenerals: late July (IL). Adults found overwintering under rotten logs. Dispersal power. Submacropterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. Reference. Löding, 1945 (ecology). Agonum (Europhilus) superioris Lindroth, 1966 Ecology. Lowlands and mountains. Borders of eutrophic marshes, ponds, lakes, swamps, fens, and bogs. Associated with open water. Open ground; soft, wet, muddy soil covered with dense vegetation (e.g., Typha, Carex, Equisetum, Sphagnum-carpets). Nocturnal; sheltering during the day under depressed grass and in moss. Biology. Seasonality: May-September. Gravids: mid-June to mid-July (QU). Tenerals: mid-June (QU). Dispersal power. Submacropterous, incapable of flight. Moderate runner. Collecting technique. Treading the Sphagnum-carpets into the water. References. Lindroth, 1966 (natural history); Larochelle, 1976d (ecology). Agonum suturale Say, 1830 (Agonum subsericeum (LeConte, 1863)) Ecology. Lowlands and mountains. Borders of rivers, ponds, lakes and reservoirs; edges of eutrophic marshes; wet pastures and cultivated fields (e.g., corn). Usually close to water. Open ground; wet clay soil covered with sparse vegetation. Nocturnal; sheltering during the day under vegetal debris, stones, and dry cow dung. Biology. Seasonality: May-July. Copulating pairs: late June (MT). Tenerals: late June (OR). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Treading the soil with the feet; pitfall trapping. References. Lindroth, 1966 (natural history); Liebherr, 1994 (taxonomy). Agonum (Agonum) tenue (LeConte, 1854) Ecology. Lowlands, mountains, subalpine and alpine zones. Margins of eutrophic marshes and ponds; marshy borders of slow rivers; flood-plain forests, low deciduous forests,

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Sphagnum-bogs, beaver houses and dams. Open ground; soft, wet, muddy or clayish soil mixed with organic debris, covered with dense vegetation (e.g., Typha, Carex). Nocturnal; sheltering during the day under dead leaves, pieces of wood, stones, and in moss. Gregarious (in winter). Biology. Seasonality: January-November. Gravids: late May (NY, QU). Tenerals: July-August. Adults found overwintering on higher and drier ground, at the edge of woods and in woods; in leaf litter, under stones, and in wood debris. Adult and larval food, in captivity: mealworms. Predators: frogs and toads. Phoretic mites found on adult body. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on trees). Collecting techniques. Treading the vegetation into the water; light trapping; beating trees; turning drift material. References. Lindroth, 1966 (natural history); Kirk, 1969 (biology, dispersal power); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Agonum (Stictanchus) texanum (LeConte, 1878) Ecology. Lowlands. Mostly margins and beds of brooks; also pastures in scrub habitats. Moderately moist or dry soil. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: April, June, August. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting technique. Light trapping. Reference. Liebherr, 1986b (ecology, dispersal power). Agonum (Europhilus) thoreyi Dejean, 1828 Ecology. Lowlands and mountains. Margins of eutrophic marshes, ponds, and lakes; swamps and drainage channels. Often associated with small bodies of water. Open ground; soil, when flooded, covered with dense emergent vegetation (e.g., Typha, Carex, Equisetum, moss (not Sphagnum)), and with emergent substrate and clumps of dead vegetation; soil, when not flooded, wet clayish or muddy, rich in organic debris, with similar vegetation. Nocturnal; sheltering during the day among dead plants (e.g., Typha) and under dead leaves. Gregarious (in winter). Larval habitat: in the hollow stems of the marsh vegetation. Biology. Seasonality: January, March-December. Gravids: May-August (AB, ON, QU). Tenerals: AprilNovember. Adults found overwintering on higher and drier ground, on the top of sand dunes, in hawthorn hedges, at the edge of woods, and in woods; in leaf litter, under stones, the loose bark of fallen trees, and in rotten logs. Adults hibernating in leaf sheaths and hollow stems of plants (Europe). Adult food, in captivity: shrimps and grass seeds. Predators: nematodes. Phoretic mites found on adult body. Defense-mechanism: when alarmed, the adult swims readily on the water surface. Dispersal power. Macropterous. Frequent flier (at dusk; to artificial lights at night; into rotary traps). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (on plants). Good swimmer. Collecting techniques. Treading the Typha-vegetation into the water; sweeping plants; light trapping; using rotary traps. References. Lindroth, 1945, 1955a, 1955b, 1963a, and 1966 (natural history); Kirk, 1975c (dispersal power); Wasner, 1979 (biology); Lindroth, ° 1986b (biology); Landry, 1994 (ecology, biology); Luff, 1993 1986 (natural history); Hurka, (ecology); Will et al. 1995 (dispersal power).

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Agonum (Agonum) trigeminum Lindroth, 1954 Ecology. Lowlands. Margins of eutrophic marshes, ponds, and lakes; marshy borders of slow brooks; swamps and low deciduous forests. Ground shaded by bushes (e.g., Alnus, Salix); soft, wet, muddy or clayish soil, rich in organic debris, covered with some vegetation (e.g., Typha, Carex, grass). Mostly nocturnal; usually sheltering during the day in leaf litter, under clumps of grass, logs, fallen trees, and stones. Biology. Seasonality: March-November. Tenerals: early July (PE), late August (QU). Adults found overwintering on higher and drier ground, at the edge of woods, in hawthorn hedges, and in fields; in leaf litter and under stones. Phoretic mites found on adult body. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Favored by spring floods along large rivers. Collecting techniques. Treading the soil with the feet; sweeping plants. References. Lindroth, 1966 (natural history); Larochelle, 1974a (ecology). Agra oblongopunctata oblongopunctata Chevrolat, 1835 Ecology. On thicket vine-covered tangles in a tropical jungle. Biology. Seasonality: JuneJuly. Dispersal power. Wing-condition unknown. Occasional climber (on plants). Collecting technique. Sweeping plants. Akephorus marinus LeConte, 1852 Ecology. Coastal lowlands. Sea beaches. Open ground; moist, sandy bare soil. Nocturnal; sheltering during the day in burrows dug in the soil, under logs, and other debris. Biology. Seasonality: April-August. Adult food, in the field: small crustaceans. Predators: swallows. Dispersal power. Brachypterous, incapable of flight. Slow runner. Strong burrower. Collecting technique. Turning debris. References. Whitehead, 1970 (dispersal power); Fedorenko, 1996 (taxonomy). Akephorus obesus (LeConte, 1866) Ecology. Coastal lowlands. Sea beaches. Open ground; moist, sandy bare soil. Nocturnal; sheltering during the day in burrows dug in the soil, under logs and other debris. Biology. Seasonality: May-August. Predators: staphylinids. Defense-mechanism: when alarmed, the adult may feign death. Dispersal power. Brachypterous, incapable of flight. Slow runner. Strong burrower. Collecting technique. Turning debris. References. Lindroth, 1961 (natural history); Whitehead, 1970 (dispersal power); Richards, 1983 (biology); Fedorenko, 1996 (taxonomy). Amara (Amara) aenea (DeGeer, 1774) Ecology. Lowlands and mountains. Lawns, parks, gardens, vacant lots, cultivated fields and their borders; meadows, roadsides, ski fields, sand pits, and orchards. Open ground;

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dry, usually sandy soil covered with some vegetation. Mostly diurnal; active in the sunshine on the ground and on plants; sheltering on cloudy or cool days under dead leaves, heaps of hay and weeds, pieces of wood, stones, dry cow dung, and basal leaves of plants. Gregarious (in winter). Biology. Seasonality: January-December. Copulating pairs: May-June (QU). Tenerals: July-September. Adults found overwintering on higher and drier ground, at the edge of fields, roadsides and woods; under dead leaves, stones, and in the soil (as deep as 7.5 cm). Adult food: lepidopterous caterpillars, coleopterous larvae, dipterous larvae and pupae, wireworms, and spiders (in the field); seeds of several plant species, corn kernels, lepidopterous caterpillars and pupae, dipterous pupae, and spiders (in captivity). Economic importance: sometimes harmful to winter cereals (Europe). Predators: toads (frequently), frogs, and spiders. Phoretic mites found on adult body. Dispersal power. Macropterous. Frequent flier (in the sunshine; to artificial lights at night; to artificial lights in captivity). Frequent in seashore drift material, which indicates flight ability. Europe: flying in spring (mostly) and in late autumn Moderate runner. Frequent climber (on plants and trees). Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Pitfall trapping; light trapping; using tanglefoot screens; sweeping or beating the vegetation; turning drift material; using window traps. References. Lindroth, 1945, 1955a, 1963a, and 1968 (natural history); Sharova, 1981 (ecology, biology); Erwin, 1981 (natural history); Lindroth, 1986 (natural history); Koch, 1989 (ecology). Amara (Amara) aeneopolita Casey, 1918 Ecology. Lowlands and mountains. Moraines, grasslands, meadows, cultivated fields (e.g., corn), vacant lots, and fencerows. Open ground; firm, rather dry soil composed of sand, clay or gravel, covered with some vegetation. Nocturnal; sheltering during the day under vegetal debris. Biology. Seasonality: June-August. Dispersal power. Macropterous. Frequent in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pitfall trapping; turning vegetal debris; turning drift material. References. Lindroth, 1955a, 1963a, and 1968 (natural history); Kirk, 1975c (ecology). Amara (Curtonotus) alpina (Paykull, 1790) Ecology. Lowlands, mountains, and alpine zone. Heaths, meadows, moraines, wastelands; the drier section of inundation areas of rivers. Open ground; rather dry, preferably sandy or gravelly soil covered with some vegetation (e.g., Empetrum). Nocturnal; sheltering during the day under stones. Biology. Seasonality: May-September. Tenerals: August (ME, NH). Cannibalism observed in captivity. Predators: foxes. Defense-mechanism: when alarmed, the adult may emit a repugnant smell from the pygidial glands. Dispersal power. Wing-polymorphic: macropterous (mostly; probably capable of flight) to brachypterous (incapable of flight). Moderate runner. Collecting techniques. Pitfall trapping; turning stones. References. Lindroth, 1945, 1955a, 1963a, and 1968 (natural history); Schmoller, 1970 (biology); Forsskåhl, 1972 (biology); Lindroth, 1986 (natural history); Luff, 1993 (biology).

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Amara (Zezea) angustata (Say, 1823) Ecology. Lowlands, mountains, subalpine and alpine zones. Meadows, cultivated fields (e.g., alfalfa), pastures, lawns, abandoned fields, vacant lots, relict prairies, moraines, gardens, gravel pits, fencerows, roadsides, vineyards, orchards (e.g., apple), open woodlands, and upper zone of river banks and lake shores. Open ground; moderately moist soil covered with meadow-type vegetation (e.g., Poa). Mostly diurnal; active in the sunshine and before sunset on plants (mostly) and shrubs, and on the first warm days of spring around the base of trees in open woodland pastures; sheltering on cloudy or cool days in leaf litter, under heaps of grass, stones, other debris, and in the soil. Biology. Seasonality: MarchDecember. Adults found overwintering on higher and drier ground, in fields and in woods; in leaf litter and in the soil. Adult food, in the field: seeds (Poa), lepidopterous caterpillars, and mites. Predators: toads and frogs. Dispersal power. Macropterous. Frequent flier (to artificial lights; to artificial lights in captivity). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (on weeds, Poa, Cornus, and Salix). Collecting techniques. Sweeping or beating the vegetation; raking the leaf litter; pitfall trapping; light trapping; turning drift material. References. Webster, 1880 (biology); Blatchley, 1910 (ecology); Lindroth, 1968 (natural history); Kirk, 1975c (ecology); Erwin, 1981 (natural history); Hieke, 2000 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Amara (Zezea) angustatoides Hieke, 2000 Ecology. Unknown. Biology. Seasonality: April-August. Tenerals: April (NY), early August (ME). Dispersal power. Unknown. References. Hieke, 2000 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Amara (Amara) anthobia A. Villa and G.B. Villa, 1833 Ecology. Lowlands. Parks, cultivated fields, and meadows. Greenhouses (occasionally). Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: MarchJuly. Tenerals: June-July (MD). Adults found overwintering (MD). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Occasional in drift material, which indicates flight ability. Strongly favored by human activities. Collecting techniques. Raking the leaf litter; pitfall trapping. References. Erwin, 1981 (natural history); Lindroth, 1986 (ecology, dispersal power); Koch, 1989 (ecology). Amara (Amara) apachensis Casey, 1884 Ecology. Mountains. Biology. Seasonality: March. Dispersal power. Unknown. Reference. Hieke, 1994 (taxonomy). Amara (Bradytus) apricaria (Paykull, 1790) Ecology. Lowlands, mountains, subalpine and alpine zones. Cultivated fields and their borders (e.g., corn, asparagus, wheat, oat, barley, flax, cabbage), grasslands, meadows,

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vacant lots, abandoned fields, lawns, golf courses, gardens, parks, courtyards, gravel pits, sand pits, sand dunes, roadsides, and forest clearings. Open ground; more or less dry soil often consisting of sand or sandy loam, covered with a somewhat close carpet of vegetation (e.g., weeds and grass). Mostly diurnal; active in the sunshine (e.g., on roads); sheltering on cloudy or cool days under stones, dead leaves, and pieces of wood. Larval habitat: in the soil of gardens (Europe). Biology. Seasonality: January, MarchDecember. Gravids: mid-July (AB). Tenerals: May-August, rarely August. Adults found overwintering in fields; in the soil. Adult food: coccinellids and wireworms (in the field); grass seeds, curculionid larvae, and lepidopterous eggs (in captivity). Predators: toads. Parasites: dipterous larvae. Dispersal power. Macropterous. Frequent flier (often swarming to artificial lights at night). Frequent in sea- and lakeshore drift material, which indicates flight ability. Moderate runner. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; using window traps; light trapping; turning drift material. References. Lindroth, 1945, 1955a, 1963a, and 1968 (natural history); Frank, 1971b (biology); Kirk, 1975c (ecology, dispersal ° 1986b (biolopower); Sharova, 1981 (ecology); Lindroth, 1986 (natural history); Hurka, gy); Koch, 1989 (ecology); Luff, 1993 (biology) Will et al. 1995 (dispersal power). Amara (Curtonotus) aulica (Panzer, 1797) Ecology. Lowlands. Meadows, pastures, vacant lots, playgrounds, roadsides, and gravel pits. Often near towns and ports. Open ground; rather dry soil often consisting of clay or sandy loam, covered with meadow-type vegetation (e.g., weeds). Nocturnal and diurnal; active in the sunshine and at night on plants and shrubs; sheltering on cloudy or cool days under stones and vegetal debris. Biology. Seasonality: March-November. Gravids: late August (QU). Tenerals: June-July, rarely August. Adults found overwintering in vacant lots and abandoned fields; in the soil at the base of plants and under stones. Adult food, in the field: seed heads of Compositae. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Frequent in seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (on weeds, Compositae, ferns, Rubus, Cornus, and Alnus). Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Sweeping or beating the vegetation; pitfall trapping; turning stones; light trapping; turning drift material. References. ° Lindroth, 1945, 1955a, 1963a, and 1968 (natural history); Sharova, 1981 (ecology); Hurka, 1986b (biology); Lindroth, 1986 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology). Amara (Amara) aurata Dejean, 1828 Ecology. Lowlands and mountains. Ridges and vicinity of snowfields. Sheltering during the day at the base of stones. Biology. Seasonality: February-May, July-September. Tenerals: April (CA). Predators: bluebirds. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pitfall trapping. References. Lindroth, 1968 (natural history); Kirk, 1975c (ecology).

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Amara (Bradytus) avida (Say, 1823) Ecology. Lowlands and mountains. Cultivated fields (e.g., corn, oat, barley, asparagus, carrot), pastures, meadows, lawns, vacant lots, fencerows, sand pits, sand dunes, gravel pits, lake shores, and orchards (e.g., apple). Open ground; moderately dry, often sandy soil covered with some vegetation (e.g., weeds). Mostly nocturnal; sometimes active on plants in the daytime; usually sheltering during the day under stones, pieces of wood and other debris, and in the soil at the base of plants. Gregarious (in winter). Biology. Seasonality: March-November. Gravids: late July to late August (AB). Tenerals: May-September; mostly in June. Adults found overwintering in vacant lots and sand dunes, at the edge of fields and woods; under stones (mostly) and in rotten logs. Adult food, in the field: strawberries and lepidopterous eggs. Predators: toads, starlings, and asilid flies. Parasites: dipterous larvae. Dispersal power. Macropterous. Frequent flier (at sunset; to artificial lights at night). Regular in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones and pieces of wood; light trapping; turning drift material; sweeping plants. References. Lindroth, 1955a, 1963a, and 1968 (natural history); Frank, 1971b (biology); Kirk, 1975c (ecology); Erwin, 1981 (natural history). Amara (Amara) basillaris (Say, 1823) Ecology. Lowlands. Macrohabitat unknown. Nocturnal; sheltering during the day in wood debris and in clumps of broomsedges. Biology. Seasonality: January, March-December. Adults found overwintering under mullein leaves and in wood debris. Predators: bluebirds and starlings. Parasites: tapeworms. Dispersal power. Macropterous. Occasional in drift material, which indicates flight ability. Moderate runner. Occasional climber (on trees). Collecting techniques. Pitfall trapping; using tanglefoot screens; beating trees. References. Blatchley, 1910 (biology); Kirk, 1969 (ecology, dispersal power), 1970 and 1975c (ecology). Amara (Zezea) belfragei G.H. Horn, 1892 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. References. Hieke, 2000 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Amara (Amara) bifrons (Gyllenhal, 1810) Ecology. Lowlands and mountains. Vacant lots, roadsides, sand pits, sand dunes, gravel pits, lawns, pastures, meadows, and open forests. Open ground; dry, mostly sandy soil covered with sparse vegetation (e.g., weeds). Mostly nocturnal; usually sheltering during the day under stones, dead leaves, heaps of hay and other grasses, pieces of wood, and in the soil at the base of plants. Biology. Seasonality: June-August. Tenerals: June-July. Adult food, in the field: plant seeds. Predators: toads (regularly). Dispersal power. Macropterous. Frequent flier (to artificial lights at night; to artifical lights in captivity). Frequent in seashore drift material, which indicates flight ability. Moderate

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runner. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; raking the leaf litter; light trapping; using window traps; turning drift material. References. Lindroth, 1945, 1955a, 1963a, and ° 1968 (natural history); Sharova, 1981 (ecology, biology); H urka, 1986b (biology); Lindroth, 1986 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology). Amara (Curtonotus) blanchardi Hayward, 1908 Ecology. Uplands. Vicinity of lake margins. Biology. Seasonality: March-September. Tenerals: May-September. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Hieke, 1994 (taxonomy) Amara (Curtonotus) bokori Csiki, 1929 Ecology. Inundation area of a river with sandy soil (AK). Biology. Seasonality: AprilSeptember. Dispersal power. Macropterous, probably capable of flight. Reference. Lindroth, 1968 (natural history). Amara (Bradytus) browni Lindroth, 1968 Ecology. Summit of a steep escarpment, on more or less moist, clayish, bare soil (NT). Biology. Seasonality: June-July. Tenerals: May-June. Dispersal power. Submacropterous, incapable of flight. Moderate runner. Reference. Lindroth, 1968 (natural history). Amara (Amara) brunnea (Gyllenhal, 1810) Ecology. Mountains. A half-shaded place, in the shade of bushes (Alnus), with fine sand (YT). Moraines, under birches, in leaf litter and moss, on sandy or gravelly soil (Europe). Nocturnal; sheltering during the day among dead leaves. Biology. Seasonality: June-July. Dispersal power. Macropterous. Flight records (Europe). Moderate runner. Collecting technique. Raking the leaf litter. References. Lindroth, 1945 and 1968 (natural history); Sharova, 1981 (ecology, biology); Lindroth, 1986 (natural history); Koch, 1989 (ecology). Amara (Amara) brunnipes Motschulsky, 1859 Ecology. Lowlands and mountains. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Reference. Hieke, 1993 (taxonomy). Amara (Amara) californica californica Dejean, 1828 Ecology. Lowlands and mountains. Cultivated fields, roadsides, and forest edges. Moderately dry soil. Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: February, April-July, September-December. Adult food, in the field: lepidopterous caterpillars. Predators: flycatchers and spiders. Parasites: podapolipid mites. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate run-

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ner. Collecting techniques. Pitfall trapping; raking the leaf litter; light trapping. Reference. Hatch, 1953 (ecology). Amara (Curtonotus) carinata (LeConte, 1848) Ecology. Lowlands and uplands. Grasslands, cultivated fields (e.g., corn, alfalfa, wheat, oat, clover, cabbage, flax), pastures, fencerows, gardens, wastelands, vacant lots, sandhills, sand pits; vicinity of alkaline lakes and reservoirs. Often near alkaline water. Open ground; moderately moist, sandy or clayish soil covered with some vegetation. Nocturnal; sheltering during the day under hay, among grass roots, under stones and dry cow dung. Biology. Seasonality: March-November. Copulating pairs: late July and early August (QU), September (SD). Gravids: August-November (ON). Tenerals: May to early July. Adult food: grass material, seeds, fungi, dipterous larvae (in the field); vegetal matter (in captivity). Predators: toads. Parasites: tapeworms. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Occasional climber (on Foxtail Grasses). Effective as a colonist. Strongly favored by human activities. Collecting techniques. Pitfall trapping; inspecting grass roots; light trapping; sweeping plants. References. Rivard, 1964c (biology; as laticollis (LeConte)); Lindroth, 1968a and 1971a (natural history); Kirk, 1975c (natural history); Doane, 1981 (ecology, biology); Hieke, 1994 (taxonomy). Amara (Bradytus) celiana (Stehr, 1949), see Amara (Bradytus) nupera G.H. Horn, 1892. Reference. Hieke, 1994 (taxonomy). Amara (Amara) chalcea Dejean, 1828 Ecology. Lowlands and mountains. Pastures, fields, and sand pits. Open ground; sandy dry soil covered with some vegetation (e.g., grass, weeds). Nocturnal; sheltering during the day under stones, dead leaves, and dry cow dung. Biology. Seasonality: JanuaryDecember. Tenerals: May (CT), August (ON). Adults found overwintering in sand pits; under dead leaves. Predators: bluebirds. Dispersal power. Macropterous. Occasional flier. Occasional in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pitfall trapping; turning stones. Reference. Lindroth, 1968 (natural history). Amara (Amara) ciudadensis (Bates, 1891) Natural History. Unknown. Amara (Amara) coelebs Hayward, 1908 Ecology. Lowlands, mountains, and subalpine zone. Grasslands, meadows, pastures, cultivated fields and their borders (e.g., corn, wheat), and sand pits. Open ground; dry, sandy soil covered with some vegetation (e.g., grass, Artemisia). Nocturnal; sheltering

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André Larochelle & Marie-Claude Larivière

during the day under stones, pieces of wood, and other debris. Biology. Seasonality: March-April, June, August-October. Tenerals: mid-July (AB, MB). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Turning stones and pieces of wood; pitfall trapping. References. Lindroth, 1968 (natural history); Kirk, 1975c (ecology, dispersal power). Amara (Bradytus) colvillensis Lindroth, 1968 Ecology. Inundation area of a river, with rather dry soil composed of clay or very fine sand, bare or covered with sparse vegetation (AK). Biology. Seasonality: June-August. Tenerals: early July (AK) Dispersal power. Macropterous, probably capable of flight. Moderate runner. References. Lindroth, 1968 (natural history); Hieke, 1999 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Amara (Amara) communis (Panzer, 1797) Ecology. Lowlands. Meadows and forests. Moderately dry soil. Nocturnal; sheltering during the day under moss and dry leaves. Plastic in its diel rhythm (Europe). Biology. Seasonality: July. Adult food, in the field (Europe): plant seeds. Dispersal power. Macropterous. Occasional in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Turning moss carpets; raking the leaf litter; pitfall trapping; using window traps. References. Lindroth, 1945 (natural history); Greenslade, 1965 (ecology, biology); Sharova, 1981 (ecology, biology); Lindroth, 1986 (natural histo° 1986b (biology); Koch, 1989 (ecology). ry); Hurka, Amara (Curtonotus) concreta (Casey, 1918), see Amara (Curtonotus) carinata (LeConte, 1848). Reference. Hieke, 1994 (taxonomy). Amara (Amara) conflata LeConte, 1855 Ecology. Lowlands and mountains. Vicinity of a snowfield. Biology. Seasonality: January-July, September-October. Predators: jays. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Amara (Amara) confusa LeConte, 1848 Ecology. Mountains. Grasslands, cultivated fields (e.g., wheat), vacant lots, and borders of fields. Open ground; dry soil composed of clay-mixed gravel, covered with some vegetation (e.g., grass). Mostly nocturnal; usualy sheltering during the day under stones or under the basal leaves of dandelions (Taraxacum). Biology. Seasonality: April-September. Tenerals: early July (BC). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Turning stones; pitfall trapping. Reference. Lindroth, 1968 (natural history).

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Amara (Amara) convexa LeConte, 1848 Ecology. Lowlands, mountains, subalpine and alpine zones. Grasslands, prairies, meadows, pastures, rangelands, cultivated fields and their borders (e.g., corn, wheat, barley), vacant lots, roadsides, sand pits, sandhills, sand blowouts, gravel pits; vicinity of water bodies (western North America). Open ground; dry, usually sandy, often soft and dusty soil covered with some vegetation (e.g., grass, weeds). Mostly nocturnal; usually sheltering during the day in the soil at the base of plants, under reindeer moss (Cladonia), stones, and dry cow dung, and in ant nests. Gregarious (up to 20 individuals under a single piece of cow dung). Biology. Seasonality: March-November. Copulating pairs: July (QU). Tenerals: August-September. Adults found overwintering in vacant lots, pastures, and sand pits; under plant rosettes (Verbascum) and stones. Adult food, in the field: coccinellids. Predators: toads (regularly), frogs, and crows. Parasites: mites. Dispersal power. Macropterous. Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; digging at the base of plants; turning reindeer moss; turning drift material. References. Lindroth, 1968 (natural history); Bell, 1971 (ecology); Kirk, 1975c (ecology). Amara (Amara) crassispina LeConte, 1855 Ecology. Lowlands and mountains. Old fields, grasslands, and pastures. Open ground; soil covered with sparse vegetation (e.g., grass). Biology. Seasonality: January-August, October-November. Adults found overwintering in fields; under plant rosettes (Verbascum) and in clumps of broomsedges. Dispersal power. Macropterous. Occasional in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Inspecting clumps of broomsedges; pitfall trapping. Reference. Kirk, 1969 (biology). Amara (Amara) cupreolata Putzeys, 1866 Ecology. Lowlands and mountains. Grasslands, relict prairies, meadows, pastures, vacant lots, lawns, golf courses, cultivated fields and their borders (e.g., corn, wheat, barley, rye, alfalfa, soybean, dry bean, tobacco, strawberry, asparagus, carrot), fencerows, roadsides, sand pits, sandhills, gravel pits, orchards (e.g., apple, peach), forest clearings, tree plantations (e.g., Pinus), and vicinity of water bodies. Open ground; dry, often sandy soil covered with some tall vegetation (e.g., weeds). Mostly diurnal; quite active in the sunshine; sheltering on cloudy or cool days under stones, dead leaves, logs, pieces of wood, heaps of hay, and reindeer moss (Cladonia). Biology. Seasonality: January-December. Copulating pairs: April (CT), May (QU). Gravids: mostly May-June (QU). Tenerals: April, June; mostly July-August. Adults found overwintering on higher and drier ground, in vacant lots, hills, sand pits, thickets, and woods; in leaf litter, wood debris, under stones, logs, and in the soil (as deep as 10 cm). Adult food: plant seeds, lepidopterous caterpillars, animal matter (in the field); various grass seeds, lepidopterous caterpillars, curculionid larvae, and spiders (in captivity). Predators: toads and frogs. Dispersal power. Wing-

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dimorphic: submacropterous (mostly; incapable of flight) or macropterous. Frequent flier (at sunset; to artificial lights at night). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on alfalfa). Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; raking the leaf litter; light trapping; turning drift material; sweeping plants. References. Dogger and Olson, 1966 (biology); Lindroth, 1968 (natural history); Kirk, 1969 (biology) and 1975c (ecology, dispersal power); Levesque et al. 1979 and 1980 (biology); Ferguson and McPherson, 1985 (biology); Barney and Pass, 1986a (biology); Epstein and Kulman, 1990 (ecology, biology); Will et al. 1995 (dispersal power). Amara (Curtonotus) daurica Motschulsky, 1845 Ecology. Prostrate Salix-grasses associations (NT). Biology. Seasonality: June-July. Dispersal power. Unknown. Reference. Hieke, 1994 (taxonomy). Amara (Curtonotus) deparca (Say, 1830) Ecology. Uplands. Biology. Seasonality: August. Dispersal power. Unknown. Reference. Hieke, 1993 (taxonomy). Amara (Amara) discors Kirby, 1837 Ecology. Mountains, subalpine and alpine zones. Moraines, meadows, pastures, vacant lots, gravel pits, sand pits, sandhills, forest clearings, vicinity of stream banks and snowfields. Open ground; rather dry, sandy soil covered with sparse vegetation. Nocturnal; sheltering during the day under stones, dead leaves, and pieces of wood. Gregarious. Biology. Seasonality: April-September. Tenerals: late July-August. Adults found overwintering in pastures; in the soil and under plant rosettes (Verbascum). Dispersal power. Wing-dimorphic: macropterous (mostly) or brachypterous (incapable of flight). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pitfall trapping; turning stones; raking dead leaves. References. Lindroth, 1963a and 1968 (natural history). Amara (Amara) ellipsis (Casey, 1918) Ecology. Lowlands, mountains, and subalpine zone. Grasslands, relict prairies, meadows, cultivated fields (e.g., wheat, barley), vacant lots, and fencerows. Open ground; rather dry, sandy or gravelly soil covered with some vegetation (e.g., grass or weeds). Nocturnal; sheltering during the day under pieces of wood. Biology. Seasonality: AprilNovember. Gravids: April to early July (AB). Tenerals: July. Adult food, in captivity: lepidopterous eggs. Parasites: dipterous larvae. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; turning pieces of wood. References. Lindroth, 1968 and 1971a (natural history); Frank, 1971b (biology); Kirk, 1975c (ecology, dispersal power); Kavanaugh, 1992 (natural history).

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Amara (Amara) emancipata Lindoth, 1968 Ecology. Lowlands and mountains. A dry hill covered with sparse pines and steppe vegetation. Biology. Seasonality: April-May. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Lindroth, 1968 (natural history). Amara (Amara) erratica (Duftschmid, 1812) Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, meadows, vacant lots, roadsides, and forest clearings. Open ground; rather dry, usually sandy soil covered with some vegetation (e.g., grass, weeds). Mostly nocturnal; usually sheltering during the day under stones, dead leaves, and pieces of wood. Biology. Seasonality: April-September. Tenerals: July-August; mostly August. Adult food, in captivity (Europe): mealworms. Predators: trushes. Dispersal power. Macropterous. Occasional flier (to artificial lights in captivity). Moderate runner. Collecting techniques. Pitfall trapping; turning stones. References. Lindroth, 1945, 1955a, 1963a, and 1968 (natural history); Bíly, 1971 and 1975 (biology); De Zordo, 1979 (biology); Lindroth, 1986 (natural ° 1986b (biology); Koch, 1989 (ecology); Luff, 1993 (biology). history); Hurka, Amara (Bradytus) exarata Dejean, 1828 Ecology. Lowlands and mountains. Cultivated fields (e.g., corn, wheat, oats, alfalfa, flax, clover), borders of fields, relict prairies, pastures, vacant lots, sand pits, upper zone of river banks, orchards (e.g., peach), edges of forests, and open forests. Open ground; dry sandy soil. Nocturnal; sheltering during the day in the soil under dead leaves, stones and pieces of wood. Gregarious (in winter). Biology. Seasonality: January-November. Tenerals: May-September; mostly May-June. Adults found overwintering on higher and drier ground, in fields, at the edge of forests, and in woods; in the soil and under dead leaves. Predators: woodpeckers and flycatchers. Dispersal power. Macropterous. Regular flier (to artificial lights at night). Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; digging into the soil; light trapping. References. Blatchley, 1910 (biology); Lindroth, 1968 (natural history); Kirk, 1975c (ecology, dispersal power); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Amara (Amara) exlineae Minsk and Hatch, 1939 Ecology. Mountains. Open parks. Nocturnal; sheltering during the day under stones. Biology. Seasonality: July-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Amara (Amara) eyrinota (Panzer, 1797) Ecology. Lowlands. Open country. Europe: fallow fileds and wastelands. Soil moderately dry, often composed of clay-mixed gravel, covered with sparse vegetation (Eu-

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André Larochelle & Marie-Claude Larivière

rope). Biology. Seasonality: March-September. Adult food, in the field (Europe): plant matter (e.g., seeds). Dispersal power. Macropterous. Flight records (Europe). Moderate runner. Clearly effective as a colonist. Collecting technique. Pitfall trapping. References. Lindroth, 1945 (natural history); Bíly, 1972 (biology); Sharova, 1981 (ecology, biolo° gy); Lindroth, 1986 (natural history); Hurka, 1986b (biology); Koch, 1989 (ecology); Luff, 1993 (biology). Amara (Amara) familiaris (Duftschmid, 1812) Ecology. Lowlands, mountains, subalpine and alpine zones. Meadows, pastures, vacant lots, cultivated fields and their borders (e.g., wheat, alfalfa, asparagus, corn, hay, raspberry), roadsides, gardens, golf courses, lawns, sand dunes, sand pits, gravel pits, orchards (e.g., apple), forest edges and clearings, open forests. Greenhouses (occasionally). Open ground; moderately dry, sandy soil covered with some vegetation (e.g., Stellaria, Polygonum). Mostly diurnal; quite active in the sunshine; sheltering on cloudy or cool days under stones, dead leaves, heaps of hay, pieces of wood, and in the soil. Biology. Seasonality: March-October, December. Tenerals: July-August. Adults found overwintering on higher and drier ground, on hills, in vacant lots, at the edge of woods, and in woods; under stones, in leaf litter, and in the soil. Adult food: dipterous larvae and pupae, elaterid larvae (in the field); grass seeds, aphids, and lepidopterous caterpillars (in captivity). Adult food, in the field (Europe): flowers or seeds of Stellaria, Poa, and Cerastium. Predators: toads. Dispersal power. Macropterous. Frequent flier (in the sunshine, at sunset; to artificial lights at night; into rotary traps). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants and trees). Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; raking the leaf litter; light trapping; turning drift material; using rotary traps; using window traps; sweeping plants; beating trees. References. Lindroth, 1945, 1955a, 1963a, and 1968 (natural history); Sharova, 1981 (ecology, biology); Erwin, 1981 (natural his° tory); Hurka, 1986b (biology); Lindroth, 1986 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology). Amara (Amara) farcta LeConte, 1855 Ecology. Lowlands and mountains. Sagebrush-steppes, rangelands, pastures, meadows, cultivated fields (e.g., alfalfa, sainfoin), sand blowouts; vicinity of human habitations, alkaline lakes, rivers, and brooks. Frequently near alkaline water. Open ground; dry, sandy or sandy-loamy soil covered with some vegetation (e.g., weeds, grass). Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under stones. Biology. Seasonality: February-October. Gravids: May and early July (AB). Tenerals: July. Adult food, in the field: plant material, weevils, flies and their larvae. Adult and larval food, in captivity: vegetal matter. Predators: spiders. Parasites: dipterous larvae. Dispersal power. Macropterous. Occasional flier. Moderate runner. Occasional climber (on plants). Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; sweeping plants. References. Lindroth, 1968 (natural

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history); Frank, 1971b (biology); Kirk, 1975c (ecology); Doane, 1981 (ecology, biology); Lester and Morrill, 1989 (ecology). Amara (Zezea) flebilis (Casey, 1918) Ecology. Unknown. Biology. Seasonality: May-June, December. Dispersal power. Brachypterous, incapable of flight. References. Hieke, 2000 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Amara (Neopercosia) fortis LeConte, 1880 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Amara (Bradytus) fulva (O.F. Müller, 1776) Ecology. Lowlands. Vacant lots, sand dunes; upper zone of sea beaches and river banks. Open ground; dry soil usually composed of fine shifting sand, covered with sparse vegetation (e.g., grass). Frequently near the sea. Nocturnal; sheltering during the day in the soil under pieces of wood, stones, and heaps of hay. Gregarious. Larval habitat: in the soil. Biology. Seasonality: May-August. Tenerals: late June. Adult food, in the field (Europe): plant material (mostly), ants, aphids, spiders, mites, and flies. Defensemechanism: body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (to artificial lights in captivity). Frequent in seashore drift material, which indicates flight ability. Moderate runner. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning pieces of wood; using window traps; sweeping; turning drift material. References. Lindroth, 1945, 1955a, 1963a, and 1968 (natural history); Sharova, 1981 (ecology, biology); ° Hurka, 1986b (biology); Lindroth, 1986 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology). Amara (Amara) gibba (LeConte, 1848) Ecology. Lowlands. Moraines, meadows, vacant lots, cultivated fields (e.g., cabbage), and sand pits. Open ground; rather dry, sandy soil covered with sparse vegetation. Nocturnal; sheltering during the day. Biology. Seasonality: May-October. Tenerals: early and late July (QU). Predators: toads and spiders. Dispersal power. Macropterous. Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strongly favored by human activities. Collecting technique. Pitfall trapping. Amara (Bradytus) glacialis (Mannerheim, 1853) Ecology. Inundation areas of river banks and lake shores. Open ground; flat, dry, sandy soil, bare or covered with sparse vegetation (e.g., Epilobium). Nocturnal; sheltering during the day under stones and pieces of wood. Biology. Seasonality: June-August. Tenerals: late June (BC), late July (ON). Dispersal power. Macropterous, probably capable of

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flight. Moderate runner. Collecting technique. Turning stones. Reference. Lindroth, 1968 (natural history). Amara (Amara) harpalina LeConte, 1855 Natural History. Unknown. Amara (Amara) hicksi Lindroth, 1968 Ecology. Unknown. Biology. Seasonality: June-August. Dispersal power. Macropterous, probably capable of flight. Amara (Curtonotus) hyperborea Dejean, 1831 Ecology. Lowlands and mountains. Moraines, moorlands, meadows, vacant lots, sand pits, gravel pits, and roadsides. Ground open or shaded by bushes (e.g., Alnus, Salix); rather dry, sandy soil covered with some vegetation (e.g., grass, Empetrum). Mostly nocturnal; usually sheltering during the day under dead leaves, stones, pieces of wood and other debris. Biology. Seasonality: April-August. Tenerals: late June to late August. Adult food, in the field: grass seeds. Predators: toads and bluebirds. Dispersal power. Macropterous. Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Strongly favored by human activities. Collecting techniques. Pitfall trapping; raking the leaf litter; turning pieces of wood; turning drift material; sweeping plants. References. Lindroth, 1955a, 1963a, and 1968 (natural history). Amara (Amara) idahoana (Casey, 1924) Ecology. Lowlands, mountains, subalpine and alpine zones. Grasslands and meadows. Open ground; soil covered with some vegetation (e.g., grass). Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under stones. Biology. Seasonality: JuneJuly. Adult food, in the field: plant seeds. Dispersal power. Wing-dimorphic: macropterous (mostly; probably capable of flight) or brachypterous (incapable of flight). Moderate runner. Occasional climber (on plants). Collecting techniques. Pitfall trapping; turning stones; sweeping plants. Reference. Lindroth, 1968 (natural history). Amara (Triaena) immunda (Casey, 1918), see Amara (Zezea) scitula Zimmermann, 1832 References. Hieke, 1994 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Amara (Amara) impuncticollis (Say, 1823) Ecology. Lowlands, mountains, subalpine and alpine zones. Grasslands, meadows, pastures, rangelands, lawns, cultivated fields (e.g., soybean, alfalfa, cabbage, hop, corn,

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oat, hay), abandoned fields, gravel pits, sand pits, roadsides, forest edges and clearings, and open forests. Open ground; moderately dry, often sandy soil covered with rich discontinuous vegetation (e.g., weeds). Diurnal (mostly) and nocturnal; active in the sunshine on the ground and on plants; sheltering on cloudy or cool days under dead leaves, stones, heaps of hay, plant rosettes (Verbascum), logs, and in moss. Biology. Seasonality: January-October. Gravids: early May (QU). Tenerals: April-September. Adults found overwintering on higher and drier ground, in the upper zone of roadsides and sand pits, in hawthorn hedges, at the edge of woods, in wood clearings, and in woods; in leaf litter, under stones, and in the soil. Adult food: grass material, plant seeds, fungi, lepidopterous caterpillars, a grasshopper (in the field); plant material (e.g., oats), fruits, lepidopterous caterpillars, carabid pupae, meat, and cheese (in captivity). Larval food, in captivity: oats. Predators: toads, frogs, bluebirds, woodpeckers, flycatchers, robins, and starlings. Dispersal power. Macropterous. Frequent flier (in the sunshine; at sunset; to artificial lights at night; to artifical lights in captivity). Moderate runner. Occasional climber (on plants, e.g., Gramineae). Strongly favored by human activities. Collecting techniques. Pitfall trapping; raking the leaf litter; turning stones; light trapping; sweeping plants. References. Lindroth, 1955a, 1963a, and 1968 (natural history); Kirk, 1975c (ecology, dispersal power); Erwin, 1981 (natural history); Ferguson and McPherson, 1985 (biology); Hieke, 2000 (taxonomy). Amara (Zezea) inexspectata Hieke, 1990 Ecology. Unknown. Biology. Seasonality: February-April, June-July, December. Dispersal power. Macropterous, probably capable of flight. References. Hieke, 1994 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Amara (Bradytus) insignis Dejean, 1831 Ecology. Lowlands and mountains. Burrows of Ground squirrels. Biology. Seasonality: February, April; fall. Dispersal power. Macropterous, probably capable of flight. Moderate runner. References. Hieke, 1995 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Amara (Bradytus) insularis G.H. Horn, 1875 Ecology. Unknown. Biology. Seasonality: spring and early summer. Dispersal power. Brachypterous, incapable of flight. References. Hieke, 1995 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Amara (Amara) interstitialis Dejean, 1828 Ecology. An open site, with rather dry soil covered with sparse vegetation (e.g., weeds) (AK). Cultivated fields (Europe). Biology. Seasonality: April, June-August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Flight records (in the sunshine; Europe). Moderate runner. Collecting technique. Pitfall trapping. References. Lindroth, 1945, 1968 and 1986 (natural history).

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Amara (Curtonotus) jacobina LeConte, 1855 Ecology. Macrohabitat unknown. Greenhouses (occasionally). Biology. Seasonality: February, April-June. Adult food, in the field: strawberries. Dispersal power. Unknown. Amara (Amara) jucunda Csiki, 1929 Natural History. Unknown. Amara (Zezea) kavanaughi Hieke, 1990 Ecology. Mountains. A moist roadside ditch covered with grass. Edges of a large vernal pond in pasture, on clay soil. Biology. Seasonality: February-June, October. Dispersal power. Macropterous, probably capable of flight. Moderate runner. References. Hieke, 1994 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Amara (Curtonotus) kurnakowi Hieke, 1994 Ecology. A beach, with soil composed of gravel, clay, and sand. An area covered with grass, Artemisia and Lupinus. Biology. Seasonality: June-July. Dispersal power. Submacropterous, probably incapable of flight. Moderate runner. Reference. Hieke, 1994 (taxonomy). Amara (Curtonotus) lacustris LeConte, 1855 Ecology. Lowlands and uplands. Grasslands, cultivated fields and their borders (e.g., wheat, barley, asparagus); vicinity of lakes and alkaline ponds. Open ground; rather dry, clay soil covered with discontinuous vegetation (e.g., grass). Nocturnal; sheltering during the day under stones and logs. Biology. Seasonality: March, May-August, October. Gravids: early August (SA). Tenerals: June-August. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; light trapping; turning stones and logs. References. Lindroth, 1968 (natural history); Frank, 1971b (biology); Kirk, 1975c (dispersal power). Amara (Amara) laevipennis Kirby, 1837 Ecology. Lowlands, mountains, and subalpine zone. Moraines, grasslands, meadows, cultivated fields (e.g., barley), vacant lots, roadsides, sand pits, gravel pits, ski fields; dry lake shores and forest clearings. Open ground; rather dry, usually sandy soil covered with sparse or moderate vegetation (e.g., grass, weeds). Nocturnal; sheltering during the day under dead leaves, stones, pieces of wood and paper, heaps of hay, and in the soil at the base of plants. Biology. Seasonality: March-October. Tenerals: late June to mid-August; mostly after mid-July. Adults found overwintering on higher and drier

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ground, in the upper zone of sand pits and in woods; under stones and in the soil under dead leaves. Predators: toads. Dispersal power. Macropterous. Occasional flier (into rotary traps). Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter; using rotary traps. References. Lindroth, 1963a and 1968 (natural history). Amara (Bradytus) latior (Kirby, 1837) Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, vacant lots, cultivated fields (e.g., corn, oat, alfalfa, cabbage, flax, barley, wheat), meadows, pastures, sand pits, gravel pits, sand dunes, roadsides, forest edges and clearings, open forests, and tree plantations (e.g., Pinus). Ground open or shaded by bushes (e.g., Alnus); moderately dry, sandy soil covered with sparse vegetation. Nocturnal; sheltering during the day in the soil under plant rosettes, dead leaves, stones, pieces of wood or cardboard, and under heaps of hay. Biology. Seasonality: April-December. Copulating pairs: September (QU). Gravids: August-November (AB, ON). Tenerals: May-July (mostly), August-October. Adults found overwintering in vacant lots, pastures, at the edge of sand pits and gravel pits, and in woods; in the soil, under plant rosettes (Verbascum), and stones. Adult food: grasshopper eggs (in the field); carabid pupae and lepidopterous eggs (in captivity). Predators: toads (frequently) and frogs. Defense-mechanism: when alarmed, the adult may emit a repugnant smell that clings to fingers when the beetle is seized. Dispersal power. Macropterous. Frequent flier (at sunset; to artificial lights at night). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; light trapping; turning drift material; digging at the base of plants. References. Lindroth, 1963a and 1968 (natural history); Rivard, 1964c (biology); Frank, 1971b (biology); Kirk, 1975c (natural history). Amara (Bradytus) lindrothi Hieke, 1990 Ecology. Uplands. Biology and Dispersal power. Unknown Amara (Amara) littoralis Mannerheim, 1843 Ecology. Lowlands and mountains. Grasslands, meadows, pastures, cultivated fields (e.g., corn, alfalfa, wheat, barley, pea, cabbage, sweet clover, turnip, carrot, bean), vacant lots, fencerows, roadsides, sand pits, gravel pits, gardens, sand blowouts, and open forests. Open ground; moderately dry, often sandy soil covered with rich discontinuous vegetation (e.g., weeds). Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under dead leaves and plant rosettes, in the soil, under stones, pieces of wood, and heaps of hay. Biology. Seasonality: February-November. Gravids: mid-May to late July (AB). Tenerals: April, July-September. Adults found overwintering in sand pits, gravel pits, and woods; under dead leaves, plant rosettes (Verbascum), stones, and in the soil. Adult food: grasshopper eggs (in the field); fruits, meat, and a dead fly (in captivity). Reared on a completely vegetarian diet. Predators: toads, crows, starlings, and swallows. Parasites: dipterous larvae and tapeworms. Dispersal power. Macropterous.

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Occasional flier (at sunset; to artificial lights in captivity). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; raking the leaf litter; turning plant rosettes. References. Lindroth, 1968 (natural history); Frank, 1971b (biology); Kirk, 1975c (ecology, dispersal power); Kavanaugh, 1992 (natural history); Boivin and Hance, 1994 (biology); Will et al. 1995 (dispersal power); Hieke, 1994 and 2000 (taxonomy). Comments. Despite its name, species not associated with water. Amara (Zezea) longula LeConte, 1855 Ecology. Lowlands and mountains. Biology. Seasonality: April-July. Predators: blackbirds, meadowlarks, flycatchers, and thrushes. Dispersal power. Macropterous, probably capable of flight. References. Hieke, 1994 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Amara (Amara) lunicollis Schiødte, 1837 Ecology. Lowlands and mountains. Grasslands, meadows, cultivated fields, vacant lots, moraines, gravel pits, ski fields, orchards (e.g., apple), edges of forests, forest clearings, thickets (Alnus), and open forests. Open ground; not too dry, gravelly or peaty soil covered with some vegetation (e.g., weeds). Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under stones, dead leaves, pieces of wood, heaps of hay, in the soil at the base of plants and under fallen branches. Biology. Seasonality: AprilNovember. Adults found overwintering in fallow fields and woods; in the soil and under dead leaves. Adult food, in the field: earwigs. Dispersal power. Macropterous. Occasional flier (at sunset; to artificial lights in captivity). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; raking the leaf litter. References. Lindroth, 1945, 1955a, 1963a and 1968 (natural history); Grenslade, 1965 ° (biology); Sharova, 1981 (ecology, biology); Hurka, 1986b (biology); Lindroth, 1986 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology); Hieke, 1994 (taxonomy). Amara (Amara) maxwelli (Casey, 1924), see Amara (Amara) nupera G.H. Horn, 1892. References. Hieke, 1994 and 1997 (taxonomy). Amara (Amara) merula (Casey, 1918) Ecology. Uplands. Dry grass openings at tree line (CO). Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Reference. Hieke, 1993 (taxonomy). Amara (Amara) musculis (Say, 1823) Ecology. Lowlands, mountains, subalpine and alpine zones. Meadows, pastures, vacant lots, cultivated fields and their borders (e.g., corn, wheat, alfalfa, carrot), sand pits,

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gravel pits, gardens, orchards (e.g., peach), forest edges and clearings, and open forests. Open ground; dry, usually sandy soil covered with sparse vegetation (e.g., Solidago). Mostly diurnal; active in the sunshine on Solidago flowers; sheltering on cloudy or cool days in the soil at the base of plants (Solidago), under plant rosettes (Verbascum), stones, pieces of wood, and heaps of hay. Gregarious (in winter). Biology. Seasonality: JanuaryNovember. Copulating pairs: late August (QU). Tenerals: May to early August; mostly spring and early summer. Adults found overwintering in vacant lots; under stones and wood debris. Adult food, in the field: plant seeds, pollen, and animal matter. Predators: toads (regularly), frogs, and starlings. Parasites: tapeworms. Defense-mechanism: when alarmed, the adult may emit a strong fetid odor. Dispersal power. Macropterous. Frequent flier (often swarming; during the day; to artificial lights at night; to artifical lights in captivity). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (on plants, e.g., Solidago, Gutierrezia, Zia). Strongly favored by human activities. Collecting techniques. Sweeping plants; digging at the base of plants. References. Blatchley, 1910 (ecology); Lindroth, 1968 (natural history); Kirk, 1969 (biology) and 1975c (ecology, dispersal power); Erwin, 1981 (natural history). Amara (Bradytus) neomexicana (Casey, 1924) Ecology. Unknown. Biology. Seasonality: May, July. Dispersal power. Unknown. References. Hieke, 1994 and 1997 (taxonomy). Amara (Amara) nexa (Casey, 1918) Natural History. Unknown. Amara (Bradytus) novella (Stehr, 1949), see Amara (Bradytus) neomexicana (Casey, 1924). References. Hieke, 1994 and 1997 (taxonomy). Amara (Amara) nupera G.H. Horn, 1892 Ecology. Uplands. Biology. Seasonality: March. Dispersal power. Unknown. References. Hieke, 1994 and 1997 (taxonomy). Amara (Percosia) obesa (Say, 1823) Ecology. Lowlands, mountains, subalpine and alpine zones. Rangelands, meadows, pastures, cultivated fields (e.g., corn, alfalfa, oat, flax, wheat, barley, cabbage, sweet clover), fencerows, vacant lots, sand pits, sandhills, sand dunes, roadsides, gravel pits, upper zone of seashore and river banks. Ground open or shaded by bushes (e.g., Alnus, Salix); dry, usually sandy soil covered with sparse vegetation. Nocturnal; sheltering during the day under stones, dead leaves, and pieces of wood. Biology. Seasonality: February-December. Gravids: August-October (ON). Tenerals: mid-May to September. Adults found

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overwintering on higher and drier ground, in vacant lots, meadows, on hills and in the upper zone of sand pits, and at the edge of woods; under stones (mostly), pieces of cardboard, and in the soil (as deep as 2.5 cm). Adult food, in the field: grasshopper eggs and nymphs. Adult and larval food, in captivity: grasshopper eggs. Repeatedly reared from grasshopper eggs. Predators: toads (frequently), bluebirds, and spiders. Parasites: tapeworms and entomogenous fungi. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous (probably capable of flight). Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones. References. Lindroth, 1963a and 1968 (natural history); Rivard, 1964c (biology); Larochelle, 1977i (dispersal power); Kirk, 1975c (natural history). Amara (Amara) otiosa Casey, 1918 Natural History. Unknown. Reference. Hieke, 2000 (taxonomy). Amara (Amara) ovata (Fabricius, 1792) Ecology. Lowlands and uplands. Vacant lots, roadsides, upper zone of river banks, and open forests. Open ground; dry, sandy or gravelly soil covered with sparse vegetation. Mostly nocturnal; usually sheltering during the day under dead leaves and other vegetal debris. Biology. Seasonality: April-June. Adults found overwintering in vacant lots; under dead leaves. Adult food, in the field (Europe): seeds, flowers or fruits of Alliaria, Barbarea, Brassica, Reseda, and Cynosurus. The larva is carnivorous. Dispersal power. Macropterous. Frequent in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Clearly effective as a colonist. Collecting techniques. Pitfall trapping; raking the leaf litter; turning drift material; sweeping plants. ° References. Lindroth, 1945 (natural history); Sharova, 1981 (ecology, biology); Hurka, 1986b (biology); Lindroth, 1986 (natural history); Koch, 1989 (ecology). Amara (Zezea) pallipes Kirby, 1837 Ecology. Lowlands, mountains, subalpine and alpine zones. Meadows, abandoned fields, vacant lots, cultivated fields and their borders (e.g., corn, wheat, barley, tobacco), roadsides, sand pits, gravel pits, upper zone of river and brook banks, and open forests. Frequently near water. Open ground; moderately moist soil covered with meadow-type grass (e.g., Poa). Mostly diurnal; active in the sunshine on plants (e.g., Poa); sheltering on cloudy or cool days under stones, dead leaves, and at the base of plants. Biology. Seasonality: April-December. Gravids: early (QU) and late (AB) July. Tenerals: April (QU), late August (AB). Adults found overwintering in fields; in the soil. Adult food, in the field: grass seeds (Poa). Predators: toads. Dispersal power. Macropterous. Occasional in seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (on plants). Strongly favored by human activities. Collecting techniques. Sweeping plants; turning stones. References. Lindroth, 1968 (natural history); Frank, 1971b (biology); Kirk, 1975c (ecology); Hieke, 1994 (taxonomy); Will et al. 1995 (dispersal power); Ball and Bousquet, 2001 (taxonomy).

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Amara (Amara) patruelis Dejean, 1831 Ecology. Lowlands, mountains, and subalpine zone. Moraines, meadows, cultivated fields and their borders (e.g., wheat, barley, asparagus, carrot), fallow fields, vacant lots, roadsides, ski fields, sand pits, gravel pits, vicinity of human habitations; lawns, forest edges and clearings. Open ground; rather dry, often sandy soil covered with sparse vegetation (e.g., weeds). Often near human habitations. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under stones, dead leaves, heaps of hay, pieces of cardboard, and in the soil. Gregarious (in winter). Biology. Seasonality: February-December. Copulating pairs: early May (QU), early June (AB); coitus lasting about one minute. Gravids: April and late July (AB). Tenerals: late June to August; mostly July. Adults found overwintering on higher and drier ground, in vacant lots, meadows, in the upper zone of sand pits, at the edge of woods, and in woods; under stones, dead leaves, pieces of wood, and in the soil. Adult food, in captivity: lepidopterous caterpillars. Parasites: dipterous larvae. Dispersal power. Macropterous. Occasional flier (in the sunshine; to artificial lights in captivity). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; raking the leaf litter. References. Lindroth, 1955a, 1963a and 1968 (natural history); Frank, 1971b (biology); Kirk, 1975c (ecology); Niemelä et al. 1992 (ecology, biology). Amara (Curtonotus) pennsylvanica Hayward, 1908 Ecology. Lowlands and mountains. Cultivated fields (e.g., corn, tobacco), pastures, old fields, hilly fields, upper zone of river banks, sand pits, gravel pits, and open forests. Open ground; dry, sandy soil covered with sparse vegetation. Nocturnal; sheltering during the day under stones. Biology. Seasonality: February-November. Tenerals: midMay to early June (CT, IL, MD, OH). Adults found overwintering (IN). Predators: crows, starlings, and woodpeckers. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Frequent in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Collecting techniques. Pitfall trapping; turning stones; turning drift material; sweeping plants. References. Blatchley, 1910 (biology); Herne, 1963 (biology); Lindroth, 1968 (natural history); Kirk, 1975c (dispersal power); Anderson et al. 1995 (ecology). Amara (Amara) pomona Casey, 1918, see Amara (Amara) brunnipes Motschulsky, 1859 Reference. Hieke, 1993 (taxonomy). Amara (Amara) pseudobrunnea Lindroth, 1968 Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, vacant lots, roadsides, and open forests. Ground shaded by bushes or open; dry sandy soil covered with sparse vegetation. Nocturnal; sheltering during the day under dead leaves, stones, and

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pieces of cardboard. Biology. Seasonality: June-September. Dispersal power. Macropterous. Frequent in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter; turning drift material. Reference. Lindroth, 1968 (natural history). Amara (Curtonotus) pterostichina Hayward, 1908 Ecology. Uplands. Biology. Unknown. Dispersal power. Macropterous. Occasional flier (during the day). Amara (Amara) quenseli (Schönherr, 1806) Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, meadows, grasslands, cultivated fields and their borders (e.g., corn, flax, barley, wheat), vacant lots, roadsides, shrub-steppes, sand pits, sand dunes, gravel pits, upper zone of lake shores, and vicinity of human habitations. Open or shaded ground, under of bushes (Alnus); dry, more or less sandy soil covered with sparse vegetation. Nocturnal; sheltering during the day in the soil (often) under stones, dead leaves, heaps of hay, plant rosettes, planks, and logs. Larval habitat: in the soil in fields. Biology. Seasonality: AprilNovember. Gravids: late July to late August (AB). Tenerals: May-August; rarely August. Adults found overwintering on higher and drier ground, in meadows, hills, and in the upper zone of sand pits; under stones. Adult food: lepidopterous eggs (in captivity); animal and plant matters (in the field, Europe). Predators: toads (frequently) and frogs. Parasites: dipterous larvae. Dispersal power. Wing-polymorphic: macropterous (capable of flight) to brachypterous (incapable of flight). Occasional flier (to artificial lights at night). Frequent in seashore drift material, which indicates flight ability. Moderate runner. Strong burrower. Strongly favored by human activities. Collecting techniques. Digging under stones and dead leaves; pitfall trapping; turning drift material. References. Darlington, 1936 (dispersal power; as remotestriata Dejean); Lindroth, 1945, 1955a, 1963a and 1968 (natural history); Frank, 1971b (biology); Kirk, 1975c (ecology, dispersal power); Bíly, 1975 (biology); Bengston and Erikstad, 1984 (dispers° al power); Hurka, 1986b (biology); Lindroth, 1986 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology). Amara (Amara) rectangula LeConte, 1855 Natural History. Unknown. Amara (Amara) rubrica Haldeman, 1843 Ecology. Lowlands. Meadows, vacant lots, cultivated fields (e.g., tobacco), sandhills, sand pits, and gravel pits. Open ground; dry, sandy soil covered with sparse vegetation (e.g., Solidago). Mostly diurnal; active in the sunshine on plants (e.g., Solidago); sheltering on cloudy or cool days under stones, in the soil at the base of plants, and under pieces of wood. Biology. Seasonality: March-November. Tenerals: early May

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(MD) to mid-June (ON, QU). Adults found overwintering in meadows, on hills, and at the edge of woods; in the soil (as deep as 7.5 cm), under stones, and in leaf litter. Adult food, in the field: seed heads of Compositae (Solidago). Predators: toads and frogs. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; to artifical lights in captivity). Moderate runner. Frequent climber (on plants, e.g., Solidago). Collecting techniques. Sweeping plants; light trapping; turning stones; digging at the base of plants. References. Lindroth, 1968 (natural history); Kirk, 1975c (dispersal power). Amara (Amara) sanjuanensis Hatch, 1949 Ecology. Unknown. Biology. Seasonality: April-July. Dispersal power. Macropterous, probably capable of flight. Amara (Bradytus) schwarzi Hayward, 1908 Ecology. Moraines, cultivated fields (e.g., corn), vacant lots, sand pits, and gravel pits. Open ground; sandy soil covered with sparse or moderate vegetation. Nocturnal; sheltering during the day under dead leaves and pieces of wood. Biology. Seasonality: JuneAugust. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Raking the leaf litter. References. Lindroth, 1968 (natural history); Kirk, 1975c (ecology, dispersal power). Amara (Zezea) scitula Zimmermann, 1832 Ecology. Lowlands and mountains. Cultivated fields (e.g, wheat); borders of fields; vicinity of alkali lakes and ponds. Open ground; clay, often alkaline soil covered with some vegetation (e.g., grass, Spergularia, Triglochin). Biology. Seasonality: March-July, September-October, December. Tenerals: October (BC). Predators: blackbirds. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pitfall trapping. References. Lindroth, 1968 (natural history); Hieke, 1994 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Amara (Amara) sera Say, 1830 Natural History. Unknown. Reference. Hieke, 2000 (taxonomy). Amara (Amara) sinuosa (Casey, 1918) Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, vacant lots, cultivated fields (e.g., barley), roadsides, gravel pits, sand pits, and railway embankments. Ground open or shaded by bushes (e.g., Alnus); dry sandy or gravelly soil covered with sparse low vegetation. Mostly nocturnal; usually sheltering during the day under dead leaves, stones, pieces of wood and cardboard, and in the soil at the base of plants. Biology. Seasonality: June-September. Gravids: late September (AB). Tenerals: June-August;

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mostly early summer. Predators: toads (regularly). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Lindroth, 1955a, 1963a (as subaenescens Csiki), and 1968 (natural history); Frank, 1971b (biology); Kavanaugh, 1992 (natural history). Amara (Amara) sodalicia Casey, 1924 Natural History. Unknown. References. Hieke, 1997 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Amara (Curtonotus) spadicea (Casey, 1918), see Amara (Curtonotus) carinata (LeConte, 1848). Reference. Hieke, 1994 (taxonomy). Amara (Amara) specularis (Casey, 1918), see Amara (Amara) apachensis Casey, 1884. Reference. Hieke, 1994 (taxonomy). Amara (Curtonotus) sponsor (Casey, 1918), see Amara (Curtonotus) blanchardi Hayward, 1908. Reference. Hieke, 1994 (taxonomy). Amara (Amara) spuria Lindroth, 1968 Ecology. Vicinity of a river bank, with sandy soil covered with sparse vegetation (AK). Biology. Seasonality: June-September. Tenerals: mid-June and mid-August (AK). Dispersal power. Macropterous, probably capable of flight. Reference. Lindroth, 1968 (natural history). Amara (Curtonotus) tartarea (Casey, 1918), see Amara (Curtonotus) blanchardi (Hayward, 1908). Reference. Hieke, 1994 (taxonomy). Amara (Amara) tenebrionella (Bates, 1882) Ecology. Mountains. Biology and Dispersal power. Unknown. Amara (Amara) texana (Putzeys, 1866) Natural History. Unknown.

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Amara (Curtonotus) thoracica Hayward, 1908 Ecology. Lowlands and mountains. Grasslands. Open ground; sandy, dry soil covered with grass. Nocturnal; sheltering during the day under debris. Biology. Seasonality: May-September. Copulating pairs: mid-September (AB). Tenerals: early June (BC). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning debris. References. Lindroth, 1968 (natural history); Hieke, 1994 (taxonomy). Amara (Curtonotus) torrida (Panzer, 1797) Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, abandoned fields, meadows, cultivated fields (e.g., wheat, barley, faba bean, fescue), fencerows, roadsides, and upper zone of sea beaches. Ground open (mostly) or shaded by bushes (Alnus); rather dry soil covered with moderately dense vegetation (e.g., grass, weeds). Mostly nocturnal; usually sheltering during the day under stones, pieces of wood or cardboard, and carpet of dead weeds or dead algae. Gregarious. Biology. Seasonality: April-October. Copulating pairs: late June to July, rarely August. Gravids: late June to mid-August (AB, QU). Tenerals: late June to August. Adult food, in the field and in captivity: lepidopterous eggs. Parasites: dipterous larvae and nematodes. Dispersal power. Macropterous. Frequent in seashore drift material, which indicates flight activity. Moderate runner. Occasional climber (on plants). Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning debris; turning drift material; sweeping plants. References. Lindroth, 1945, 1955a, 1963a and 1968 (natural history); Frank, 1971b (biology); Forsskåhl, 1972 (biology); Lindroth, 1986 (natural history). Amara (Amara) turbata Casey, 1918 Ecology. Lowlands and uplands. Biology. Seasonality: April. Dispersal power. Macropterous, probably capable of flight. References. Hieke, 1994 and 2000 (taxonomy). Amara (Bradytus) vegasensis (Stehr, 1949), see Amara (Bradytus) novella (Stehr, 1949). Reference. Hieke, 1994 (taxonomy). Amara (Amara) volatilis (Casey, 1918) Ecology. Lowlands. Biology and Dispersal power. Unknown. Amblygnathus evansi Ball and Maddison, 1987 Ecology. Shallow Typha-marshes and palm forests, on damp black soil. Vicinity of marshy areas on drier, sandy places (Mexico). Biology. Seasonality: August (USA); January, April-

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September, December (Mexico). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting technique. Light trapping. Reference. Ball and Maddison, 1987 (natural history). Amblygnathus iripennis (Say, 1823) Ecology. Lowlands. Macrohabitat unknown. Nocturnal; sheltering during the day under pieces of wood and other debris. Biology. Seasonality: January-December. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting techniques. Turning debris; light trapping. Reference. Ball and Maddison, 1987 (biology, dispersal power). Amblygnathus mexicanus Bates, 1882 Ecology. Lowlands. A palm forest, with wet, black soil. Nocturnal; sheltering during the day under fallen palm fronds. Biology. Seasonality: January-September, November. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Reference. Ball and Maddison, 1987 (natural history). Amblygnathus subtinctus (LeConte, 1866) Ecology. Lowlands. Borders of marshes. Moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: February, June-July, September-November. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Treading the soil with the feet. Reference. Ball and Maddison, 1987 (biology, dispersal power). Amerinus linearis (LeConte, 1863) Ecology. Vicinity of swampy places. Previously flooded habitats. Wet, sandy soil. Mostly nocturnal; usually sheltering during the day among dead leaves. Biology. Seasonality: January-July, except May. Adults found overwintering in leaf litter. Dispersal power. Wing-dimorphic: macropterous (probably capable of flight) or brachypterous (incapable of flight). Occasional flier (during the day; to artificial lights at night). Moderate runner. Occasional climber (on plants). Collecting techniques. Raking the leaf litter; sweeping plants. References. Ulke, 1903 (ecology); Lindroth, 1968 (dispersal power). Ameroduvalius jeanneli jeanneli Valentine, 1952 Ecology. Caves: along pool margins. Moderately moist, muddy soil. Sheltering during the day under dead leaves and debris or running on the ground. Biology. Seasonality: July-November, except September. Adult food, in the field: worms (tubificids) and dipterous larvae. Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Ameroduvalius jeanneli rockcastlei Valentine, 1952 Ecology. Caves: along streams subjected to sudden flooding. Wet, silty soil. Biology. Seasonality: unknown. Adult food, in the field: small live organisms. Able to survive in water for over four hours (in captivity). Dispersal power. Brachypterous, incapable of flight. Fast runner. Amphasia (Amphasia) interstitialis (Say, 1823) Ecology. Lowlands and mountains. Deciduous flood-plain forests. Shaded ground; wet, clayish or muddy soil, rich in organic debris. Nocturnal; sheltering during the day under dead leaves, logs, and stones. Biology. Seasonality: February-December. Tenerals: April (rarely; MD), late July (WV), August (VT), late November (IN). Adults found overwintering in woods and at the edge of woods; under stones, in leaf litter, and in rotten logs. Adult food: vegetal matter (mostly) and insects (in the field); mealworms and diplopods (in captivity). Larval food, in captivity: mealworms. Predators: toads, bluebirds, flycatchers, robins, and thrushes. Dispersal power. Macropterous. Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on trees). Collecting techniques. Raking the leaf litter; pitfall trapping; pyrethrum spraying; beating trees. References. Blatchley, 1910 (ecology, biology); Lindroth, 1968 (natural history); Erwin, 1981 (natural history); Snider, 1984 (biology); Anderson et al. 1995 (ecology); Will et al. 1995 (dispersal power). Amphasia (Pseudamphasia) sericea (T.W. Harris, 1828) Ecology. Lowlands. Grasslands, meadows, pastures, cultivated fields (e.g., corn), vacant lots, sand pits, sand dunes, gravel pits, and orchards (e.g., apple). Open ground; dry, sandy soil covered with moderate or dense vegetation. Diurnal and nocturnal; during the day, active on grass heads and stems, burrowing under the roots of grasses and sedges, or sheltering in leaf litter. Biology. Seasonality: February-November. Tenerals: April (ON). Adults found overwintering on higher and drier ground, in vacant lots and on hillocks; under dead leaves. Adult food, in the field: grass seeds (Poa, Agrostis) and pollen, lepidopterous caterpillars, and other insects. Predators: crows. Dispersal power. Macropterous. Frequent flier (often swarming to artificial lights at night). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (mostly on plants; on trees). Strongly favored by human activities. Collecting techniques. Pitfall trapping; light trapping; raking the leaf litter; turning drift material; using tanglefoot screens; sweeping plants. References. Forbes, 1880 (biology); Webster, 1880 (biology); Blatchley, 1910 (ecology, dispersal power); Lindroth, 1968 (natural history); Kirk, 1969, 1970 (dispersal power) and 1975c (ecology, dispersal power); Will et al. 1995 (dispersal power). Anatrichis minuta (Dejean, 1831) Ecology. Lowlands and uplands. Swamp-forests, flood-plain forests, marshes; marshy edges of lakes, ponds, and slow rivers and brooks. Somewhat shaded ground; soft, wet,

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muddy or silty soil, rich in organic debris, covered with some vegetation. Nocturnal; sheltering during the day in the soil under vegetal debris (e.g., leaf litter). Biology. Seasonality: January-September, December. Tenerals: early July to mid-August. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet; raking the leaf litter; light trapping. References. Leng, 1915 (dispersal power); Frost, 1929b (ecology); Kirk, 1969 (dispersal power); Lee, 1978 (dispersal power); Bousquet, 1996a (taxonomy, ecology). Anatrichis oblonga G.H. Horn, 1891 Ecology. Lowlands. Swamp-forests, edges of marshes, and marshy borders of ponds. Nocturnal; sheltering during the day in the soil. Shaded or open ground; wet, muddy soil covered with some vegetation. Biology. Seasonality: December, April, July-August, October. Dispersal power. Macropterous, probably capable of flight. Strong runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Reference. Bousquet, 1996a (taxonomy). Anatrichis picea (Motchulsky, 1865), see Oodinus pseudopiceus Bousquet, 1996a. Reference. Bousquet, 1996a (taxonomy). Anchomenus aeneolus (LeConte, 1854) Ecology. Lowlands and mountains. River banks (mostly), lake shores, and reservoir margins. Close to water. Open ground; wet, stony-gravelly bare soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: June-September. Copulating pairs: late July (OR). Tenerals: mid-June to late July. Predators: robins. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Turning stones. References. Lindroth, 1966 (natural history); Liebherr, 1991a (ecology). Anchomenus funebris (LeConte, 1854) Ecology. Lowlands and mountains. Banks of large rivers. Open ground; wet, gravelly or stony soil, somewhat mixed with mud. Nocturnal; sheltering during the day. Biology. Seasonality: April-May, July, December. Eggs are laid on peat moss. Adults found overwintering. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Turning stones. Reference. Liebherr, 1991a (natural history). Anchomenus quadratus (LeConte, 1854) Ecology. Lowlands and mountains. Margins of brooks and rivers. Open ground; moist soil covered with some vegetation. Nocturnal; sheltering during the day under stones.

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Biology. Seasonality: June-August. Tenerals: early and late July (OR, WA). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Turning stones. References. Lindroth, 1966 (natural history); Liebherr, 1991a (ecology). Anchonoderus quadrinotatus G.H. Horn, 1878 Ecology. Borders of slow-running water courses. Wet sandy-muddy soil. Diurnal and nocturnal; during the day, running on the ground or sheltering under stones. Biology and Dispersal power. Unknown. Collecting technique. Turning stones. Reference. Ball and Bousquet, 2001 (ecology). Anchonoderus schaefferi Liebke, 1928 Ecology. Borders of slow-running water courses. Wet sandy-muddy soil. Diurnal and nocturnal; during the day, running on the ground or sheltering under stones. Biology and Dispersal power. Unknown. Collecting technique. Turning stones. Reference. Ball and Bousquet, 2001 (ecology). Anillaspis caseyi Jeannel, 1963 Ecology. Caves. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Anillaspis explanatus (G.H. Horn, 1888) Ecology. Caves. Biology. Seasonality: February. Dispersal power. Brachypterous, incapable of flight. Anillinus barberi Jeannel, 1963 Ecology. Forests: along river banks. Cool shaded ground; rather moist soil covered with thick leaf litter. Nocturnal; sheltering during the day under large large stones and in flood debris. Biology. Seasonality: March-May, October. Dispersal power. Brachypterous, incapable of flight. Slow runner. Collecting technique. Turning stones. Reference. Erwin, 1981 (natural history). Anillinus carolinae Casey, 1918, see Anilllinus fortis (G.H. Horn, 1868). Reference. Barr, 1995 (taxonomy). Anillinus dohrni (Ehlers, 1884) Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight.

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Anillinus dunavani Jeannel, 1963 Ecology. Mountains. Forests: along ravines. Shaded ground; moist soil covered with thick leaf litter. Nocturnal; sheltering during the day in leaf litter and under large stones. Biology. Seasonality: June, August. Dispersal power. Brachypterous, incapable of flight. Slow runner. Collecting techniques. Raking the leaf litter; turning stones. Reference. Kirk, 1970 (ecology). Anillinus elongatus Jeannel, 1963 Ecology. Lowlands. Biology. Seasonality: March-April. Dispersal power. Brachypterous, incapable of flight. Anillinus fortis (G.H. Horn, 1868) Ecology. Mountains. Forests (e.g., Picea, Abies). Shaded ground; rather moist soil covered with thick leaf litter. Nocturnal; sheltering during the day under embedded stones and in leaf litter. Associated with carrion. Biology. Seasonality: March-July, September-October. Defense-mechanism: when alarmed, the adult runs aimlessly instead of looking for a new shelter. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Slow runner. Collecting techniques. Turning stones; raking the leaf litter. References. Blatchley, 1910 (ecology, biology); Barr, 1969 (taxonomy, ecology); Kirk, 1970 (ecology; as carolinae Casey); Barr, 1995 (taxonomy). Anillinus indianae Jeannel, 1963 Ecology. Unknown. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Anillinus longiceps Jeannel, 1963 Ecology. Caves. Biology. Seasonality: March. Dispersal power. Brachypterous, incapable of flight. Anillinus sinuaticollis Jeannel, 1963 Ecology. Unknown. Biology. Seasonality: March. Dispersal power. Brachypterous, incapable of flight. Anillinus steevesi Barr, 1995 Ecology. Unknown. Biology. Seasonality: April, June. Dispersal power. Brachypterous, incapable of flight. Reference. Barr, 1995 (taxonomy).

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Anillinus turneri Jeannel, 1963 Ecology. Macrohabitat unknown. Nocturnal; sheltering during the day in the soil and in leaf litter. Biology. Seasonality: February, April, November-December. Dispersal power. Brachypterous, incapable of flight. Slow runner. Anillinus valentinei (Jeannel, 1963) Ecology. Caves. Sheltering under small wood debris. Biology. Seasonality: June, October. Dispersal power. Brachypterous, incapable of flight. References. Barr, 1969 (ecology) and 1995 (ecology, taxonomy). Anillinus virginiae Jeannel, 1963 Ecology. Mountains. Biology. Seasonality: September, November. Dispersal power. Brachypterous, incapable of flight. Anillodes affabilis (Brues, 1902) Ecology. Ant nests. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Slow runner. Anillodes debilis (LeConte, 1853) Ecology. Mountains. Forests. Shaded ground; rather moist soil. Nocturnal; sheltering during the day under stones and dead leaves. Biology. Seasonality: November. Dispersal power. Brachypterous, incapable of flight. Slow runner. Collecting techniques. Turning stones; raking the leaf litter. Anillodes minutus Jeannel, 1963 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Anillodes sinuatus Jeannel, 1963 Ecology. An orchard (peach). Nocturnal; sheltering during the day in the soil. Biology. Seasonality: February, May, November. Dispersal power. Brachypterous, incapable of flight. Anillodes walkeri Jeannel, 1963 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Anisodactylus (Anisodactylus) agricola (Say, 1823) Ecology. Lowlands. Flood-plain forests (e.g., Gleditsia, Platanus). Shaded ground; wet soil

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composed of sandy loam or sand. Nocturnal; sheltering during the day under stones (mostly) and in leaf litter. Biology. Seasonality: February-November. Tenerals: March-May, August, October; mostly March-April. Predators: bluebirds, cardinals, and crows. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones; night collecting with a headlamp. References. Erwin, 1981 (natural history); Will et al. 1995 (dispersal power); Noonan, 1996 (natural history). Anisodactylus (Anadaptus) alternans LeConte, 1851 (Anisodactylus (Anadaptus) nivalis G.H. Horn, 1880) Ecology. Lowlands and mountains. Borders of eutrophic marshes, lakes, and reservoirs; montane meadows. Open ground; soft, wet, muddy soil, sometimes sand-mixed, covered with some vegetation (e.g., Juncus, Typha). Nocturnal; sheltering during the day under fallen plants, pieces of wood, fallen trees, dry cow dung, stones, and pieces of cracked clay. Biology. Seasonality: February-April, July. Tenerals: late July (OR). Adult food, in the field and in captivity: grasshopper nymphs. Dispersal power. Macropterous. Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pitfall trapping; turning debris. References. Hatch, 1953 (ecology; as nivalis G.H. Horn); Noonan, 2001 (taxonomy, natural history; Harpalus alternans Motschulsky, 1845 is treated as a Incertae Sedis case) Anisodactylus (Anadaptus) alternans (Motschulsky, 1845), see Anisodactylus (Anadaptus) alternans LeConte, 1851. Reference. Noonan, 2001 (taxonomy) Anisodactylus (Aplocentroides) amaroides LeConte, 1851 Ecology. Mountains. Biology. Seasonality: May, July. Dispersal power. Macropterous, probably capable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Anisodactylus (Gynandrotarsus) anthracinus (Dejean, 1829) Ecology. Mountains. Meadows, grasslands, pastures, vacant lots, roadsides, deserts, and open forests (e.g., Quercus, Pinus). Open ground; moist soil composed of clay or sand, covered with dense vegetation. Nocturnal; sheltering during the day under stones, dead leaves, dry cow dung, fallen tree bark, and logs. Biology. Seasonality: January, May-November. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Collecting techniques. Pitfall trapping; turning debris. Anisodactylus (Anisodactylus) binotatus (Fabricius, 1787) Ecology. Lowlands. Vicinity of cities. Cultivated fields, lawns, dumps, peat bogs; upper banks of lakes and reservoirs. Greenhouses (occasionally). Often near water. Open ground; moderately moist, composed of clay, clay-mixed sand, or loam, with dense

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vegetation (e.g., grass, sedges). Nocturnal; sheltering during the day under stones. Biology. Seasonality: March-November. Tenerals: June-October; mostly August. Adult food, in the field (Europe): worms, mollusks, and strawberries; the larva is carnivorous. Dispersal power. Macropterous. Flight records (Europe); flying in spring (mostly) and in late summer. Frequent in lakeshore drift material, which indicates flight ability. Moderate runner. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; turning drift material. References. Lindroth, 1945 and 1968 (natural history); Sharova, 1981 (ecology, biology); Lindroth, 1986 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology); Noonan, 1996 (natural history). Anisodactylus (Aplocentroides) caenus (Say, 1823) Ecology. Lowlands and mountains. Low deciduous forests. Shaded ground; moist or wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: January, MarchAugust, October-November. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; light trapping. References. Allen and Thompson, 1977 (ecology); Will et al. 1995 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Anisodactylus (Anisodactylus) californicus Dejean, 1829 Ecology. Lowlands and mountains. Borders of eutrophic marshes and ponds; borders of lakes, reservoirs, and slow rivers; cultivated fields (e.g., strawberry) and orchards (e.g., apple). Greenhouses (often). Open ground; soft, moist or wet soil composed of mud, clay or loam, covered with moderate or dense vegetation (e.g., grass and herbaceous plants). Nocturnal; sheltering during the day under stones, pieces of wood, plant mats, and in the soil. Biology. Seasonality: January-December. Tenerals: MayDecember. Adult food: strawberries and grasshopper nymphs (in the field), lepidopterous caterpillars and rolled oats (in captivity). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning debris. References. Lindroth, 1968 (natural history); Noonan, 1996 (taxonomy, natural history). Anisodactylus (Anisodactylus) carbonarius (Say, 1823) Ecology. Lowlands and mountains. Grasslands, meadows, pastures, cultivated fields (e.g., corn, alfalfa, asparagus, barley), abandoned fields, vacant lots, roadsides, and orchards (e.g, peach). Open ground; moderately moist soil, often consisting of sandy loam or clay, covered with moderate or dense vegetation. Nocturnal; sheltering during the day in the soil under dead grass and other debris. Biology. Seasonality: February-December. Tenerals: April-August; mostly June-August. Adult food, in captivity: lepidopterous caterpillars. Predators: crows, robins, and starlings. Parasites: mites. Defense-mechanism: when alarmed, the adult may emit a repugnant Brachinus-like smoke from the pygidial glands. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in seashore drift

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material, which indicates flight ability. Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; digging under debris. References. Kirk, 1969 (dispersal power) and 1975c (ecology); Erwin, 1981 (natural history). Anisodactylus (Anisodactylus) consobrinus LeConte, 1851 Ecology. Lowlands and mountains. River and brook banks; lake shores and pond edges. Open ground; moist, muddy soil covered with some grass and other herbaceous plants. Nocturnal; sheltering during the day under stones, fallen tree bark, and other debris. Biology. Seasonality: February-December. Tenerals: March, June-October; mostly July-August. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Turning stones; pitfall trapping. Reference. Noonan, 1996 (natural history). Anisodactylus (Anadaptus) depressicollis (Motschulsky, 1859), see Anisodactylus (Anisodactylus) californicus Dejean, 1829. Reference. Noonan, 1996 (taxonomy). Anisodactylus (Anadaptus) discoideus Dejean, 1831 Ecology. Lowlands. Borders of rivers and brooks; ponds and pools in sand pits and gravel pits; alluvial places, cultivated fields (e.g., corn), and golf courses. Open ground; soft, wet soil composed of fine sand (somewhat silt-mixed), bare or covered with sparse vegetation. Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering during the day in burrows dug in the soil (mostly), under well-embedded logs, stones, pieces of wood, and dead leaves. Gregarious. Biology. Seasonality: February, April-November. Copulating pairs: early June (QU). Gravids: early June and mid-July (QU). Tenerals: August-September (QU). Adults found overwintering on higher and drier ground, in the upper zone of river banks, sand pits and gravel pits, on the top of hillocks and mounds; in burrows dug in the soil (mostly; as deep as 15 cm) and under stones. Able to survive winter inundations. Adult food, in the field: vegetal matter, insects, and other animal matter. Adult and larval food, in captivity: mealworms. Predators: frogs, toads, flycatchers, robins, thrushes, woodpeckers, and raccoons. Defense-mechanism: when alarmed, the adult may escape by digging into the soil; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (in the sunshine; to artificial lights at night; to artifical lights in captivity). Slow runner. Strong burrower. Collecting techniques. Pitfall trapping; light trapping; turning embedded logs and stones. References. Blatchley, 1910 (ecology); Dogger and Olson, 1966 (biology); Lindroth, 1968 (natural history); Larochelle, 1974a (ecology); Kirk, 1975c (ecology, dispersal power); Erwin, 1981 (natural history); Noonan, 2001 (natural history). Anisodactylus (Gynandrotarsus) dulcicollis (LaFerté-Sénectère, 1841) Ecology. Lowlands and mountains. Pastures, cultivated fields (e.g., tobacco, strawberry, sweet potato, cotton, soybean, and rice), vacant lots, gardens, orchards (e.g., peach),

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and open forests. Open ground; sandy soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day under stones, dead plants, dry cow dung, at the base of plants, under the loose bark of rotten logs, and in burrows dug in the soil. Biology. Seasonality: January-December. Tenerals: February (AL). Adults found overwintering in vacant lots and at the edge of woods; in burrows dug in the soil, under dead leaves, and under the loose bark of fallen trees. Predators: ducks. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Fast runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; digging the soil; raking the leaf litter; light trapping. Anisodactylus (Anisodactylus) furvus LeConte, 1863 Ecology. Lowlands and mountains. Cultivated fields (e.g., alfalfa, cotton, soybean), grasslands, pastures, pecan fields, orchards (e.g., peach), blowouts, upper zone of brook borders. Open ground; sandy soil covered with some vegetation. Nocturnal; sheltering during the day under dead leaves, stones, logs, and fallen tree bark. Biology. Seasonality: FebruaryDecember. Tenerals: June-July. Adults found overwintering (SC). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants, e.g., Lespedeza; on shrubs). Collecting techniques. Pitfall trapping; turning debris; sweeping plants; beating shrubs. References. Kirk, 1970 (ecology); Noonan, 1996 (natural history). Anisodactylus (Gynandrotarsus) haplomus Chaudoir, 1868 Ecology. Lowlands. A coniferous forest (Pinus), on well-drained, sandy soil. Nocturnal; sheltering during the day. Biology. Seasonality: March-August, November-December. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Fast runner. Collecting techniques. Pitfall trapping; light trapping. Anisodactylus (Gynandrotarsus) harpaloides (LaFerté-Sénectère, 1841) Ecology. Lowlands. Pastures and cultivated fields (e.g., cotton, soybeans). Open ground; well-drained soil covered with sparse vegetation. Nocturnal; sheltering during the day under dry cow dung and stones. Biology. Seasonality: January, March-September. Tenerals: September (TN). Adult food, in the field: heteropterans. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in beach drift material, which indicates indirect flight. Fast runner. Collecting techniques. Pitfall trapping; turning debris. Anisodactylus (Anisodactylus) harrisii LeConte, 1863 Ecology. Lowlands and mountains. Borders of marshes, ponds, lakes, rivers, brooks, and irrigation canals; swamps. Moist pastures and meadows, relict prairies, cultivated fields (e.g, alfalfa, corn, oat), vacant lots, gardens, fencerows, gravel pits, rubbish dumps, and orchards (e.g., apple). Frequently near water. Open ground; moderately moist, often clay-

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ish or muddy soil covered with meadow-type vegetation. Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering during the day under stones, pieces of wood, dead leaves, and mats of dead plants. Gregarious (in winter). Biology. Seasonality: March-December. Copulating pairs: April-May (QU); coitus lasting 18-35 minutes. Gravids: early May to mid-June (QU). Tenerals: April-June (rarely), July-November; mostly August-September. Adults found overwintering on higher and drier ground, in vacant lots, in the upper zone of sand pits, gravel pits, and irrigation canals, on sandy hillocks and hills, in field hedges, at the edge of woods, and in woods; in burrows dug in the soil (mostly; as deep as 15 cm) dug under stones, also under dead leaves and plants, and plant rosettes (Verbascum). Adult food: plant seeds (in the field); lepidopterous caterpillars, earthworms, mealworms, and meat (in captivity). Larval food, in captivity: mealworms. Predators: frogs, toads, pheasants, and thrushes. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Frequent flier (to artificial lights at night; to artifical lights in captivity). Moderate runner. Frequent in seashore drift material, which indicates flight ability. Occasional climber (on plants). Favored by spring floods along large rivers. Collecting techniques. Pitfall trapping; light trapping; turning stones; turning drift material; sweeping plants. References. Lindroth, 1968 (natural history); Kirk, 1975c (ecology); Will et al. 1995 (dispersal power); Noonan, 1996 (natural history). Anisodactylus (Anisodactylus) kirbyi Lindroth, 1953 Ecology. Lowlands. Borders of eutrophic ponds, marshes, and canalised brooks; swamps, flood-plain forests, and wet meadows. Open ground or in the shade of trees; wet, usually sand-mixed soil, rich in organic debris, covered with dense depressed vegetation (e.g., grass, Carex). Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering durring the day under mats of dead plants, dead leaves, logs, stones, and pieces of fallen tree bark. Biology. Seasonality: February-December. Copulating pairs: May-June (QU). Gravids: early May to early June (QU). Tenerals: March-October; mostly after mid-summer. Adults found overwintering on higher and drier ground, in the upper zone of sand dunes, sand pits, gravel pits, and hills, in vacant lots and thickets, at the edge of woods, and in woods; under stones, in burrows dug in the soil (as deep as 15 cm), under dead leaves, logs, pieces of fallen bark, the loose bark of fallen tree-stumps, and in rotten logs. Adult and larval food, in captivity: mealworms. Predators: frogs and toads. Phoretic mites often infesting the adult body. Defense-mechanism: when alarmed, the adult may escape by diving into the water. Dispersal power. Macropterous. Frequent flier (in the sunshine; to artificial lights at night; to artifical lights in captivity). Occasional in sea- and lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Collecting techniques. Treading the vegetation into the water; light trapping; sweeping plants. References. Lindroth, 1968 (natural history); Noonan, 1996 (natural history). Anisodactylus (Pseudaplocentrus) laetus Dejean, 1829 Ecology. Lowlands. Borders of ponds and lakes. Golf courses (occasionally). Open ground; wet, muddy soil, rich in organic debris. Nocturnal; sheltering during the day

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under dead leaves and other dead organic debris (e.g., dead crayfish). Biology. Seasonality: January-October. Copulating pairs: late July (AL). Adults found overwintering in wood debris. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting techniques. Light trapping; turning debris; using Japanese beetle traps; using interception traps. References. Leng, 1915 (ecology, dispersal power); Kirk, 1969 (ecology, biology), 1970, and 1975c (dispersal power). Anisodactylus (Anisodactylus) lodingi Schaeffer, 1911 Ecology. Lowlands. Wet situations. Nocturnal; sheltering during the day under debris and cow dung. Biology. Seasonality: February-June, September-November. Adults found overwintering (AL). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Good swimmer. References. Leng, 1915 (ecology); Löding, 1945 (ecology); Noonan, 1996 (species possibly extinct). Anisodactylus (Anisodactylus) melanopus (Haldeman, 1843) Ecology. Lowlands. Borders of eutrophic marshes, ponds, and lakes; swamps; swampforests; marshy edges of slow brooks and rivers. Open or somewhat shaded ground; wet, muddy soil, rich in organic debris, covered with dense vegetation. Nocturnal; sheltering during the day under dead leaves, plant mats, logs, and boards. Biology. Seasonality: February-December. Tenerals: June-September. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Treading the vegetation into the water; turning debris. References. Kirk, 1969 (dispersal power) and 1975c (ecology); Erwin, 1981 (natural history); Noonan, 1996 (natural history). Anisodactylus (Gynandrotarsus) merula (Germar, 1824) Ecology. Lowlands and mountains. Moraines, cultivated fields (e.g., alfalfa, soybean, cotton, corn, strawberry), pastures, abandoned fields, vacant lots, roadsides, gardens, sand hills, sand pits, gravel pits, orchards (e.g., peach), forest clearings, and open forests. Open ground; very dry, sandy soil covered with sparse vegetation. Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering during the day under dead leaves, plant rosettes, stones, reindeer moss (Cladonia), pieces of cardboard, dry cow dung, logs, and in burrows dug in the soil. Gregarious. Biology. Seasonality: March-October. Gravids: early July (QU). Tenerals: April (rarely), July-August, October. Adults found overwintering in vacant lots, pastures, hills, and gravel pits; under plant rosettes (Verbascum), stones, in the soil at the base of plants (Gramineae), and under heaps of dead grass. Adult food: plant seeds and weevils (in the field); mealworms and earthworms (in captivity). Larval food, in captivity: mealworms. Predators: toads (frequently). Dispersal power. Macropterous. Frequent flier (to artificial lights at night; to artifical lights in captivity). Fast runner. Occasional climber (on plants). Strongly favored by human activities. Collecting techniques. Pitfall trapping;

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light trapping; turning debris; sweeping plants. References. Lindroth, 1968 (natural history); Kirk, 1975c (ecology). Anisodactylus (Anisodactylus) nigerrimus (Dejean, 1831) Ecology. Lowlands and mountains. Vacant lots, sandhills, pastures, cultivated fields (e.g., carrot), old fields, sand pits, gravel pits, roadsides, forest edges and clearings. Open ground; firm, very dry, sandy soil covered with sparse vegetation (e.g., grass, herbs). Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering during the day under stones, dead leaves, heaps of hay and weeds, reindeer moss (Cladonia), and pieces of wood and cardboard. Biology. Seasonality: January-December. Gravids: mid-June (QU). Tenerals: January-November; mostly after mid-July. Adults found overwintering in woods, at the edge of woods, on hills, in vacant lots and gravel pits; under stones, dead leaves, reindeer moss (Cladonia), pieces of wood, and in moss. Adult and larval food, in captivity: mealworms. Predators: toads. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Frequent flier (in the sunshine; at sunset). Frequent in seashore drift material, which indicates flight ability. Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; turning drift material. References. Lindroth, 1963a (natural history); Kirk, 1969 (biology); Larochelle, 1974a (ecology); Erwin, 1981 (natural history); Anderson et al. 1995 (ecology); Noonan, 1996 (natural history). Anisodactylus (Anisodactylus) nigrita Dejean, 1829 Ecology. Lowlands and uplands. Borders of eutrophic marshes, ponds, and lakes; borders of rivers, irrigation canals, and roadside ditches; swamps, swamp-forests; beaver houses, pools in gravel pits and sand pits; vacant lots. Frequently near water. Open ground; wet soil usually consisiting of sand-mixed clay, covered with dense depressed vegetation carpet (e.g., grass, Carex). Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering during the day under mats of plant debris, pieces of wood, dead leaves, stones, and in the soil. Gregarious (in winter). Biology. Seasonality: February-December. Copulating pairs: April (ON), mid-May (VA). Gravids: April (QU). Tenerals: May-October; mostly after mid-July. Adults found overwintering on higher and drier ground, in sand dunes, gravel pits, vacant lots, and meadows, at the edge of woods, and in woods; under stones (mostly), in the soil (as deep as 15 cm), in rotten logs, and under the loose bark of logs and fallen trees. Adult and larval food, in captivity: mealworms. Predators: toads, bluebirds, and woodpeckers. Dispersal power. Macropterous. Frequent flier (in the sunshine; to artificial lights in captivity). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants, e.g., Carex). Collecting techniques. Treading the vegetation into the water; turning mats of plant debris; light trapping; sweeping plants. References. Lindroth, 1968 (natural history); Kirk, 1975c (ecology, biology); Noonan, 1996 (natural history).

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Anisodactylus (Anadaptus) nivalis G.H. Horn, 1880, see Anisodactylus (Anadaptus) alternans LeConte, 1851. Reference. Noonan, 2001 (taxonomy). Anisodactylus (Gynandrotarsus) opaculus (LeConte, 1863) Ecology. Lowlands and uplands. Grasslands, cultivated fields (e.g., alfalfa), pastures, orchards (e.g., apple), mesquite scrub-deserts, and open forests. Nocturnal; sheltering during the day in the soil under the leaf litter, stones and dry cow dung. Biology. Seasonality: January-August, November. Adult food, in the field: seeds and other vegetal tissue. Predators: bluebirds. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Fast runner. Occasional climber (on trees). Collecting techniques. Pitfall trapping; light trapping; using tanglefoot screens; beating trees. Anisodactylus (Gynandrotarsus) ovularis (Casey, 1914) Ecology. Lowlands and mountains. Pastures, grasslands, meadows, rangelands, cultivated fields (e.g., tobacco, corn, alfalfa, soybean), old fields, edges of fields, fencerows, lawns, orchards, and open forests. Open ground; sandy soil covered with some vegetation (e.g., grass, herbs). Nocturnal; sheltering during the day in the soil at the base of plants, under dry cow dung, stones, and pieces of wood. Biology. Seasonality: January, April-December. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Frequent in lake- and seashore drift material, which indicates flight ability. Fast runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; digging at the base of plants; light trapping; turning drift material. References. Kirk, 1975c (ecology); Anderson et al. 1995 (ecology). Anisodactylus (Anadaptus) pitychrous LeConte, 1861 Ecology. Lowlands, uplands, and subalpine zone. Sagebrush areas, meadows, grassy clearings, roadside ditches, and open forests. Open ground; moist or wet soil covered with grass or herbaceous plants. Nocturnal; sheltering during the day under stones, dry cow dung, and pieces of wood. Biology. Seasonality: July. Predators: asilids (Diptera). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Noonan, 2001 (taxonomy, natural history). Anisodactylus (Anadaptus) porosus (Motschulsky, 1845) (Anisodactylus viridescens LeConte, 1861) Ecology. Mountains. Roadside ditches and meadows. Open ground; moist or wet soil covered with grass or herbaceous plants. Nocturnal; sheltering during the day under stones, died cow dung, dead plants, and pieces of wood. Biology. Seasonality: MayJune, August. Dispersal power. Macropterous, probably capable of flight. Moderate

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runner. Collecting technique. Pitfall trapping. References. Bousquet, 1997a (taxonomy); Noonan, 2001 (taxonomy, natural history). Anisodactylus (Anisodactylus) pseudagricola Noonan, 1996 Ecology. Lowlands. Macrohabitat unknown. Taken by treading a Sphagnum-mat. Biology. Seasonality: April-June, September. Dispersal power. Wing-condition unknown. Moderate runner. Reference. Noonan, 1996 (taxonomy, ecology). Anisodactylus (Anadaptus) rudis LeConte, 1863 Ecology. Lowlands and mountains. Meadows, grassy clearings, and roadsides. Open ground; moist or wet soil covered with grass or herbaceous plants. Nocturnal; hiding during the day under stones, dry cow dung, and pieces of wood. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Occasional climber (on plants). Collecting techniques. Pitfall trapping; sweeping plants. Reference. Noonan, 2001 (taxonomy, natural history). Anisodactylus (Gynandrotarsus) rusticus (Say, 1823) Ecology. Lowlands, mountains, subalpine and alpine zones. Vacant lots, pastures, grasslands, meadows, rangelands, ploughed fields, cultivated fields and their borders (e.g., corn, hay, oat, wheat, cotton, sweet clover, alfalfa, soybean, asparagus, cabbage, tobacco, strawberry, peanut), old fields, golf courses, lawns, sandhills, sand pits, gravel pits, roadsides, orchards (e.g., peach, plum), pine plantation, and open forests. Open ground; dry sandy soil covered with thin but frequently tall vegetation (e.g., grass). Mostly nocturnal; active in the spring sunshine (e.g., over plowed ground); usually sheltering during the day in the soil (mostly) at the base of plants, under dead leaves, heaps of hay, dead plants, stones, reindeer moss (Cladonia), in clumps of broomsedges, under pieces of wood, and clods of earth. Gregarious. Biology. Seasonality: January-December. Gravids: mostly JuneJuly (ON). Tenerals: April-October; mostly from mid-July. Adults found overwintering on higher and drier ground, on hills and hillocks, in vacant lots, sand pits, gravel pits, meadows, and roadsides; under stones, plant rosettes (Verbascum), dead leaves, logs, and in the soil at the base of plants. Adult food: vegetal matter (e.g., fungi), lepidopterous caterpillars and other insects (in the field); grass seeds, corn kernels, mealworms, earthworms, and slugs (in captivity). Larval food, in captivity: mealworms. Predators: frogs, toads (frequently), bluebirds, quails, cardinals, prairie chickens, crows, woodpeckers, flycatchers, starlings, and ducks. Parasites: tapeworms. Dispersal power. Macropterous. Frequent flier (in the sunshine; to artificial lights at night). Frequent in lake- and seashore drift material, which indicates flight activity. Fast runner. Occasional climber (on trees). Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones and plant rosettes; light trapping; turning drift material; beating trees. References. Blatchley, 1910 (ecology, dispersal power); Herne, 1963 (biology); Rivard, 1964c (biology); Lindroth, 1968 (natural history); Kirk, 1969 (biology, dispersal power) and 1975c (ecology); Erwin, 1981 (natural history); Anderson et al. 1995 (ecology).

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Anisodactylus (Anadaptus) sanctaecrucis (Fabricius, 1798) Ecology. Lowlands and mountains. Borders of rivers, brooks, lakes, ponds, reservoirs, pools, marshes, and alluvial places. Sand pits, gravel pits, roadside ditches, vacant lots, ski fields, pastures, meadows, grasslands, cultivated fields (e.g., corn, wheat, alfalfa, soybean, tobacco, barley, oat, potato, carrot, asparagus, cabbage) and their borders, orchards (e.g., apples, peach), and open forests. Open ground; soft, wet or moist, often sandy soil covered with dense vegetation (e.g., grass, etc.). Mostly nocturnal; quite active in the spring sunshine; usually sheltering during the day in burrows dug in the soil, under dead leaves, stones, mats of dead plants, logs, and pieces of wood. Gregarious. Biology. Seasonality: January, March-December. Copulating pairs: May (QU). Gravids: May to early July (SD). Gravids: mid-May to late June (IL, QU, VT). Tenerals: late May-October; mostly from mid-summer on. Adults found overwintering on higher and drier ground, in woods and at the edge of woods (mostly), in the upper zone of sand pits, gravel pits, sand dunes, and roadsides; in leaf litter, in the soil (as deep as 7.5 cm), under logs, pieces of fallen bark, the loose bark of fallen trees, and in rotten logs. Adult food: vegetal matter (e.g., grass) and lepidopterous caterpillars (in the field); various plant seeds, lepidopterous caterpillars, dipterous pupae, immature stages of weevils, scarabaeid larvae, earthworms, mealworms, dog food, and chicken eggs (in captivity). Larval food, in captivity: mealworms, dog food, and chicken eggs. Predators: frogs, toads (regularly), bluebirds, quails, catbirds, cowbirds, flycatchers, robins, starlings, and swallows. Parasites: Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult escapes by flying readily; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (in the sunshine; to artificial lights at night; to artifical lights in captivity). Frequent in lake- and seashore drift material, which indicates flight ability. Slow runner. Occasional climber (on plants and trees). Strongly favored by human activities. Collecting techniques. Pitfall trapping; light trapping; treading the soil with the feet; turning drift material; using emergence traps; sweeping plants. References. Herne, 1963 (biology); Lindroth, 1968 (natural history); Kirk, 1975c (ecology, dispersal power) and 1977 (biology); Hsin et al. 1979 (biology); Boivin and Hance, 1994 (biology); Chen and Willson, 1996 (biology); Noonan, 2001 (natural history). Anisodactylus (Anisodactylus) similis LeConte, 1851 Ecology. Lowlands and mountains. Meadows, cultivated fields (e.g., pea), clearings, hills, coastal canyons, orchards (e.g., apple), and open forests. Greenhouses (occasionally). Open ground; moderately moist soil covered with dense vegetation (e.g., grass and herbaceous plants). Nocturnal; sheltering during the day under pieces of wood, fallen bark and tree trunks, dry cow dung, and stones. Biology. Seasonality: January-December. Tenerals: April-September; mostly June-July. Predators: flycatchers. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning debris. Reference. Noonan, 1996 (natural history).

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Anisodactylus (Gynandrotarsus) texanus Schaeffer, 1910 Ecology. Lowlands. Gravel pits, roadside ditches; vicinity or water bodies in arid areas, scrub-deserts, and open forests. Open or shaded ground; dry or moist, gravelly or clay soil covered with sparse vegetation. Nocturnal; sheltering during the day under stones, dead leaves, plant heaps, and fallen tree branches. Biology. Seasonality: February-July. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Collecting techniques. Pitfall trapping; turning debris. Anisodactylus (Spongopus) verticalis (LeConte, 1848) Ecology. Lowlands and mountains. Deciduous flood-plain forests. Shaded ground; moist or wet, clayish or muddy soil, rich in organic debris. Nocturnal; sheltering during the day under logs, fallen trees, stones, the loose bark of trees, under pieces of fallen bark, and under dead leaves. Biology. Seasonality: March-September. Copulating pairs: early June (QU). Tenerals: late May-September; mostly from mid-summer on. Adults found overwintering on higher and drier ground, in woods; under stones. Adult and larval food, in captivity: mealworms. Predators: toads, crows, and raccoons. Parasites: mites. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pitfall trapping; turning logs and fallen trees. References. Blatchley, 1910 (ecology); Gilbert, 1957 (biology); Lindroth, 1968 (natural history); Larochelle, 1974a (ecology); Kirk, 1975c (ecology); Erwin, 1981 (natural history). Anisodactylus (Anadaptus) viridescens LeConte, 1861, see Anisodactylus (Anadaptus) porosus (Motschulsky, 1845). References. Noonan, 1996 (taxonomy); Bousquet, 1997a (taxonomy). Apenes (Apenes) angustatus Schwarz, 1878 Ecology. Lowlands. Biology. Seasonality: June-August. Dispersal power. Macropterous, probably capable of flight. Apenes (Apenes) lucidulus (Dejean, 1831) Ecology. Lowlands and mountains. Deciduous forests (e.g., Quercus, Carya). Shaded ground; soil rather moist, covered with thick leaf litter. Nocturnal; sheltering during the day among dead leaves and under stones. Biology. Seasonality: March-September. Copulating pairs: April (CT). Adults found overwintering in woods; in leaf litter. Dispersal power. Wing-dimorphic: macropterous (mostly; capable of flight) or brachypterous (incapable of flight). Frequent flier (to artificial lights at night). Frequent in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter; light trapping; turning drift material; using berlese funnels. References. Erwin, 1981 (natural history); Lenski, 1981 (dispersal power).

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Apenes (Apenes) nebulosus LeConte, 1866 Ecology. Unknown. Biology. Seasonality: June-August. Dispersal power. Macropterous, probably capable of flight. Apenes (Apenes) opacus LeConte, 1851 Ecology. Lowlands. Coniferous forests (e.g., Pinus). Shaded ground; rather moist, sandy soil. Nocturnal; sheltering during the day near the base of trees and tree-stumps, under pieces of wood, clumps of grass, in leaf litter, and under stones. Biology. Seasonality: January-April, June-July. Adults found overwintering in wood debris. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning debris; sifting clumps of grass roots. References. Leng, 1915 (ecology); Kirk, 1969 (biology) and 1970 (ecology). Apenes (Apenes) pallidipes (Chevrolat, 1835) Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Apenes (Apenes) sinuatus (Say, 1823) Ecology. Lowlands and mountains. Deciduous (e.g., Quercus, Carya) or coniferous forests (e.g., Pinus); open woodlands. Shaded ground; occurring at the base of trees and tree-stumps; soil rather moist, covered with thick leaf litter. Nocturnal; sheltering during the day under dead leaves, logs, fallen tree bark, and other debris. Biology. Seasonality: January-December. Adults found overwintering in woods; in leaf litter, under pieces of wood, and in tree hollows. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pitfall trapping; light trapping; raking the leaf litter. References. Blatchley, 1910 (ecology); Kirk, 1969 (natural history) and 1970 (ecology); Lindroth, 1969a (dispersal, power). Apristus actuosus Casey, 1920 Ecology. Lowlands. Biology. Seasonality: May. Dispersal power. Macropterous, probably capable of flight. Apristus agitatus Casey, 1920 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Apristus cephalus Casey, 1920 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight.

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Apristus constrictus Casey, 1920 Ecology. Lowlands and mountains. Vicinity of rivers (mostly), brooks, and lakes. Close to water. Open ground; well-drained, dry, sandy or gravelly bare soil. Diurnal; active in the sunshine; sheltering on cloudy or cool days among gravel or under stones. Biology. Seasonality: May-July. Tenerals: mid-July (WA). Defense-mechanism: when alarmed, the adult escapes by running and flying readily; the beetle is difficult to pursue; body color matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed). Fast runner. Collecting techniques. Hunting with a butterfly net; raking the loose gravel. Reference. Lindroth, 1969a (ecology). Apristus latens (LeConte, 1848) Ecology. Lowlands and mountains. Vicinity of rivers, brooks, and lakes. Close to water or at some distance from it. Open ground; well-drained, dry, sandy or gravelly bare soil. Diurnal; active in the sunshine; sheltering on cloudy or cool days among gravel, under stones, pieces of wood or cardboard, and at the base of plants. Biology. Seasonality: May-September. Tenerals: early August (MN, TX). Defense-mechanism: when alarmed, the adult escapes by running and flying readily; the beetle is difficult to pursue; body color matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed). Fast runner. Collecting techniques. Hunting with a butterfly net; raking the loose gravel. Reference. Bell, 1971 (ecology). Apristus laticollis LeConte, 1851 Ecology. Lowlands and mountains. Vicinity of rivers, brooks, ditches, and lakes; garden walks. Open ground; dry, bare soil. Diurnal; active in the sunshine. Biology. Seasonality: April-August. Defense-mechanism: when alarmed, the adult escapes by running and flying readily; the beetle is difficult to pursue. Dispersal power. Macropterous. Frequent flier (when alarmed). Fast runner. Collecting techniques. Hunting with a butterfly net; raking the soil. Apristus liratus Casey, 1920 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Apristus nevadensis Casey, 1920 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Apristus pugetanus Casey, 1920 Ecology. Lowlands and mountains. River banks and lake shores. Open ground; welldrained, dry, sandy or gravelly bare soil. Mostly diurnal; active in the sunshine; sheltering

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on cloudy or cool days under stones. Biology. Seasonality: May-August. Tenerals: late August (UT). Defense-mechanism: when alarmed, the adult escapes by running and flying readily; the beetle is difficult to pursue; body color matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed). Fast runner. Collecting techniques. Collecting with a butterfly; raking the soil. Reference. Lindroth, 1969a (ecology). Apristus subdeletus Casey, 1920 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Fast runner. Apristus subsulcatus (Dejean, 1826) Ecology. Lowlands. Vicinity of rivers (mostly), brooks, and ponds. Gravel pits, sand pits, abandoned fields, cultivated fields (e.g., soybean), pastures, and roadsides. Open ground; well-drained, dry, sandy or gravelly bare soil. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days among gravel, under stones, pieces of wood, and other debris. Biology. Seasonality: January-October. Tenerals: April (rarely), late August, and September (QU). Adults found overwintering on higher and drier ground, in the upper zone of river banks; in leaf litter. Defense-mechanism: when alarmed, the adult escapes by running and flying readily; the beetle is difficult to pursue; body color matching the soil. Dispersal power. Macropterous. Frequent flier (in the bright sunshine). Fast runner. Collecting techniques. Hunting with a butterfly net; raking the soil; using an aspirator. References. Lindroth, 1969a (natural history); Erwin, 1981 (natural history); Will et al. 1995 (ecology). Apristus thoracicus Casey, 1920 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Apristus tuckeri Casey, 1920 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Ardistomis morio (Dejean, 1831) Ecology. Lowlands. Biology. Seasonality: March-April. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Ardistomis obliquatus Putzeys, 1846 Ecology. Lowlands. Banks of river and brooks, pond margins, seashore, swamps, swampforests, and wet meadows. Open ground; wet soil composed of sand or humus, sometimes saline, covered with some vegetation. Nocturnal; sheltering during the day in burrows dug in the soil and under grass or seaweeds. Biology. Seasonality: January-

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September, December. Adults found overwintering (SC). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Slow runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping; sweeping grass. References. Leng, 1915 (ecology, dispersal power); Löding, 1945 (ecology). Ardistomis schaumii LeConte, 1857 Ecology. Lowlands. Borders of lakes, ponds, marshes, and slow rivers; swamps, swampforests and margins of irrigation canals; wet meadows and seashore. Open ground; soft, wet, muddy or clayish soil covered with sparse vegetation (e.g., grass) or bare. Nocturnal; sheltering during the day in burrows dug in the soil, under dead leaves or weeds. Biology. Seasonality: January-August, October-November. Tenerals: early June (FL), late July (AL, TX). Adults found overwintering on higher and drier ground, in the upper zone of sand pits, river banks, and roadsides; in the soil (as deep as 4 cm). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Slow runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping. Reference. Nichols, 1988 (ecology, dispersal power). Asaphidion alaskanum Wickham, 1919 Ecology. Lowlands. Banks of rivers and brooks. At a considerable distance from water. Ground open or half-shaded (e.g., by Salix-bushes); soil composed of fine sand, covered with sparse vegetation. Diurnal (mostly) and nocturnal; active in the sunshine; sheltering on cloudy or cool days under dead leaves and other debris. Biology. Seasonality: June-August. Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Raking the leaf litter. Reference. Lindroth, 1963b (natural history). Asaphidion flavipes (Linné, 1761) Ecology. Lowlands. Light deciduous forests (mostly; along stream banks), swamps, meadows, vacant lots, city yards and gardens. Greenhouses (occasionally). Associated with habitats rich in springtails. Open or half-shaded ground; moist or wet, clayish or sandy soil covered with sparse vegetation and leaf litter. Diurnal (mostly) and nocturnal; active in the sunshine; sheltering on cloudy or cool days under dead leaves (mostly), fallen tree bark, and wood debris. Biology. Seasonality: February-September. Copulating pairs: May (CT). Eggs are laid singly in moist soil. Adults found overwintering under dead leaves. Adult food, in the field (Europe): springtails and mites. Defensemechanism: when threatened, the adult stays stationary; body color-pattern matching surroundings. Dispersal power. Macropterous. Occasional flier (during the day). Fast runner. Collecting techniques. Raking the leaf litter; pitfall trapping. References. Lindroth, 1945 (natural history); Bauer, 1971 (biology); Sharova, 1981 (ecology, biology); Krinsky, 1981 (ecology, biology); Lindroth, 1985 (natural history); Koch, 1989 (ecology); Loreau, 1985 (biology); Luff, 1993 (biology). Comments. Body shape and quick movements similar to ants.

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Asaphidion yukonense Wickham, 1919 Ecology. Lowlands and mountains. Stream banks. Open ground; moist, more or less clay soil covered with patches of tiny moss. Diurnal (mostly) and nocturnal; active in the sunshine; sheltering on cloudy or cool days under dead leaves. Biology. Seasonality: May-August. Adult food, in captivity: mealworms. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Raking the leaf litter. Reference. Lindroth, 1963b (natural history). Aspidoglossa subangulata (Chaudoir, 1843) Ecology. Lowlands. Borders of rivers, brooks, lakes, and ponds; wet meadows, moist pastures, cultivated fields (e.g., alfalfa, soybean), and low deciduous forests. Open or slightly shaded ground; soft, wet or moist, muddy or clayish soil, sometimes mixed with sand, bare or covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil, under dead leaves, stones, debris, and the loose bark of logs. Biology. Seasonality: January-September, November. Tenerals: mid-July (LA), late July (TX). Adult food, in the field: weevil larvae. Predators: catbirds and ducks. Parasites: pyemotid mites and Laboulbeniales (Fungi). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Slow runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping. References. Ulke, 1903 (ecology, dispersal power); Blatchley, 1910 (ecology); Leng, 1915 (ecology, dispersal power); Kirk, 1969 (dispersal power). Athrostictus punctatulus (Putzeys, 1878) Ecology. Forests (mostly), pastures, and gardens. Mostly shaded ground; dry soil. Biology and Dispersal power. Unknown. References. Bousquet and Larochelle, 1993 (distribution); Ball and Bousquet, 2001 (taxonomy). Atranus pubescens (Dejean, 1828) Ecology. Lowlands and mountains. Beaver houses (in muddy walls). Wooded brook edges: in heaps of mud and vegetal debris. Flood-plain forests and caves (southeastern USA). Open or shaded ground; wet, muddy or sandy soil, bare or covered with sparse vegetation. Nocturnal; sheltering during the day in mud cracks, under well-embedded pieces of wood and logs, in heaps of mud, twigs, and dead leaves, and under stones. Gregarious. Associated in beaver houses with Platypatrobus lacustris Darlington and Pterostichus castor Goulet and Bousquet. Biology. Seasonality: January-November. Tenerals: late June to September, November; mostly from mid-July on. Adults found overwintering in woods; under logs, tree-stumps, and the loose bark of fallen trees. Adult and larval food, in captivity: mealworms. Parasites: Laboulbeniales (Fungi). Phoretic mites found on adult body. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Strong burrower. Good swimmer. Favored by spring floods along streams and by laws protecting the beaver habitat. Collecting techniques. Inspecting

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the walls of beaver houses; dismembering the heaps of mud, twigs, and leaves along brooks. References. Ulke, 1903 (ecology); Blatchley, 1910 (ecology); Lindroth, 1968 (natural history); Erwin, 1981 (natural history). Axinopalpus biplagiatus (Dejean, 1825) Ecology. Lowlands and uplands. Grasslands, meadows, prairies, pastures, abandoned fields, cultivated fields, roadsides, sand hills, and sand dunes. Open ground; well-drained, dry, sandy soil covered with sparse vegetation (e.g. grass). Mostly diurnal; active in the sunshine on the soil (mostly) or on flowers; sheltering on cloudy or cool days under stones, dead leaves, pieces of wood, and other debris. Gregarious (up to 80 individuals found under a single stone). Biology. Seasonality: January-December. Tenerals: mid-July (QU). Adults found overwintering on hills and roadsides; in leaf litter and under the loose bark of trees. Predators: flickers (woodpeckers). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants and trees). Moderately favored by human activities. Collecting techniques. Lifting large stones from sandy dry fields; pitfall trapping; light trapping; sweeping plants; using tanglefoot screens. References. Blatchley, 1910 (ecology); Lindroth, 1969a (ecology, dispersal power); Bell, 1971 (ecology); Larochelle, 1974a (ecology); Kirk, 1975c (ecology, dispersal power); Lavigne, 1978 (ecology). Axinopalpus californicus (Motschulsky, 1845) Natural History. Unknown. Axinopalpus coloradensis Casey, 1920 Natural History. Unknown. Axinopalpus demissus Casey, 1920 Natural History. Unknown. Axinopalpus denticulatus Hatch, 1949 Ecology. Unknown. Biology. Seasonality: May. Dispersal power. Unknown. Axinopalpus fusciceps LeConte, 1851 Ecology. Macrohabitat unknown. Nocturnal; sheltering during the day under debris (CA). Biology. Seasonality: March, July. Dispersal power. Unknown. Axinopalpus habilis Casey, 1920 Natural History. Unknown.

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Axinopalpus illectus Casey, 1920 Natural History. Unknown. Axinopalpus pratti Hatch, 1949 Ecology. Unknown. Biology. Seasonality: June. Dispersal power. Unknown. Axinopalpus utahensis Tanner, 1928 Natural History. Unknown. Axinopalpus vittatus Hatch, 1949 Natural History. Unknown. Aztecarpalus schaefferi Ball, 1970 Ecology. Lowlands. Forests. Mexico: open forests (Quercus) and a thorn forest, near streams or moist places. Nocturnal; sheltering during the day under stones and in thin leaf litter (Mexico). Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Moderate runner. References. Ball, 1970 (ecology, dispersal power); Ball and Bousquet, 2001 (ecology). Badister (Badister) elegans LeConte, 1880 Ecology. Lowlands and mountains. Macrohabitat unknown. Nocturnal; sheltering during the day. Biology. Seasonality: January, March, June, August-September. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Badister (Badister) ferrugineus Dejean, 1831 Ecology. Lowlands. Shores of an alkaline lake: open, moist, muddy-sandy soil covered with sparse vegetation (OR). Nocturnal; sheltering during the day under mats of plant debris. Biology. Seasonality: June-August. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Badister (Badister) flavipes flavipes LeConte, 1853 Ecology. Lowlands. Flood-plain forests: along ponds. Shaded ground; wet soil. Nocturnal; sheltering during the day under vegetal debris. Biology. Seasonality: March-June, August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Lifting vegetal debris. Reference. Erwin, 1981 (natural history).

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Badister (Badister) flavipes laticeps Blatchley, 1910 Ecology. A low, damp situation in a secondary flood-plain forest (MD). A shaded, moist river bank (AR). Shaded ground; moist soil, bare or covered with depressed vegetation. Nocturnal; sheltering during the day. Biology. Seasonality: April-October. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Badister (Baudia) grandiceps Casey, 1920 Ecology. Lowlands. Borders of eutrophic marshes, ponds, and lakes; swamps and open swamp-forests. Often associated with quite small bodies of water. Somewhat shaded ground; wet, clayish or muddy soil, rich in organic debris, covered with dense vegetation (e.g., Typha, Menyanthes, Solanum, Carex). Nocturnal; sheltering during the day in leaf litter, under dead plants and pieces of fallen tree bark. Biology. Seasonality: January, April-September, November-December. Tenerals: August (MN, NE, QU). Adults found overwintering on higher and drier ground, at the edge of woods, in woods, and on hillocks; in leaf litter (mostly) and in the soil (as deep as 10 cm). Dispersal power. Macropterous. Frequent flier (to artificial lights at night; to artifical lights in captivity). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Treading the vegetation into the water; light trapping. References. Lindroth, 1969a (natural history); Kirk, 1975c (dispersal power). Badister (Badister) maculatus LeConte, 1853 Ecology. Lowlands. Flood-plain forests (e.g., cypress). Shaded ground; wet soil. Nocturnal; sheltering during the day under loose bark and vegetal debris. Biology. Seasonality: April-July, October. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Occasional climber (on trees). Collecting techniques. Lifting vegetal debris; light trapping; beating trees. References. Ulke, 1903 (dispersal power); Blatchley, 1910 (ecology); Kirk, 1969 (dispersal power). Badister (Badister) neopulchellus Lindroth, 1954 Ecology. Lowlands and mountains. Borders of eutrophic marshes, ponds, and lakes; marshy edges of pools; swamps, swamp-forests, and flood-plain forests. Shaded ground; wet or moist, muddy or clayish soil, rich in organic debris, covered with dense tall plants (e.g., Typha and Salix). Nocturnal; sheltering during the day in leaf litter, floating mats of dead plants, under moss, and at the base of plants. Biology. Seasonality: January-November. Tenerals: August-September. Adults found overwintering on higher and drier ground, on hillocks, hills, in sand dunes, pastures, at the edge of woods, in woods and hawthorn hedges; in leaf litter, under stones, and in the soil (as deep as 7.5 cm). Active at low temperatures under snow in October-November. Adult food, in the field: carabid eggs. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; to artifical lights in captivity). Occasional in lakeshore drift material, which indicates

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flight ability. Moderate runner. Moderately favored by human activities. Collecting techniques. Treading the vegetation into the water; light trapping. References. Lindroth, 1969a (natural history); Kirk, 1975c (dispersal power). Badister (Badister) notatus Haldeman, 1843 Ecology. Lowlands and mountains. Steep slopes of disturbed habitats: roadsides, gravel pits, sand pits, vacant lots, hills, pastures, relict prairies, cultivated fields, gardens, fencerows, and forest clearings. Open ground; warm, dry, gravelly (mostly) or sandy soil covered with scattered stones and moderate or dense vegetation (e.g., grass). Nocturnal; sheltering during the day under stones. Biology. Seasonality: January-December. Tenerals: late June (QU), early August (NE). Adults found overwintering on higher and drier ground, on hills, hillocks, in vacant lots, pastures, and the upper zone of gravel pits and sand pits; under stones, in leaf litter, in the soil (7.5 cm), and under reindeer moss (Cladonia). Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (to artificial lights at night). Moderate runner. Moderately favored by human activities. Collecting techniques. Pitfall trapping; lifting small stones. References. Lindroth, 1969a (natural history); Larochelle, 1974a (ecology); Kirk, 1975c (ecology); Erwin, 1981 (natural history). Badister (Badister) obtusus LeConte, 1878 Ecology. Lowlands and mountains. Deciduous forests and their edges, hawthorn hedges, thickets, and cultivated fields (e.g., barley). Shaded ground; rather moist, gravelly or stony soil covered with thick leaf litter. Nocturnal; sheltering during the day among dead leaves and under stones. Biology. Seasonality: January, March-October. Tenerals: August (ON, QU). Adults found overwintering in woods and hawthorn hedges; in leaf litter and under stones (mostly), also under pieces of wood. Dispersal power. Macropterous. Occasional flier (into rotary traps). Moderate runner. Collecting techniques. Raking and sifting the leaf litter; using rotary traps. References. Lindroth, 1969a (natural history); Larochelle, 1974a (ecology). Badister (Baudia) ocularis Casey, 1920 Ecology. Lowlands. Borders of eutrophic marshes, ponds, and lakes; swamps and wet meadows. Somewhat shaded ground; wet, clayish or muddy soil, rich in organic debris, covered with dense vegetation (e.g., Typha, Menyanthes, Solanum, Carex). Nocturnal; sheltering during the day in leaf litter and moss, and at the base of plants. Biology. Seasonality: March-October. Tenerals: August (QU)-September (NS). Adults found overwintering on higher and drier ground, in woods and at the edge of woods; in leaf litter (mostly), in the soil, and in rotten logs. Parasites: Laboulbeniales (Fungi). Phoretic mites found on adult body. Dispersal power. Macropterous. Occasional flier (to artificial lights at night; to artifical lights in captivity). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting technique. Treading the vegetation into the water. Reference. Lindroth, 1969a (natural history).

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Badister (Baudia) parviceps Ball, 1959 Ecology. Lowlands. Low deciduous forests: along lakes, ponds, eutrophic marshes, and slow rivers; flood-plain forests. At a certain distance from water. Shaded ground; moderately moist soil covered with thick leaf litter. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: March-May, July, October-November. Adults found overwintering on higher and drier ground, in woods and in the upper zone of sand pits; in leaf litter. Dispersal power. Macropterous. Regular flier (to artificial lights at night). Moderate runner. Collecting techniques. Treading the vegetation into the water; light trapping; sifting the leaf litter; using berlese funnels. References. Lindroth, 1969a (natural history); Ball, 1992 (ecology). Badister (Badister) pulchellus LeConte, 1848 Ecology. Lowlands. Borders of a small canalised brook, shaded by Salix, Ulmus, Acer, and Cornus, and covered with dense tall vegetation (e.g., Solanum, Typha) (ON). Nocturnal; sheltering during the day. Biology. Seasonality: June. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Treading the vegetation into the water. Reference. Lindroth, 1969a (natural history). Badister (Baudia) reflexus LeConte, 1880 Ecology. Lowlands and mountains. Flood-plain forests, low deciduous forests, borders of eutrophic ponds and marshes; swamps. Somewhat shaded ground; wet, clayish or muddy soil covered with thick leaf litter. Nocturnal; sheltering during the day under dead leaves and other vegetal debris. Biology. Seasonality: January-October. Adults found overwintering in woods; in leaf litter. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on ferns). Collecting techniques. Pouring water over the ground; raking the leaf litter from the soil; sweeping ferns. References. Ulke, 1903 (ecology); Kirk, 1969 (biology); Erwin, 1981 (natural history). Badister (Baudia) submarinus Motschulsky, 1859 Ecology. Lowlands. Biology. Seasonality: October. Dispersal power. Macropterous, probably capable of flight. Badister (Trimorphus) transversus Casey, 1920 Ecology. Lowlands. Swamps, swamp-forests, low deciduous forests, flood-plain forests, and borders of eutrophic marshes. Shaded ground; wet, clayish or muddy soil, rich in organic debris. Nocturnal; sheltering during the day under dead leaves, pieces of fallen tree bark, and other vegetal debris. Biology. Seasonality: April-November. Tenerals: late May (ON). Adults found overwintering on higher and drier ground, in woods

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and vacant lots; in rotten tree-stumps and under stones. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; into rotary traps). Moderate runner. Occasional climber (on plants). Collecting techniques. Pouring water over the ground; treading the soil with the feet; by light trapping; using rotary traps; sweeping plants. References. Larochelle, 1974a (ecology); Kirk, 1975c (dispersal power); Will et al. 1995 (dispersal power). Badister (Badister) vandykei Ball, 1959 Ecology. Unknown. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Batenus (Platynidius) angustatus angustatus (Dejean, 1828) Ecology. Lowlands and mountains. Low woods (deciduous, mixed, coniferous): often along streams, ravines, and slopes. Shaded ground; moist soil covered with thick leaf litter. Mostly nocturnal; usually sheltering during the day under the loose bark of fallen or standing trees, under logs, stones, and moss; sometimes active in the daytime on trees; active at night on tree trunks and on the ground. Biology. Seasonality: April-November. Tenerals: May-November; mostly before mid-summer. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Frequent climber (on trees, bushes, and plants). Collecting techniques. Pitfall trapping; beating or sweeping the vegetation; night collecting with a headlamp. References. Liebherr, 1986a (dispersal power) and 1988b (ecology, dispersal power); Morvan, 2000 (taxonomy). Batenus (Platynidius) angustatus hypolithos (Say, 1823) Ecology. Lowlands and mountains. Deciduous forests (e.g., Acer, Quercus) bordering rivers and brooks; woods slopes, forest clearings, edges and roads; neighboring fields. Shaded ground; moist soil, rich in leaf litter, often covered with scattered stones. Nocturnal; sheltering during the day under stones (mostly), the loose bark of logs, fallen trees, and tree-stumps, under dead leaves and moss carpets. Biology. Seasonality: AprilOctober. Tenerals: late May to September; mostly from mid-July on. Adult food, in the field: lepidopterous immature stages. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Frequent climber (on tree-stumps, logs, fallen and standing trees). Collecting techniques. Pitfall trapping; turning stones; beating trees; using malaise traps. References. Blatchley, 1910 (ecology); Lindroth, 1966 (natural history); Liebherr, 1988b (dispersal power); Morvan, 2000 (taxonomy). Batenus (Batenus) cincticollis (Say, 1823) Ecology. Lowlands. Low forests (deciduous, mixed) along lakes, ponds, and brooks; flood-plain forests, swamp-forests, swamps, and beaver houses. Shaded ground; wet, muddy, clayish (often) or peaty soil, rich in organic debris. Mostly nocturnal; sometimes active during the day; usually sheltering in leaf litter, moss, under stones, loose

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bark, in tree hollows, on trees and plants. Gregarious. Biology. Seasonality: JanuaryDecember. Tenerals: April (IL), September (VT). Adults found overwintering at the border of ponds; under vegetal debris and in moss. Adult food, in the field: corn. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Frequent in drift material (sea- and lake shores), which indicates flight ability. Moderate runner. Frequent climber (on trees and plants). Collecting techniques. Pitfall trapping; raking the leaf litter; beating or sweeping the vegetation; turning drift material; using tanglefoot screens. References. Blatchley, 1910 (biology); Lindroth, 1966 (natural history); Kirk, 1969 (ecology, dispersal power) and 1970 (biology); Erwin, 1981 (natural history); Morvan, 2000 (taxonomy). Batenus (Batenus) mannerheimii (Dejean, 1828) Ecology. Lowlands, mountains, subalpine and alpine zones. Coniferous forests (mostly Picea; Larix, Abies); thickets (Alnus) bordering lakes, ponds, and slow rivers; Sphagnumbogs (with Sarracenia, Comarum, Rubus); open Carex-marshes; open subalpine scrubland clearings (Picea); moist alpine meadows. Europe: old coniferous forests (mostly spruce mires). Open or shaded ground; often soft wet peaty soil covered with Sphagnum-moss, Sarracenia, Comarum or Rubus. Mostly nocturnal; usually sheltering during the day in moss carpets or under the loose bark of dead or dying standing trees (e.g., Picea, Larix, Abies, Acer). Sometimes active during the day on shrubs and bushes (e.g., Alnus, Salix). Biology. Seasonality: April-October. Gravids: June-July (QU). Tenerals: June-August; mostly August. Adult food, in the field: coleopterous larvae. Adult and larval food, in captivity: mealworms. Predators: toads. Also eaten by insectivorous plants (Sarracenia). Phoretic mites found on adult body. Dispersal power. Submacropterous, probably incapable of flight. Moderate runner. Frequent climber (on trees, shrubs, and bushes). Collecting techniques. Pitfall trapping; treading the moss into water; beating the vegetation. References. Lindroth, 1945 and 1955a (natural history); Krogerus, 1960 (ecology); Lindroth, 1963a, 1966 and 1986 (natural history); Niemelä et al. 1987 (ecology); Lindelöw, 1990 (ecology, biology); Niemelä et al. 1992 (ecology, biology); Morvan, 2000 (taxonomy). Bembidion acticola Casey, 1884 Ecology. Lowlands. Biology. Seasonality: April-June, August, October. Dispersal power. Unknown. Bembidion actuosum Casey, 1918 (Bembidion debilicolle Casey, 1924) Ecology. Mountains. Margins of small streams. Half-shaded ground; moist soil covered with moss. Nocturnal; sheltering during the day among moss. Biology. Seasonality: May-August. Tenerals: late May. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Turning moss carpets. Reference. Lindroth, 1963b (natural history; as debilicolle Casey).

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Bembidion (Trepanedoris) acutifrons LeConte, 1879 Ecology. Lowlands and mountains. Margins of lakes, ponds, sloughs, and slow brooks. Open ground; wet, clay soil covered with some vegetation (e.g., Typha, Juncus, grass). Nocturnal; sheltering during the day among vegetal debris. Biology. Seasonality: AprilAugust. Dispersal power. Wing-dimorphic: macropterous (mostly; capable of flight) or brachypterous (incapable of flight). Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Treading the soil with the feet; pouring water over the ground. References. Lindroth, 1963b (natural history); Kirk, 1975c (dispersal power). Bembidion (Bembidion) adductum Casey, 1918 Natural History. Unknown. Bembidion (Notaphus) aeneicolle (LeConte, 1848) Ecology. Lowlands and uplands. Margins of alkaline lakes, ponds, cattle water holes, and slow rivers and brooks. Close to water. Open ground; moist, clayish, feebly alkaline soil covered with sparse vegetation (e.g., Salicornia, Atriplex, Distichlis). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in cracks in the soil, under vegetal debris and stones. Biology. Seasonality: May-October. Tenerals: late June, mostly August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Hunting with a butterfly net; pouring water over the ground; treading the soil with the feet. References. Lindroth, 1963b (natural history); Bell, 1971 (ecology); Kirk, 1975c (ecology, dispersal power). Bembidion (Odontium) aenulum Hayward, 1901 Ecology. Lowlands and mountains. Bars and banks of rivers. Open ground; moist, bare soil consisting of very coarse sand, sometimes mixed with fine clay. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in soil cracks, burrows dug in the ground, and under pieces of wood. Biology. Seasonality: May, JulySeptember. Defense-mechanism: when alarmed, the adult escapes by flying readily; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (in the bright sunshine, when alarmed). Fast runner. Collecting techniques. Hunting with a butterfly net; pouring water over the ground. Reference. Choate and Miliotis, 1979 (ecology). Bembidion (Furcacampa) affine Say, 1823 Ecology. Lowlands and mountains. Margins of rivers, ponds, lakes, and pools; flood-plain forests, swamp-forests, eutrophic marshes, and swamps. Open or slightly shaded ground; wet, muddy or clayish soil covered with moderate or dense vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in leaf litter and under pieces of wood. Biology. Seasonality: February-November. Tenerals: July-August (MD).

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Adults found overwintering (AL, IN, SC). Defense-mechanism: body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in drift material, which indicates flight ability. Moderate runner. Collecting techniques. Treading the soil with the feet; pouring water over the ground; light trapping; using car nets. References. Blatchley, 1910 (biology); Lindroth, 1963b (natural history); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Bembidion (Bracteon) alaskense Lindroth, 1962 Ecology. Lake shores and river banks. Open ground; moist, sandy bare soil, sometimes mixed with clay or silt. Mostly diurnal; active in the bright sunshine; sheltering on cloudy or cool days in soil cracks and burrows dug in the ground. Defense-mechanism: when alarmed, the adult escapes by flying readily; body color-pattern matching the soil. Biology. Seasonality: June-July. Dispersal power. Macropterous. Frequent flier (when alarmed; in the bright sunshine). Fast runner. Collecting technique. Hunting with a butterfly net. Reference. Maddison, 1993 (ecology). Bembidion (Hirmoplataphus) alpineanum Casey, 1924 Ecology. Alpine zone. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion (Ochthedromus) americanum Dejean, 1831 Ecology. River banks, lake and pond shores. Open ground; soft, wet, sandy, muddy or clayish soil, bare or covered with sparse vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in cracks in the soil, in leaf litter, under mats of dead plants, and at the base of plant tufts (Carex). Biology. Seasonality: March-December. Tenerals: late July (WV), August (VT). Adult food, in the field: a mosquito larva. Defense-mechanism: when alarmed, the adult escapes by running and flying readily like a tiny Cicindela; the beetle is difficult to pursue; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; to artificial lights at night). Fast runner. Collecting techniques. Pouring water over the ground; hunting with a butterfly net; using tanglefoot screens. References. Löding, 1945 (ecology); Kirk, 1969 (ecology, dispersal power); Erwin, 1981 (natural history). Bembidion (Trepanedoris) ampliceps Casey, 1918 Ecology. Unknown. Biology. Seasonality: May-August. Dispersal power. Macropterous, probably capable of flight. Bembidion (Trepanedoris) anguliferum (LeConte, 1852) Ecology. Margins of a small brook, on clay soil. Biology. Seasonality: June-July. Dispersal power. Macropterous, probably capable of flight.

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Bembidion (Notaphus) approximatum (LeConte, 1852) Ecology. Unknown. Biology. Seasonality: May-July, October. Dispersal power. Macropterous, probably capable of flight. Bembidion (Eupetedromus) aratum (LeConte, 1852) Ecology. Stream margins. Open ground; moist or wet, gravelly-stony soil, mixed with silt, covered with sparse vegetation. Nocturnal; sheltering during the day among gravel and under stones. Biology. Seasonality: July. Dispersal power. Macropterous. Regular flier (to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; raking the soil. Bembidion (Plataphodes) arcticum Lindroth, 1963 Ecology. Margins of brooks, ponds, and pools. Close to water. Open ground; moist, gravelly-stony, bare soil. Biology. Seasonality: June, August. Tenerals: August (AK). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pouring water over the ground; raking the soil. Reference. Lindroth, 1963b (natural history). Bembidion (Odontium) arizonae Lindroth, 1963 Ecology. Unknown. Biology. Seasonality: May, August-September. Dispersal power. Macropterous, probably capable of flight. Bembidion (Notaphus) auxiliator Casey, 1924 Ecology. Unknown. Biology. Seasonality: May-July, October. Dispersal power. Macropterous, probably capable of flight. Bembidion (Hirmoplataphus) avidum Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion (Bracteon) balli Lindroth, 1962 Ecology. Uplands. Banks and bars of rivers. Open ground; wet, sandy bare soil. Mostly diurnal; active in the bright sunshine; sheltering on cloudy or cool days in soil cracks, burrows dug in the ground, under stones and pieces of wood. Biology. Seasonality: JulySeptember. Defense-mechanism: when alarmed, the adult escapes by flying readily; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; in the bright sunshine). Fast runner. Collecting technique. Hunting with a butterfly net. References. Lindroth, 1963b (natural history); Maddison, 1993 (ecology).

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Bembidion (Plataphus) basicorne Notman, 1920 Ecology. Lowlands and mountains. Margins and bars of cold, swift forest streams. Shaded ground; wet, gravelly, bare soil covered with scattered stones. Nocturnal; sheltering during the day among gravel and under stones. Biology. Seasonality: March-September. Tenerals: late July to September. Dispersal power. Macropterous. Occasional in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pouring water over the ground; raking the loose gravel. Bembidion (Ochthedromus) bifossulatum (LeConte, 1852) Ecology. Lowlands and mountains. Margins of saline flats and ponds; edges of reservoirs and cattle water holes; river banks and lake shores; cultivated fields (e.g., corn, wheat) and meadows. Close to water. Open ground; clayish or muddy soil, bare or covered with sparse vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under stones and logs. Biology. Seasonality: February-September. Tenerals: late July (VA) to mid-September (BC). Defense-mechanism: when alarmed, the adult escapes by running and flying readily like a tiny Cicindela; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Fast runner. Collecting techniques. Pouring water over the ground; hunting with a butterfly net. References. Lindroth, 1963b and 1971a (natural history); Bell, 1971 (ecology); Kirk, 1975c (ecology, dispersal power); Will et al. 1995 (dispersal power). Bembidion bimaculatum (Kirby, 1837) Ecology. Lowlands and mountains. Banks of large rivers. At some distance from water. Cultivated fields and their borders (e.g., barley, fescue). Open ground; moist, clay soil covered with sparse vegetation. Biology. Seasonality: June-September. Gravids: late July to early September (AB). Tenerals: June (AK, BC, YT). Adult food: dipterous pupae (in the field); lepidopterous eggs (in captivity). Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pitfall trapping. References. Lindroth, 1963b (natural history); Frank, 1971b (biology); Doane, 1981 (ecology, biology). Bembidion (Odontium) bowditchii LeConte, 1878 Ecology. Mountains. Banks of large rivers. Open ground; wet, silty or muddy bare soil. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in soil cracks and burrows dug in the ground. Biology. Seasonality: MayJuly. Defense-mechanism: when alarmed, the adult escapes by flying readily. Dispersal power. Macropterous. Frequent flier (when alarmed; in the bright sunshine). Fast runner. Collecting techniques. Pouring water over the ground; hunting with a butterfly net.

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Bembidion (Plataphodes) brachythorax Lindroth, 1963 Ecology. Stream banks. At some distance from water. Nocturnal; sheltering during the day in leaf litter (Salix). Biology. Seasonality: May-August. Tenerals: August (AK). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Raking the leaf litter. Reference. Lindroth, 1963b (natural history). Bembidion (Plataphodes) breve (Motschulsky, 1845) (Bembidion incertum (Motschulsky)) Ecology. Lowlands, mountains, subalpine and alpine zones. Borders of snowfields (mostly); margins of streams, drainage canals, ponds, and lakes. Associated with cold water. Open ground; moist soil. Nocturnal; sheltering during the day under stones and logs. Biology. Seasonality: May-September. Tenerals: late August (AK). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Turning stones. References. Lindroth, 1963b (natural history; as incertum (Motschulsky)); Kavanaugh, 1992 (natural history; as incertum (Motchulsky)). Bembidion (Hydriomicrus) brevistriatum Hayward, 1897 Ecology. Unknown. Biology. Seasonality: February-August, October. Dispersal power. Macropterous, probably capable of flight. Bembidion (Peryphus) bruxellense Wesmael, 1835 Ecology. Lowlands and mountains. Borders of temporary pools, marshes, and slow brooks; beaver houses, vacant lots, gardens, roadside ditches, meadows, sand pits, and gravel pits. Open ground; firm moist, clayish (mostly), gravelly or sandy soil covered with sparse vegetation. Nocturnal; sheltering during the day under small stones, pieces of cardboard or wood, heaps of hay, at the base of plants, and in cracks in the soil. Biology. Seasonality: May-October. Copulating pairs: late June (QU). Tenerals: July-August. Adult food, in the field (Europe): insects (e.g., chironomids, aphids). Dispersal power. Macropterous. Occasional flier (to artificial lights in captivity). Moderate runner. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones. References. Lindroth, 1945, 1955a (as rupestre Linné), 1963a, 1963b and 1985 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology). Bembidion (Hydriomicrus) californicum Hayward, 1897 Ecology. Unknown. Biology. Seasonality: May-June. Dispersal power. Macropterous, probably capable of flight. Bembidion callens Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight.

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Bembidion (Trepanedoris) canadianum Casey, 1924 Ecology. Lowlands and mountains. Borders of brooks, rivers, rivulets, roadside ditches, and marshes; moist meadows and cultivated fields (e.g., barley). Open ground; moist or wet, clay soil covered with sparse or moderate vegetation. Nocturnal; sheltering during the day under stones. Biology. Seasonality: May-July, September-November. Active at low temperatures under snow in October-November. Adult food, in captivity: lepidopterous eggs. Dispersal power. Wing-dimorphic: macropterous (probably capable of flight) or brachypterous (incapable of flight). Moderate runner. Collecting technique. Pitfall trapping. References. Lindroth, 1963b (natural history); Frank, 1971b (biology); Larochelle, 1974a (ecology). Bembidion (Odontium) carinatum (LeConte, 1852) Ecology. River banks. Biology. Seasonality: January, August. Defense-mechanism: when alarmed, the adult escapes by flying readily. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Hunting with a butterfly net. Bembidion (Bracteon) carinula Chaudoir, 1868 Ecology. Lowlands and uplands. Banks and bars of large rivers and lakes. Close to water. Open ground; moderately moist, bare soil composed of pure, sometimes rather coarse sand. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days in soil cracks, burrows dug in the ground, under dead fallen plants and pieces of wood. Biology. Seasonality: April-October. Copulating pairs: June-July (QU). Tenerals: mid-July to August. Predators: toads. Defense-mechanism: when alarmed, the adult escapes by flying readily like a tiny Cicindela; the beetle is difficult to pursue; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; in the bright sunshine). Fast runner. Collecting technique. Hunting with a butterfly net. References. Blatchley, 1910 (ecology); Lindroth, 1955a, 1963a, 1963b and 1985 (natural history); Maddison, 1993 (ecology). Bembidion (Plataphus) carolinense Casey, 1924 Ecology. Lowlands and mountains. Banks of cold, swift rivers and brooks. Close to water. Open ground; wet, gravelly, bare soil. Nocturnal; sheltering during the day among gravel and under stones. Gregarious. Biology. Seasonality: April-October. Copulating pairs: early July (QU). Tenerals: late June to September. Defense-mechanism: when alarmed, the adult may escape by flying. Dispersal power. Macropterous. Frequent flier (when alarmed; to artificial lights in captivity). Moderate runner. Collecting techniques. Pouring water over the ground; raking the loose gravel. References. Lindroth, 1955a, 1963a and 1963b (natural history).

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Bembidion (Notaphus) castor Lindroth, 1963 Ecology. Lowlands and mountains. Borders of rivers, brooks, ponds, lakes, and marshes; sea beaches. At some distance from water. Open ground; moist or wet, bare soil consisting of fine sand, usually clay-mixed, covered with a thin layer of organic debris. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in leaf litter, under fallen dead branches, and stones. Biology. Seasonality: January-December. Copulating pairs: May and early July (QU). Tenerals: June-September. Adults found overwintering on higher and drier ground, at the edge of woods, and on hillocks; in leaf litter (mostly) and under small stones. Predators: frogs and toads. Phoretic mites found on adult body. Defense-mechanism: body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (in the sunshine; to artificial lights at night; to artifical lights in captivity). Moderate runner. Occasional climber (on plants). Collecting techniques. Raking the leaf litter; pouring water over the ground; sweeping. References. Lindroth, 1963a and 1963b (natural history); Kirk, 1975c (ecology); Erwin, 1981 (natural history). Bembidion (Lionepha) castum Casey, 1918 Ecology. Lowlands and mountains. Edges of mixed and deciduous forests. Shaded ground; moderately moist, sandy (preferably) or silty soil covered with sparse vegetation (e.g., tiny moss). Nocturnal; sheltering during the day in leaf litter and under plant rosettes. Biology. Seasonality: January, March-November. Tenerals: May-August; mostly July-August. Dispersal power. Wing-dimorphic: macropterous (mostly; probable capable of flight) or brachypterous (incapable of flight). Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter; liting stones. References. Lindroth, 1963b (natural history); Kavanaugh, 1992 (natural history). Bembidion (Pseudoperyphus) chalceum Dejean, 1831 Ecology. Lowlands and mountains. Banks and bars of large rivers (mostly), lake shores, and sea beaches. Sometimes at a considerable distance from water. Open ground; moderately moist bare soil consisting of gravel or coarse sand. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day among gravel, under stones and dead leaves. Gregarious. Biology. Seasonality: January, March-September, November. Copulating pairs: mid-July (QU). Tenerals: mid-July to September. Adults found overwintering on higher and drier ground, in the upper zone of river banks and on sandy-gravelly hillocks; under dead leaves. Predators: frogs and toads. Phoretic mites found on adult body. Defense-mechanism: when alarmed, the adult escapes by flying readily; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; to artificial lights at night; to artifical lights in captivity). Moderate runner. Collecting techniques. Pouring water over the ground; raking the loose gravel. References. Lindroth, 1955a, 1963a, 1963b (natural history); Kirk, 1975c (dispersal power).

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Bembidion (Lionepha) chintimini Erwin and Kavanaugh, 1981 Ecology. Mountains. Biology. Seasonality: May. Dispersal power. Brachypterous, incapable of flight. Bembidion (Trepanedoris) clemens Casey, 1918 Ecology. Uplands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion (Notaphus) coloradense Hayward, 1897 Ecology. Uplands. Borders of ponds, sloughs, marshes, lakes, reservoirs, and slow rivers and brooks; wet meadows. Close to water. Open ground; wet clay soil, sometimes moderately alkaline, covered with sparse vegetation (e.g., Eleocharis, Carex, Equisetum). Adults respond to chemical cues associated with green algae. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under mats of plants and stones. Biology. Seasonality: April-September. Tenerals: mid-July to mid-August. Adult food, in the field: a coccinellid. Defense-mechanism: when alarmed, the adult may escape by flying; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. References. Lindroth, 1963b (natural history); Evans, 1983 (ecology). Bembidion (Bracteon) colvillense Lindroth, 1965 Ecology. Banks of a small river inlet, on bare sandy soil, with clayish spots (AK). Diurnal; active in the bright sunshine. Biology. Seasonality: unknown. Tenerals: August (AK). Defense-mechanism: when alarmed, the adult escapes by flying readily; body colorpattern matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; in the bright sunshine). Fast runner. Collecting technique. Hunting with a butterfly net. Reference. Lindroth, 1969a (natural history). Bembidion commotum Casey, 1918 Ecology. Mountains, subalpine and alpine zones. Margins of snowfields; alpine meadows (especially after snowmelt), pine forests, sagebrush areas, and brook margins. Open or shaded ground; moist, gravelly or rocky soil, sometimes covered with moss. Nocturnal; sheltering during the day under stones. Biology. Seasonality: April, June-October. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; turning stones. Reference. Lindroth, 1963b (natural history).

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Bembidion (Plataphodes) complanulum (Mannerheim, 1853) Ecology. Mountains, subalpine and alpine zones. Margins of snowfields (mostly); alpine meadows; margins of streams and drainage channels. Open ground; moderately moist soil covered with some vegetation. Nocturnal; sheltering during the day under stones. Biology. June-August. Tenerals: late June (BC), August (AB, AK). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Turning stones. References. Lindroth, 1963b (natural history); Kavanaugh, 1992 (natural history). Bembidion (Plataphodes) compressum Lindroth, 1963 Ecology. Mountains. Banks of cold, swift brooks and rivers. Open ground; moist, bare, stony-gravelly soil. Nocturnal; sheltering during the day under stones and among gravel. Biology. Seasonality: June-August. Tenerals: early August (AK). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Raking the soil. Reference. Lindroth, 1963b (natural history). Bembidion (Hirmoplataphus) concolor (Kirby, 1837) Ecology. Lowlands, mountains, and subalpine zone. Banks of rivers (mostly) and brooks; lake shores. Associated with cold, swift water. Open ground; wet bare soil consisting of gravel or coarse sand. Nocturnal; sheltering during the day under stones and among gravel. Gregarious. Biology. Seasonality: February, April-September, November. Tenerals: mid-July to August. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pouring water over the ground; raking the soil. References. Lindroth, 1955a (as longulum LeConte), 1963a and 1963b (natural history). Bembidion (Trepanedoris) concretum Casey, 1918 Ecology. Lowlands and mountains. Borders of eutrophic marshes, ponds, and lakes; swamps; marshy borders of slow rivers and brooks; peat bogs; borders of irrigation canals and roadside ditches; wet meadows. Open ground; wet, muddy, clayish or peaty soil, rich in organic debris, covered with dense vegetation (e.g., Carex, Typha, moss (never Sphagnum)). Nocturnal; sheltering during the day in leaf litter, under heaps of grass, in moss, at the base of plants, under stones, and in the soil. Gregarious. Biology. Seasonality: January-December. Tenerals: July-August, October; mostly from mid-July on. Adults found overwintering on higher and drier ground, on sandy hillocks, sand dunes, in field hedges, and at the edge of woods; in leaf litter (mostly), under heaps of grass, small stones, and in rotten logs. Phoretic mites found on adult body. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; to artifical lights in captivity). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Treading the vegetation into the water; light trapping. References. Lindroth, 1955a (as anguliferum LeConte), 1963a, and 1963b (natural history); Kirk, 1975c (ecology, dispersal power).

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Bembidion (Odontium) confusum Hayward, 1897 Ecology. Lowlands and mountains. Banks and bars of rivers and brooks (mostly); shores of ponds and lakes. Close to water. Open ground; wet, sandy bare soil, often mixed with fine clay. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in holes dug in the ground, in heterocerid soil galleries, and under stones and vegetal debris. Gregarious. Biology. Seasonality: January-December. Copulating pairs: early June (QU). Tenerals: July-October. Overwintering in the larval, pupal, and adult stages (OK). Adult food, in the field: small insects. Larval food, in the field: heterocerid larvae, springtails, and larvae of its own species. Predators: toads. Defense-mechanism: when alarmed, the adult escapes by running and flies readily like a tiny Cicindela, or emits a spray from the pygidial glands; the beetle is difficult to pursue; body colorpattern matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; in the bright sunshine; to artificial lights at night). Fast runner. Collecting techniques. Pouring water over the ground; colllecting with a butterfly net. References. Lindroth, 1963b (natural history); Kirk, 1975c (ecology, dispersal power); Erwin, 1981 (natural history); Kaufmann, 1986 (biology). Bembidion (Trepanedoris) connivens (LeConte, 1852) Ecology. Lowlands and mountains. Borders of eutrophic marshes and ponds; marshy borders of slow rivers and brooks; wet meadows. Usually at some distance from water. Ground open or shaded by trees (e.g., Populus); wet soil consisting of clay or fine sand, covered with dense vegetation (e.g., grass). Nocturnal; sheltering during the day under vegetal debris and stones. Biology. Seasonality: January, March, May-October. Tenerals: early July (ID, OR). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Treading the vegetation into the water. Reference. Lindroth, 1963b (natural history). Bembidion consanguineum Hayward, 1897 Ecology. Uplands. Stream margins; shores of small lakes. Open ground; moist or wet soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: AprilAugust. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Turning stones. Bembidion consimile Hayward, 1897 Ecology. Uplands. Borders of saline lakes and ponds; small muddy flats. Open ground; moist, clayish saline soil, bare or covered with sparse vegetation (e.g., Distichlis, Atriplex). Nocturnal; sheltering during the day in cracks in the soil. Biology. Seasonality: March, June-August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. References. Lindroth, 1963b (natural history); Kirk, 1975c (dispersal power).

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Bembidion (Notaphus) conspersum Chaudoir, 1868 Ecology. Coastal lowlands. Salt meadows. Open ground. Biology. Unknown. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Bembidion constricticolle Hayward, 1897 Ecology. Uplands. Biology. Seasonality: May. Dispersal power. Brachypterous, incapable of flight. Bembidion constrictum (LeConte, 1848) Ecology. Lowlands. Sea beaches; borders of lagoons, salt marshes, and salt ponds. Open ground; wet, clayish saline soil. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under mats of dead algae and grass, in cracks in the soil, and under stones. Biology. Seasonality: April-November. Tenerals: early July (NS). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Lifting mats of dead algae and grass. References. Lindroth, 1963b (natural history); Kirk, 1975c (dispersal power). Bembidion (Notaphus) consuetum Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion contractum Say, 1823 Ecology. Coastal lowlands. Salt marshes, saline ponds, salt meadows, sea beaches (upper limit of tidal zone), lagoons, and banks of estuarine rivers. Open ground; flat, wet, sandy soil mixed with clay and covered with a thin crust of green algae. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in cracks in the soil, at the base of plants, under algal mats, and under stones. Biology. Seasonality: January-November. Copulating pairs: early July (NS). Tenerals: late June-August. Adult food, in the field: a dead fly. Defense-mechanism: body color-pattern matching the soil. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; turning debris. References. Leng, 1915 (ecology, dispersal power); Lindroth, 1955a, 1963a, and 1963b (natural history). Bembidion (Semicampa) convexulum Hayward, 1897 Ecology. Upper zone of river and brook banks; margins of pools, ponds, small lakes, and eutrophic marshes; mud flats. Open ground; soil consisting of sand-mixed clay, covered with some vegetation (e.g., grass, Carex, Juncus, Scirpus). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under dead plants, dead

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leaves, pieces of wood and stones. Biology. Seasonality: June-July. Tenerals: early and mid-July (ID, MT, WA). Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous (probably capable of flight). Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Reference. Lindroth, 1963b (natural history). Bembidion (Notaphus) cordatum (LeConte, 1848) Ecology. Lowlands and uplands. Borders of ponds and lakes. Close to water with emerging vegetation. Open ground; wet muddy soil composed of sand-mixed clay. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under vegetal debris and stones. Biology. Seasonality: March-October. Tenerals: mid-August (MN). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. References. Bell, 1971 (ecology); Kirk, 1975c (ecology, dispersal power); Will et al. 1995 (dispersal power). Bembidion (Odontium) coxendix Say, 1823 Ecology. Lowlands and mountains. Banks of rivers and brooks. Open ground; soft, moist, sandy or clayish, bare soil. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in soil cracks or burrows dug in the ground. Biology. Seasonality: April-November. Defense-mechanism: when alarmed, the adult escapes by flying readily; the beetle is difficult to pursue; body color matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed). Fast runner. Collecting techniques. Pouring water over the ground; collecting with a butterfly net. References. Blatchley, 1910 (ecology); Lindroth, 1963b (natural history); Kirk, 1975c (dispersal power). Bembidion (Plataphus) curtulatum Casey, 1918 (Bembidion flebile Casey, 1918) Ecology. Mountains. Margins of usually large streams, irrigation canals, and snowfields. Open ground; moist, gravelly, bare soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: May-July. Tenerals: late May and late July (BC). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Turning stones. Reference. Lindroth, 1963b (natural history; as flebile Casey). Bembidion (Peryphanes) dauricum (Motschulsky, 1844) Ecology. Lowlands. Tundra. Open ground; rather dry soil consisting of fine sand or peat, covered with sparse vegetation, sometimes bare. Biology. Seasonality: June-September. Tenerals: late August (AK). Dispersal power. Wing-dimorphic: macropterous (probably capable of flight) or brachypterous (incapable of flight). Moderate runner. References. Lindroth, 1945, 1963b and 1985 (natural history).

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Bembidion (Notaphus) debiliceps Casey, 1918 Ecology. Margins of pools and eutrophic marshes. Open ground; moist, clayish or peaty soil covered with dense vegetation. Biology. Seasonality: May-June. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Treading the vegetation into the water. Reference. Lindroth, 1963b (natural history). Bembidion dejectum Casey, 1884 Ecology. Uplands. Borders of saline waters, e.g., lakes. Open ground; moist, clayish saline soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: May-August, October. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pouring water over the ground; turning stones. Reference. Lindroth, 1963b (natural history). Bembidion (Emphanes) diligens Casey, 1918 Ecology. Uplands. Borders of saline lakes, ponds, sloughs, reservoirs, slow rivers and brooks; salt flats. Open ground; moist, clayish or muddy soil, bare or covered with moderately dense vegetation (e.g., Distichlis, Salicornia). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in cracks in the soil, at the base of plants, and under stones. Biology. Seasonality: April-August. Tenerals: late June to August; mostly August. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. References. Lindroth, 1963b (natural history); Kirk, 1975c (ecology); Ball and Bousquet, 2001 (taxonomy). Bembidion (Lionepha) disjunctum Lindroth, 1963 Ecology. Mountains. Margins of cold streams. Open ground; moist, gravelly soil. Biology. Seasonality: May-July, September. Tenerals: September (CA). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pouring water over the ground; raking the gravel. Bembidion (Notaphus) dorsale Say, 1823 Ecology. Lowlands and uplands. Banks of large rivers. At some distance from water. Ground open (mostly) or shaded by scattered trees (e.g., Salix, Populus); moist, clay soil, bare or almost so. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under debris. Biology. Seasonality: February, May-July. Adults found overwintering under pieces of wood. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Pouring water over the ground. References. Lindroth, 1963b (natural history); Kirk, 1975c (dispersal power).

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Bembidion (Odontium) durangoense Bates, 1891 Ecology. Unknown. Biology. Seasonality: September. Dispersal power. Unknown. Bembidion (Metallina) dyschirinum LeConte, 1861 Ecology. Mountains. Sandhills, moraines, vacant lots, roadsides, and forest edges. Open or half-shaded ground, under bushes (e.g., Alnus); well-drained, dry, sandy soil, partially bare and covered with dead leaves or tiny moss. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in leaf litter and under stones. Biology. Seasonality: May-September. Tenerals: early August (AB). Dispersal power. Wing-dimorphic: brachypterous (incapable of flight) or macropterous (probably capable of flight). Fast runner. Collecting techniques. Pitfall trapping; raking the leaf litter; turning stones. References. Lindroth, 1963b (natural history); Kavanaugh, 1992 (natural history); Ball and Bousquet, 2001 (taxonomy). Bembidion egens Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion (Peryphodes) ephippigerum (LeConte, 1852) Ecology. Coastal lowlands. Seashore, salt meadows, and salt flats. Open ground; moist, sandy, saline soil. Biology. Seasonality: April-August. Dispersal power. Wing-dimorphic: macropterous (probably capable of flight) or brachypterous (incapable of flight). Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil. Reference. Lindroth, 1963b (natural history). Bembidion (Lionepha) erasum LeConte, 1859 Ecology. Lowlands and mountains. Margins of cold small rivers, rills, lakes, and snowfields. Shaded ground; moist, gravelly soil. Nocturnal; sheltering during the day among moss, dead leaves, and under stones. Biology. Seasonality: April-September. Tenerals: July-August. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pouring water over the ground; raking the gravel. References. Lindroth, 1963b (natural history); Erwin and Kavanaugh, 1981 (natural history). Bembidion (Notaphus) evidens Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion (Plataphus) falsum Blaisdell, 1902 Ecology. Mountains. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight.

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Bembidion (Plataphodes) farrarae Hatch, 1950 Ecology. Mountains, subalpine and alpine zones. Margins of snowfields (mostly) and lakes. Open ground; moist soil covered with sparse vegetation. Nocturnal; sheltering during the day under stones. Biology. Seasonality: June-August. Tenerals: late August (AK). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Turning stones. References. Lindroth, 1963b (natural history); Kavanaugh, 1992 (natural history). Bembidion (Peryphus) femoratum Sturm, 1825 Ecology. Lowlands. Abandoned fields. Open ground; moderately moist soil consisting of clay or clay-mixed sand, covered with sparse vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under stones, clods of clay, and in soil crevices. Biology. Seasonality: June-July. Dispersal power. Macropterous. Flight records (Europe). Moderate runner. Collecting technique. Turning stones. References. Lindroth, 1945 (natural history); Sharova, 1981 (ecology, biology); Meissner, 1983 (ecology, biology); Lindroth, 1985 (natural history); Koch, 1989 (ecology). Bembidion (Liocosmius) festivum Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous. Flight records (Europe). Bembidion flavopictum Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion flohri Bates, 1878 (Bembidion henshawi Hayward, 1897) Ecology. Uplands. Borders of saline lakes and ponds. Open ground; moist, clayish saline soil, bare or covered with sparse vegetation (e.g., Distichlis, Atriplex). Nocturnal; sheltering during the day in cracks in the soil, under pieces of wood, and at the base of plants. Biology. Seasonality: May-September. Tenerals: mid-July (MB), late July (UT). Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Reference. Lindroth, 1963b (natural history; as henshawi Hayward). Bembidion (Trepanedoris) fortestriatum (Motschulsky, 1845) Ecology. Lowlands, mountains, and subalpine zone. Borders of eutrophic marshes, lakes, ponds, and slow rivers in forested areas; swamps; wet meadows. Open or slightly shaded ground; wet, clayish or muddy soil, rich in organic debris, covered with dense vegetation (e.g., Carex, Comarum, Menyanthes, Smilacina). Mostly nocturnal; sometimes ac-

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tive in the sunshine; usually sheltering during the day in leaf litter and moss. Biology. Seasonality: January, March-September, November-December. Tenerals: mid-July to late September. Adults found overwintering on higher and drier ground, at the edge of woods, on sandy hillocks and mounds, and in vacant lots; in leaf litter (mostly) and under small stones. Dispersal power. Macropterous. Occasional flier (to artificial lights in captivity). Occasional in lakeshore drift material, which indicates flight. Moderate runner. Collecting technique. Treading the vegetation into the water. References. Lindroth, 1963a and 1963b (natural history); Kavanaugh, 1992 (natural history). Bembidion (Bracteon) foveum Motschulsky, 1844 Ecology. Lowlands and uplands. River banks (mostly) and lake shores. Often around backwater pools. Close to the water. Open ground; sandy, bare soil mixed with clay or silt. Mostly diurnal; active in the bright sunshine; sheltering on cloudy or cool days in soil cracks and burrows dug in the ground. Biology. Seasonality: June-September. Defense-mechanism: when alarmed, the adult escapes by running and flying readily like a tiny Cicindela; the beetle is difficult to pursue; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; in the sunshine). Fast runner. Collecting techniques. Pouring water over the ground; hunting with a butterfly net. References. Lindroth, 1963b (natural history); Maddison, 1993 (ecology). Bembidion (Trepanedoris) frontale (LeConte, 1848) Ecology. Lowlands and mountains. Margins of eutrophic marshes, ponds, and pools; marshy margins of slow rivers and brooks; swamps. Often along very small bodies of standing water. Ground shaded by bushes (e.g., Salix) or deciduous trees; wet, clay soil, mixed with organic debris, covered with dense vegetation and thick leaf litter. Mostly nocturnal; usually sheltering during the day under dead leaves. Biology. Seasonality: January-December. Copulating pairs: May (QU). Tenerals: May-November; mostly from mid-summer on. Adults found overwintering on higher and drier ground, in woods, at the edge of woods, and in abandoned fields; mostly in leaf litter (usually lying upside down under the lower side of leaves) and under small stones. Adult food: mosquito eggs (in the field); meat (in captivity). Predators: toads. Defense-mechanism: when alarmed, the adult escapes by digging into the leaf litter. Dispersal power. Macropterous. Frequent flier (to artificial lights in captivity). Frequent in drift material, which indicates flight ability. Moderate runner. Regular climber (on plants and trees). Favored by spring floods along large rivers. Collecting techniques. Raking the leaf litter; sweeping or beating the vegetation; using an aspirator; turning drift material. Reference. Lindroth, 1963b (natural history). Bembidion fuchsii Blaisdell, 1902 Ecology. Mountains. Margins of small brooks. Open ground; moist, gravelly-sandy soil. Nocturnal; sheltering during the day among gravel and sand. Biology. Seasonality: JulySeptember. Dispersal power. Macropterous, probably capable of flight. Moderate runner.

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Bembidion (Trichoplataphus) fugax (LeConte, 1848) Ecology. Lowlands and mountains. River banks. Open ground; moist, gravelly soil, sometimes mixed with sand and silt. Nocturnal; sheltering during the day under stones. Biology. Seasonality: April-September. Tenerals: July and August (MD). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pouring water over the ground; turning stones. Reference. Erwin, 1981 (natural history). Bembidion (Plataphus) gebleri turbatum Casey, 1918 Ecology. Uplands and subalpine zone. Banks of cold, mostly large rivers and brooks. Open ground; moist, gravelly bare soil. Biology. Seasonality: June-August. Tenerals: late July (AB, BC). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pouring water over the ground; raking the gravel. Bembidion (Plataphus) gordoni Lindroth, 1963 Ecology. Mountains. Banks of small swift streams. Biology. Seasonality: March, May, July-September. Tenerals: late July (BC). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Lindroth, 1963b (natural history). Bembidion (Eupetedromus) graciliforme Hayward, 1897 Ecology. Lowlands and mountains. Borders of pools (often small ones) and eutrophic marshes; swamps and swamp-forests along lakes and slow rivers. Open or moderately shaded ground; wet, muddy soil covered with rather dense vegetation (e.g., depressed grass) and thick leaf litter. Nocturnal; sheltering during the day under dead leaves, pieces of wood, dead grass, at the base of plants, and in the soil. Biology. Seasonality: January-December. Tenerals: mid-July to October. Adults found overwintering on higher and drier ground, in woods, at the edge of woods, in the upper zone of sand pits and gravel pits, in hawthorn hedges and vacant lots; in leaf litter (mostly), under small stones, dead grass, and the loose bark of fallen trees. Adult food, in captivity: a piece of earthworm. Predators: toads. Defense-mechanism: body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; to artifical lights in captivity). Moderate runner. Collecting techniques. Treading the soil with the feet; pouring water over the ground; light trapping. References. Lindroth, 1963b (natural history); Kirk, 1975c (dispersal power). Bembidion (Trichoplataphus) grandiceps Hayward, 1897 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight.

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Bembidion (Notaphus) graphicum Casey, 1918 Ecology. Lowlands, mountains, and subalpine zone. Margins of pools, ponds, lakes, reservoirs, slow rivers, brooks, and marshes. Open ground; wet clay soil, alkaline or moderately saline, bare or covered with sparse vegetation (e.g., Distichlis). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in cracks in the soil, under plant debris, and under stones. Biology. Seasonality: May-June, August, October-November. Tenerals: late June to September. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Reference. Lindroth, 1963b (natural history). Bembidion (Peryphanes) grapii Gyllenhal, 1827 Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, vacant lots, meadows, cultivated fields (e.g., barley), roadsides, sand pits, gravel pits, forest edges, and open forests. Ground open or shaded by bushes (e.g., Alnus, Salix); rather dry, gravelly soil mixed with very fine sand, covered with sparse vegetation (e.g., tiny moss). Nocturnal; sheltering during the day under stones, dead leaves, pieces of wood or cardboard, and in moss. Biology. Seasonality: February, April-November. Tenerals: mid-July to September. Adults found overwintering in meadows and wood clearings; under stones and in moss. Dispersal power. Wing-polymorphic: macropterous (capable of flight) to brachypterous (incapable of flight). Frequent flier (to artificial lights at night). Frequent in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pitfall trapping; lifting small stones; light trapping; turning drift material. References. Lindroth, 1945, 1955a, 1963a, 1963b and 1985 (natural history); Pearson, 1964 (dispersal power); Luff, 1993 (biology). Bembidion (Plataphus) gratiosum Casey, 1918 Ecology. Uplands and subalpine zone. Banks of cold streams (mostly); lake and pond margins. Open ground; moist soil. Biology. Seasonality: May-August, October. Tenerals: late July to late August. Dispersal power. Macropterous, probably capable of flight. Moderate runner. References. Lindroth, 1963b and 1966 (natural history). Bembidion hageni Hayward, 1897 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion (Plataphodes) haruspex Casey, 1918 Ecology. Mountains, subalpine and alpine zones. Margins of small forest brooks. Associated with cold, swift water. More or less shaded ground; moist soil. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Lindroth, 1963b (natural history).

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Bembidion (Blepharoplataphus) hastii C.R. Sahlberg, 1827 Ecology. Lowlands and mountains. River banks. Shores of lakes and pools (in the tundra). Moist, gravelly, bare soil. Biology. Seasonality: June-August. Tenerals: mid-July (BC). Dispersal power. Macropterous. Flight records (Europe). Moderate runner. Collecting technique. Raking the loose gravel. References. Lindroth, 1945, 1963b, and 1985 (natural history). Bembidion (Bracteon) hesperium Fall, 1910 Ecology. Lowlands and mountains. River banks. Close to the water edge. Open ground; moist, sandy soil, bare or occasionally covered with sparse vegetation. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days in soil cracks or burrows dug in the ground. Biology. Seasonality: March, May-September. Defense-mechanism: when alarmed, the adult escapes by running and flying readily like a tiny Cicindela; the beetle is difficult to pursue; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; in the bright sunshine). Fast runner. Collecting technique. Hunting with a butterfly net. Reference. Maddison, 1993 (ecology). Bembidion (Pseudoperyphus) honestum Say, 1823 Ecology. Lowlands and mountains. Banks of river and brooks. Open ground; moist, gravelly, bare soil. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day among gravel or under stones. Gregarious. Biology. Seasonality: April-October, December. Tenerals: late July (NS), early August (VT). Defense-mechanism: when alarmed, the adult may escape by flying. Dispersal power. Macropterous. Frequent flier (when alarmed; to artificial lights in captivity). Moderate runner. Collecting techniques. Pouring water over the ground; raking the loose gravel. Reference. Lindroth, 1963b (natural history); Erwin, 1981 (natural history). Bembidion (Liocosmius) horni Hayward, 1897 Ecology. Brook margins. Moist, gravelly soil. Biology. Seasonality: June-July. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Bembidion (Hirmoplataphus) humboldtense Blaisdell, 1902 Ecology. An alpine meadow (found soon after snowmelt (CA)). Biology. Seasonality: April-July. Dispersal power. Macropterous, probably capable of flight. Bembidion (Plataphus) hyperboraeorum Munster, 1923 Ecology. Stream banks. Lake shores (Europe). Close to water. Moist, gravelly, bare soil. Nocturnal; sheltering during the day among gravel and stones. Biology. Seasonality: July-September. Dispersal power. Macropterous. Flight records (Europe). Moderate

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runner. Collecting technique. Raking the soil. References. Lindroth, 1945, 1963b and 1985 (natural history). Bembidion (Notaphus) idoneum Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion (Eupetedromus) immaturum Lindroth, 1954 Ecology. Lowlands and mountains. Borders of lakes, ponds, eutrophic marshes, and slow rivers; low forests. Ground open or shaded by bushes (e.g., Alnus, Salix); soft, wet, muddy soil covered with some vegetation (e.g., Carex). Mostly nocturnal; usually sheltering during the day under dead leaves, mats of dead plants, pieces of wood, and stones. Biology. Seasonality: February, April-September, November. Tenerals: midMay to November; mostly from mid-July on. Adults found overwintering on higher and drier ground, on hillocks and at the edge of woods; in leaf litter (mostly) and in the soil. Defense-mechanism: body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; to artifical lights in captivity). Moderate runner. Collecting techniques. Treading the soil with the feet; pouring water over the ground; light trapping. References. Lindroth, 1955a, 1963a, and 1963b (natural history). Bembidion impotens Casey, 1918 Ecology. Lowlands and mountains. Borders of rivers, brooks, lakes, ponds, and pools. Open ground; wet, clayish or sandy soil, bare or covered with sparse vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under dead leaves, grass, and in cracks in the soil. Biology. Seasonality: January, March-October. Tenerals: late July (BC). Adults found overwintering at the edge of woods; in leaf litter. Predators: frogs. Defense-mechanism: body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in drift material, which indicates flight ability. Moderate runner. Clearly effective as a colonist: quick to establish in defaunated streams. Collecting techniques. Treading the soil with the feet; pouring water over the ground; light trapping. References. Lindroth, 1963b (natural history); Kirk, 1975c (ecology, dispersal power); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Bembidion (Plataphodes) improvidens Casey, 1924 Ecology. Mountains, subalpine and alpine zones. Alpine meadows. Open ground; moist, soil covered with some vegetation (e.g., grass). Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Moderate runner.

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Bembidion (Bracteon) inaequale Say, 1823 Ecology. Lowlands and mountains. Banks of rivers and brooks; edges of ponds and pools (including temporary ones, e.g., in sand pits and gravel pits); lake shores. Open ground; wet soil, clay soil (often sand-mixed), bare or covered with sparse vegetation. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days in soil cracks and burrows dug in the ground, at the base of plants, under dead leaves, pieces of wood, and stones. Biology. Seasonality: March-October. Copulating pairs: May (QU). Tenerals: mid-July to September. Adults found overwintering on higher and drier ground, in the upper zone of river banks and lake shores; in the soil under dead leaves. Adult food, in the field: dying or dead arthropods. Phoretic mites found on adult body. Defense-mechanism: when alarmed, the adult escapes by running and flying readily like a tiny Cicindela; the beetle is difficult to pursue; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; in the bright sunshine; to artificial lights in captivity). Fast runner. Collecting techniques. Pouring water over the ground; hunting with a butterfly net. References. Lindroth, 1963b (natural history); Erwin, 1981 (natural history); Kavanaugh, 1992 (natural history); Maddison, 1993 (ecology). Bembidion (Eupetedromus) incrematum LeConte, 1860 Ecology. Lowlands and mountains. Marshy borders of pools, ponds, lakes, reservoirs, slow rivers, and brooks; swamps and mud flats. More or less shaded ground; wet, muddy soil, mixed with organic debris (e.g., dead leaves and twigs) and covered with sparse vegetation. Mostly nocturnal; sheltering during the day under dead leaves, dead grass, pieces of wood, and at the base of plants; sometimes active on cloudy days. Biology. Seasonality: May-November. Tenerals: early July to late August; mostly from mid-July on. Adults found overwintering on higher and drier ground, at the edge of woods and on hillocks; under dead leaves. Predators: frogs. Defense-mechanism: body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (to artificial lights in captivity). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Treading the soil with the feet; pouring water over the ground. References. Lindroth, 1955a, 1963a, and 1963b (natural history); Kavanaugh, 1992 (natural history). Bembidion (Notaphus) indistinctum Dejean, 1831 Ecology. Coastal lowlands. Seashore, salt marshes, and seaside lagoons. Open ground; wet, clayish saline soil covered with some Salicornia-plants. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under vegetal debris. Biology. Seasonality: January, April-August, November. Tenerals: late May (BC). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. References. Lindroth, 1963b (natural history); Kavanaugh, 1992 (natural history).

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Bembidion (Notaphus) insulatum (LeConte, 1852) Ecology. Uplands. Borders of alkaline ponds, lakes, and sloughs; salt meadows; salt flats in cultivated fields (e.g., corn). Open ground; moist, clayish, alkaline or saline soil covered with sparse vegetation (e.g., Salicornia). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in soil cracks or burrows dug in the ground. Biology. Seasonality: May-November. Tenerals: early August (MB, WY). Predators: avocets, gadwalls, and stilts. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet. References. Lindroth, 1963b (natural history); Kirk, 1975c (ecology). Bembidion (Pseudoperyphus) integrum Casey, 1918 Ecology. Mountains. Biology. Seasonality: August. Dispersal power. Unknown. Bembidion (Notaphus) intermedium (Kirby, 1837) Ecology. Lowlands and mountains. River banks (mostly); shores of lakes, ponds, and reservoirs; edges of pools in sand pits and gravel pits. Open ground; moist or wet soil consisting of clay or clay-mixed sand, bare or covered with sparse vegetation (e.g., Carex, Juncus). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under dead leaves and at the base of plants. Biology. Seasonality: February, AprilDecember. Tenerals: late June to mid August. Adults found overwintering on higher and drier ground, at the edge of woods, in the upper zone of sand pits and gravel pits; in leaf litter, under stones and pieces of wood. Predators: ducks. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. References. Blatchley, 1910 (ecology, biology); Lindroth, 1963b and 1971a (natural history); Kirk, 1975c (ecology, dispersal power). Bembidion (Eurytrachelus) interventor Lindroth, 1963 Ecology. Lowlands and mountains. Banks of large rivers. At some distance from water. Open ground; sandy soil, mixed with clay, covered with some vegetation. Mostly nocturnal; sometimes active in the bright sunshine around the lower limit of the scarce vegetation (e.g., Salix); usually sheltering during the day in soil cracks and burrows dug in the ground, and under twigs and fallen branches. Biology. Seasonality: April-August, October-November. Tenerals: late July to early August (AB, SA). Defense-mechanism: when alarmed, the adult escapes by flying readily. Dispersal power. Wing-dimorphic: macropterous (capable of flight) or brachypterous (incapable of flight). Occasional flier (when alarmed; to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Fast runner. Collecting techniques. Pitfall trapping; pouring water over the ground. References. Lindroth, 1963b (natural history); Ball and Bousquet, 2001 (taxonomy).

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Bembidion (Trechonepha) iridescens (LeConte, 1852) Ecology. Lowlands and mountains. Vicinity of trickles and small brooks with cold water. Sometimes at a considerable distance from water. Shaded ground; moist soil consisting of silt or organic debris. Nocturnal; sheltering during the day among moss, in leaf litter, and under stones. Biology. Seasonality: February-December. Tenerals: late June (CA), late July (BC). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. References. Lindroth, 1963b (natural history); Kavanaugh, 1992 (natural history). Bembidion (Notaphus) jacobianum Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion (Bembidion) jucundum G.H. Horn, 1895 Natural History. Unknown. Bembidion (Plataphus) kalumae Lindroth, 1963 Ecology. Alpine zone. Banks of small streams originating from glaciers. Associated with milky-like cold water. Moist, gravelly, bare soil. Biology. Seasonality: June-July. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Lindroth, 1963b (natural history). Bembidion kuprianovi Mannerheim, 1843 Ecology. Lowlands and mountains. Margins of running waters, e.g., small swift streams. Open ground; moist, gravelly-stony, bare soil. Nocturnal; sheltering during the day among gravel and under stones. Biology. Seasonality: April, June-August. Tenerals: late July (BC), early August (AK). Dispersal power. Macropterous. Occasional flier (into rotary traps). Moderate runner. Collecting techniques. Raking the soil; using rotary traps. Reference. Lindroth, 1963b (natural history). Bembidion (Actedium) lachnophoroides Darlington, 1926 Ecology. River banks. Close to water. Open ground; wet, bare soil composed of rather coarse sand. Nocturnal; sheltering during the day in the soil. Biology. Seasonality: July, September. Tenerals: early August (AB). Defense-mechanism: body color-pattern matching the soil. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pouring water on the ground. Reference. Lindroth, 1963b (natural history).

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Bembidion (Peryphanes) lacunarium (Zimmermann, 1869) Ecology. Lowlands, mountains, subalpine and alpine zones. Margins of cold, swift rivulets, brooks, and rivers (in forests); lake shores; wet forested places, at a certain distance from water (high elevation; WV); alpine meadows. Caves (frequently). Shaded ground; moist, gravelly, stony or rocky, bare soil. Nocturnal; sheltering during the day under stones, among gravel, under dead leaves and moss. Biology. Seasonality: AprilOctober. Tenerals: early July (VA) to early September (KY). Adults found overwintering in the upper zone of stream banks; under dead leaves. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous (probably capable of flight). Moderate runner. Collecting technique. Turning stones. References. Lindroth, 1963b (natural history); Erwin, 1981(natural history); Will et al. 1995 (ecology, dispersal power). Bembidion (Metallina) lampros (Herbst, 1784) Ecology. Lowlands. Cultivated fields, abandoned fields, vacant lots, gardens, parks, backyards. Greenhouses (occasionally). Open ground; moderately dry, clayish or gravelly soil covered with sparse low vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day. Biology. Seasonality: March-July, November. Oviposition: April to early June (NF). Tenerals: early to late June (NF). Adults found overwintering in ploughed fields (North America); in hedges (Europe). Life span: a year or more (Europe). Adult food, in the field (Europe): small arthropods, insect eggs, and seedlings of trees and crucifers. Adults are aggressive and cannibalistic. Economic importance (Europe): may play an important role in the control of of the Cabbage Root Fly; sometimes a pest of forest seedlings. Defense-mechanism: when alarmed, the adult may slowly emit a discharge from the pygidial glands. Dispersal power. Wing-polymorphic: macropterous (capable of flight) to brachypterous (incapable of flight). Flight records (Europe). Fast runner. Occasional climber (on plants; Europe). Clearly effective as a colonist: quick to establish in recently modified habitats. Strongly favored by human activities. Collecting techniques. Pitfall trapping; using window traps; sweeping plants. References. Lindroth, 1945, 1963a, and 1963b (natural history); Mitchell, 1963a (biology) and 1963b (natural history); Finlayson and Campbell, 1976 (biology); Sharova, 1981 (ecology, biology); Langor and Larson, 1983 (natural history); Lindroth, 1985 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology); Ball and Bousquet, 2001 (taxonomy). Bembidion (Bracteon) lapponicum Zetterstedt, 1828 Ecology. Lowlands and uplands. Banks of large rivers; lake shores. Open ground; wet, bare sandy soil, sometimes clay-mixed. Mostly diurnal; active in the sunshine; sheltering on hot days in rather dense vegetation or, on cloudy or cool days, in soil cracks and burrows dug in the ground. Biology. Seasonality: May-September. Tenerals: September (NT). Defense-mechanism: when alarmed, the adult escapes by running and flying readily like a tiny Cicindela; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; in the bright sunshine). Fast runner. Collecting techniques.

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Pouring water over the ground; hunting with a butterfly net. References. Lindroth, 1963b and 1985 (natural history); Maddison, 1993 (ecology); Luff, 1993 (biology). Bembidion (Notaphus) latebricola Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion (Lymneops) laticeps (LeConte, 1858) Ecology. Coastal lowlands. Seashore: intertidal zone. Open ground; wet, rocky, saline soil. Nocturnal; sheltering in airbells among narrow rifts of fissured rocks covered at high tide. Biology and Dispersal power. Unknown. Reference. Cooper, 1976 (ecology). Bembidion (Plataphodes) laxatum Casey, 1918 Ecology. Mountains, subalpine and alpine zones. Margins of brooks and snowfields. Open ground; moist soil. Biology. Seasonality: July-September. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Bembidion lecontei Csiki, 1928 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion lenae Csiki, 1928 Ecology. Inundation areas of river banks. At considerable distance from water. Ground shaded by bushes (e.g., Alnus); rather dry soil composed of fine sand, bare or covered with sparse vegetation (e.g., Equisetum). Nocturnal; sheltering during the day in leaf litter and under stones. Biology. Seasonality: June-August. Tenerals: early August (AK). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Turning debris. Reference. Lindroth, 1963b (natural history). Bembidion (Bracteon) levettei Casey, 1918 Ecology. Lowlands and mountains. Lake shores; pond margins (even temporary ones; often in sand pits); banks of rivers and brooks. Open ground; moderately moist soil consisting of sand or clay-mixed sand, bare or covered with sparse vegetation. Mostly diurnal; active in the bright sunshine; sheltering on cloudy or cool days in soil cracks, burrows dug in the ground, and under pieces of wood. Biology. Seasonality: AprilNovember. Tenerals: early July (MI) to August (QU). Adults found overwintering on higher and drier ground, on the top of sand dunes; in rotten logs. Defense-mechanism: when alarmed, the adult escapes by running and flying readily like a tiny Cicindela; the beetle is difficult to pursue; body color-pattern matching the soil. Dispersal power.

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Macropterous. Frequent flier (when alarmed; in the bright sunshine; to artificial lights in captivity). Moderately favored by human activities. Collecting techniques. Pouring water over the ground; hunting with a butterfly net. References. Lindroth, 1963a and 1963b (natural history); Maddison, 1993 (ecology).. Bembidion (Hydrium) levigatum Say, 1823 Ecology. Lowlands and mountains. River banks. Open ground; moist or wet, sandy or sandy-clayish soil, bare or covered with sparse vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in cracks in the soil and under stones. Biology. Seasonality: March-October. Tenerals: July-August; mostly from mid-July on. Adults found overwintering on higher and drier ground, on hilly slopes; under stones. Defense-mechanism: when alarmed, the adult escapes by running and flying readily; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; in the bright sunshine; to artificial lights at night). Fast runner. Collecting techniques. Pouring water over the ground; treading the soil with feet; hunting with a butterfly net. References. Blatchley, 1910 (ecology); Everly and Everly, 1936 (ecology); Kirk, 1975c (dispersal power); Erwin, 1981 (natural history). Bembidion (Lionepha) lindrothellus Erwin and Kavanaugh, 1981 Ecology. Mountains. Vicinity of a stream, on open ground with dry, gravelly soil. Nocturnal; sheltering under dandelion rosettes (Taraxacum). Biology. Seasonality: June, August. Tenerals: June (AK). Dispersal power. Wing-dimorphic: macropterous (probably capable of flight) or brachypterous (incapable of flight). Moderate runner. Bembidion (Bracteon) lorquinii Chaudoir, 1868 Ecology. Lowlands and mountains. River banks. Open ground; moist, bare sand. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days in soil cracks and burrows dug in the ground. Biology. Seasonality: April-July. Defense-mechanism: when alarmed, the adult escapes by flying readily. Dispersal power. Macropterous. Frequent flier (when alarmed; in the bright sunshine). Fast runner. Collecting techniques. Pouring water over the ground; hunting with a butterfly net. Reference. Maddison, 1993 (ecology). Bembidion luculentum Casey, 1918 Ecology. Coastal lowlands. Salt marshes. Open ground; moist, saline soil. Biology. Seasonality: June. Dispersal power. Wing-condition unknown. Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Bembidion (Lionepha) lummi Erwin and Kavanaugh, 1981 Ecology. Lowlands and mountains. Brook margins. Biology. Seasonality: April-May, JulyAugust. Dispersal power. Macropterous, probably capable of flight. Moderate runner.

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Bembidion (Plataphodes) manningense Lindroth, 1969 Ecology. Mountains. River banks. Shaded ground. Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: May-September. Tenerals: August (BC, NT). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Lindroth, 1963b (natural history; as ochropus Lindroth). Bembidion (Hydriomicrus) marinianum Casey, 1924 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion (Terminophanes) mckinleyi carneum Lindroth, 1963 Ecology. Mountains. Banks of cold rivers and brooks, often originating from glaciers. Open ground; moist, gravelly, bare soil. Biology. Seasonality: May-August. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Raking the loose gravel. Reference. Müller-Motzfeld, 1998 (taxonomy). Bembidion (Terminophanes) mckinleyi mckinleyi Fall, 1926 Ecology. Mountains. Banks of rivers and brooks (often originating from glaciers). Associated with cold water. Open ground; moist, gravelly, bare soil. Biology. Seasonality: June-August. Tenerals: mid-June (AK). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Raking the loose gravel; pouring water over the ground. References. Lindroth, 1963b (natural history); MüllerMotzfeld, 1998 (taxonomy). Bembidion mexicanum Dejean, 1831 Ecology. Uplands. Stream margins. Open ground; moist, gravelly soil mixed with some silt, bare or covered with sparse vegetation. Biology. Seasonality: May-August. Tenerals: late May (AZ). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Bembidion mimus Hayward, 1897 Ecology. Lowlands and mountains. Borders of ponds, eutrophic marshes, bogs, swamps, fens, pools (in gravel pits), slow rivers, brooks, and irrigation canals; wet meadows, and cultivated fields (e.g., carrot). Open or half-shaded ground; moist, peaty (mostly), clayish or sandy soil, rich in organic debris, with bare spots surrounded by sparse vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under dead leaves, dead plants, pieces of wood, small stones, heaps of hay, in cracks in the soil , and at the base of plants. Biology. Seasonality: January, March-November. Tenerals: mid-July and late August. Adults found overwintering on higher and drier ground, in the upper zone of sand pits, hillocks, and hills, in vacant lots, mead-

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ows, pastures, swamps, field hedges, at the edge of woods, and in woods; under dead leaves, moss, dead fallen plants, and stones. Predators: frogs. Defense-mechanism: body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (in late afternoon; at sunset; to artificial lights at night). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Moderately favored by human activities. Collecting techniques. Pouring water over the ground; treading the soil with the feet. References. Lindroth, 1955a, 1963a, and 1963b (natural history); Kirk, 1975c (ecology, dispersal power). Bembidion modocianum Casey, 1924 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion mormon Hayward, 1897 Ecology. Uplands. Margins of alkaline lakes, ponds, and slow rivers. Open ground; moist, clayish, alkaline soil, bare or covered with sparse vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in cracks in the soil and at the base of plants. Biology. Seasonality: May-September. Tenerals: late July (UT). Dispersal power. Macropterous. Frequent flier (when alarmed; to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. References. Hatch, 1953 (ecology); Kirk, 1975c (dispersal power). Bembidion (Semicampa) morulum LeConte, 1863 Ecology. Coastal lowlands. Borders of a small brook in the tundra, on open ground with moderately moist, peaty soil. Biology. Seasonality: May-August. Tenerals: early and mid-August (MB). Dispersal power. Wing-dimorphic: macropterous (capable of flight) or brachypterous (incapable of flight). Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. References. Lindroth, 1963a and 1963b (natural history). Bembidion (Liocosmius) mundum (LeConte, 1852) Ecology. Lowlands and mountains. Banks of rivers and brooks. Open or shaded ground; wet, sandy soil, bare or almost so. Nocturnal; sheltering during the day under dead leaves, dead fallen plants, and stones. Gregarious. Biology. Seasonality: May-August, October. Tenerals: mid-July (WA). Dispersal power. Macropterous, capable of flight. Moderate runner. Collecting techniques. Pouring water oder the ground; treading the soil with the feet. Reference. Lindroth, 1963b (natural history). Bembidion (Semicampa) muscicola Hayward, 1897 Ecology. Lowlands and mountains. Moist deciduous forests (along water bodies); swamp-forests, swamps, Sphagnum-bogs, eutrophic marshes; marshy borders of ponds,

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small lakes, slow rivers and brooks. Shaded ground; firm, moist, clayish or muddy soil, rich in organic debris, covered with some vegetation (e.g., Carex). Nocturnal; sheltering during the day under dead leaves, dead grass, and in moss. Biology. Seasonality: January-December. Tenerals: April-May, mid-July to August, October. Adults found overwintering on higher and drier ground, in the upper zone of marshes and lake shores, in woods, at the edge of woods, and in hawthorn hedges; under dead leaves, the loose bark of standing rotten tree-stumps, in clumps of moss and grass, under small stones, and in the soil. Adult food, in the field: mosquito eggs. Dispersal power. Wing-dimorphic: macropterous (capable of flight) or brachypterous (incapable of flight). Occasional flier (to artificial lights in captivity). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on tree-stumps). Collecting techniques. Pouring water over the ground; raking the leaf litter. Reference. Lindroth, 1963b (natural history). Bembidion (Bembidion) mutatum Gemminger and Harold, 1868 Ecology. Lowlands and mountains. Moraines, vacant lots, meadows, pastures, cultivated fields (e.g., wheat, barley), roadsides, sandhills, sand pits, gravel pits, and vineyards. Open ground; moderately moist soil consisting of fine sand, with organic debris, covered with tiny moss or very sparse vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under stones, pieces of wood, and in moss. Biology. Seasonality: March-November. Gravids: mid-July (QU) and early August (AB). Adults found overwintering on higher and drier ground, in woods; under the loose bark of standing tree-stumps. Adult food: dipterous pupae (in the field); lepidopterous eggs (in captivity). Dispersal power. Wing-dimorphic: macropterous (mostly; capable of flight) or brachypterous (incapable of flight). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants, vines, and trees). Collecting techniques. Pitfall trapping; turning stones; sweeping plants. References. Lindroth, 1955a and 1963b (natural history); Frank, 1971b (biology). Bembidion nebraskense LeConte, 1863 Ecology. Mountains. Stream banks. At some distance from water. Ground half-shaded by bushes (e.g., Alnus); rather dry soil covered with sparse vegetation (e.g., Equisetum). Nocturnal; sheltering during the day in cracks in the soil. Biology. Seasonality: MaySeptember. Tenerals: early summer (BC), early July (ID), early August (WY). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Reference. Lindroth, 1963b (natural history). Bembidion neocoerulescens Bousquet, 1993 Ecology. Stream margins. Biology. Seasonality: December. Dispersal power. Macropterous, probably capable of flight. Moderate runner.

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Bembidion nevadense Ulke, 1875 Ecology. Mountains. Borders of small streams. More or less shaded ground; moist soil covered with some vegetation (e.g., moss). Nocturnal; sheltering during the day usually among moss. Biology. Seasonality: June-September. Tenerals: late May (BC). Dispersal power. Wing-dimorphic: macropterous (probably capable of flight) or brachypterous (capable of flight). Moderate runner. Collecting technique. Pouring water over the ground. Reference. Lindroth, 1963b (natural history). Bembidion (Notaphus) nigripes (Kirby, 1837) Ecology. Lowlands and mountains. Seashore, salt meadows, and alkali sloughs; margins of eutrophic marshes, lakes, and slow brooks and rivers. Open ground; rather firm, moist or wet, clay soil, often saline or alkaline, covered with sparse vegetation (e.g., grass). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under algal carpets, dead plants, stones, logs, pieces of wood, and in cracks in the soil. Biology. Seasonality: March-October. Copulating pairs: early July (ID). Tenerals: late June to early August. Predators: ducks. Defense-mechanism: body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; to artifical lights in captivity). Moderate runner. Collecting techniques. Lifting algal carpets; pouring water over the ground. References. Lindroth, 1955a, 1963a, and 1963b (natural history); Kirk, 1975c (ecology, dispersal power). Bembidion nigrocoeruleum Hayward, 1897 Ecology. Unknown. Biology. Seasonality: May. Dispersal power. Macropterous, probably capable of flight. Bembidion (Lymneops) nigropiceum (Marsham, 1802) Ecology. Seashore (Europe): close to the high tide mark; open ground; wet, rocky or sandy, saline soil; sheltering under stones. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Reference. Lindroth, 1974 (ecology, dispersal power). Comments. Doubtfully established in North America. Bembidion (Hirmoplataphus) nigrum Say, 1823 Ecology. Lowlands and mountains. Banks of small rivers and brooks. Open ground; moist, bare soil composed of gravel lying on a mixture of sand and clay. Nocturnal; sheltering during the day among gravel, under stones and dead leaves, and in cracks in the soil. Gregarious. Biology. Seasonality: April-October. Tenerals: late May (QU) to early September (MA); mostly from mid-July on. Adults found overwintering on higher and drier ground, in sand pits; under dead leaves. Predators: frogs. Defense-mechanism: when alarmed, the adult may emit a Brachinus-like spray from the pygidial glands; body color matching the soil. Dispersal power. Macropterous. Frequent flier (to artifi-

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cial lights in captivity). Moderate runner. Collecting techniques. Pouring water over the ground; raking the loose gravel. References. Lindroth, 1963b (natural history); Larochelle, 1974a (ecology); Erwin, 1981 (natural history). Bembidion (Eurytrachelus) nitidum (Kirby, 1837) Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, vacant lots, meadows, grasslands, cultivated fields (e.g., corn, oat, flax, wheat, barley, cabbage), roadsides, sand pits, and gravel pits. Open ground; dry sandy soil covered with sparse or moderate vegetation (e.g., Artemisia, grass). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under stones and dead leaves (mostly), plant rosettes (Verbascum), and heaps of hay. Biology. Seasonality: March-November. Tenerals: JulyAugust. Adults found overwintering in sand pits, meadows, vacant lots, at the edge of woods, and in woods; under stones, dead leaves, plant rosettes (Verbascum), and in the soil (as deep as 10 cm). Adult food: dipterous larvae (in the field); dipterous eggs, earthworms, meat, and a corn kernel (in captivity). Predators: toads. Parasites: nematodes. Defense-mechanism: when alarmed, the adult escapes by running. Dispersal power. Macropterous. Occasional flier (on cloudy days after rain; to artificial lights in captivity). Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; raking the leaf litter. References. Lindroth, 1963b (natural history); Frank, 1971b (biology); Kirk, 1975c (ecology); Ball and Bousquet, 2001 (taxonomy). Bembidion nogalesium Casey, 1924 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion (Notaphus) nubiculosum Chaudoir, 1868 Ecology. Uplands. River banks. Biology. Seasonality: January, April, June, August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Bembidion nudipenne Lindroth, 1963 Ecology. Uplands. Cultivated fields (e.g., alfalfa, wheat, barley, turnip), vacant lots, and pastures. Open ground; rather dry, sandy soil covered with sparse vegetation. Nocturnal; sheltering during the day in cracks in the soil. Biology. Seasonality: April-September. Gravids: late September (AB). Adult food, in the field: dipterous larvae. Dispersal power. Macropterous. Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strongly favored by human activities. Collecting technique. Pitfall trapping. Reference. Lindroth, 1963b (natural history). Bembidion oberthueri Hayward, 1901 Ecology. Lowlands and mountains. Banks of rivers (mostly) and brooks; pond margins (close to rivers). Open ground; moist or wet soil composed of sand-mixed clay, bare or

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covered with sparse vegetation (e.g., Juncus, Equisetum). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in the soil and under stones. Biology. Seasonality: April-October. Tenerals: July-October; mostly from mid-July on. Adults found overwintering on higher and drier ground, at the edge of woods; under dead leaves. Predators: toads. Defense-mechanism: body color matching the soil. Dispersal power. Macropterous. Frequent flier (to artificial lights in captivity; sometimes at sunset). Moderate runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Lindroth, 1963b (natural history). Bembidion (Eurytrachelus) obliquulum LeConte, 1859 Ecology. River margins. Biology. Seasonality: January-February, May-June. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Ball and Bousquet, 2001 (taxonomy). Bembidion (Amerizus) oblonguloides Lindroth, 1963 Ecology. Lowlands and mountains. Coniferous and mixed forests: often along seepages and rivulets. Shaded ground; moist soil, rich in organic debris, moss, and stones. Nocturnal; sheltering during the day deep under stones, logs, and in clumps of moss. Biology. Seasonality: June-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning stones. References. Lindroth, 1963b (natural history); Kavanaugh, 1992 (natural history). Bembidion (Amerizus) oblongulum (Mannerheim, 1852) Ecology. Lowlands and uplands. Coniferous forests. Shaded ground; soil rich in fine, dark organic debris. Nocturnal; sheltering during the day under well-embedded stones. Biology. Seasonality: April-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting technique. Turning well-embedded stones. References. Lindroth, 1963b (natural history); Kavanaugh, 1992 (natural history). Bembidion (Peryphus) obscurellum (Motschulsky, 1845) Ecology. Lowlands and mountains. River banks (drier parts), pond margins, shores of lakes and reservoirs; edges of cattle water holes; salt flats, cultivated fields (e.g., wheat, barley), meadows, abandoned fields, rubbish dumps, playgrounds, sand pits, sand dunes, gravel pits, and roadsides. Open ground; moist or wet, sandy, clayish or muddy soil covered with sparse vegetation. Nocturnal; sheltering during the day under stones, pieces of wood, dead plants, and in the soil (e.g., between clumped grass roots). Biology. Seasonality: February, April-November. Copulating pairs: May (AB), June (SA), late July (QU). Gravids: late May (AB). Tenerals: early June to early August. Adults found overwintering on higher and drier ground, on hillocks; under stones, dead leaves, and in rotten logs. Adult food: dipterous larvae and adult coccinellids (in the field); dipterous eggs (in captivity). Dispersal power. Macropterous. Frequent flier (often swarming

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by the hundreds to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strongly favored by human activities. Clearly effective as a colonist: quick to establish in recently modified habitats. Collecting techniques. Pouring water over the ground; treading the soil with the feet; pitfall trapping; light trapping. References. Lindroth, 1963b and 1971a (natural history); Bell, 1971 (ecology); Frank, 1971b (biology); Kirk, 1975c (ecology, dispersal power); Finlayson and Campbell, 1976 (biology); Doane, 1981 (ecology, biology); Will et al. 1995 (dispersal power). Bembidion obscuripenne Blaisdell, 1902 Ecology. Mountains, subalpine and alpine zones. An alpine meadow (found after snowmelt), on wet soil. Biology. Seasonality: July, September. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Bembidion (Notaphus) obscuromaculatum (Motschulsky, 1859) Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion obtusangulum LeConte, 1863 Ecology. Uplands. Borders of saline lakes, ponds, and sloughs. At some distance from water. Open ground; moist, clayish saline soil covered with some vegetation (e.g., Spergularia, Salicornia, Atriplex). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in cracks in the soil. Biology. Seasonality: March-August. Tenerals: mid-July to early August (MB, SA). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Reference. Lindroth, 1963b (natural history). Bembidion obtusidens Fall, 1922 Ecology. Uplands. Borders of saline lakes, ponds, sloughs, marshes, and slow streams. Open ground; moist, clayish saline soil, bare or covered with sparse vegetation (e.g., Distichlis, Atriplex). Attracted to volatile metabolites emitted by filamentous blue-green algae. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in cracks in the soil. Biology. Seasonality: May-August. Tenerals: mid-July (SA) to early August (WY). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. References. Lindroth, 1963b (natural history); Evans, 1982 (ecology). Bembidion (Phyla) obtusum Audinet-Serville, 1821 Ecology. Lowlands and mountains. Vacant lots, pastures, meadows, cultivated fields (e.g., onion, alfalfa, cabbage), hills, roadsides, sand pits, gravel pits, edges of seepages, irrigation canals, brooks, and rivers; forest edges and open forests. Open or slightly

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shaded ground; moderately dry, clay soil covered with sparse or moderate vegetation. Mostly nocturnal; sometimes active on roads in the spring sunshine; usually sheltering during the day under stones, dead leaves, heaps of hay, in cracks in the soil, and in moss. Gregarious (in winter). Biology. Seasonality: January, March-December. Tenerals: March-June, August-November; mostly from August on. Adults found overwintering on higher and drier ground, in fields, at the edge of woods, and in woods; in leaf litter (mostly) and in moss. Dispersal power. Brachypterous, incapable of flight (North America). Europe: wing-dimorphic: macropterous (probable capable of flight) or brachypterous (incapable of flight). Occasional flier (to artificial lights in captivity). Moderate runner. Occasional climber (on plants and trees). Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Raking the leaf litter; pitfall trapping; sweeping plants. References. Lindroth, 1945, 1963b, and 1985 (natural history); Koch, 1989 (ecology); Hoebeke et al. 1991 (natural history). Bembidion (Plataphodes) occultator Notman, 1919 Ecology. Lowlands and mountains. Wooded areas: along rivers, brooks, rills, eutrophic marshes, lakes, and ponds; swamps. More or less shaded ground, under bushes (e.g., Alnus, Salix, Myrica); rather firm, wet sandy or clayish soil, often mixed with organic debris, covered with low sparse vegetation. Nocturnal; sheltering during the day in leaf litter (mostly), under dead plants, and logs. Gregarious. Biology. Seasonality: May-September. Tenerals: mid-July to early August. Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Occasional flier (to artificial lights in captivity). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Collecting technique. Raking the leaf litter. References. Lindroth, 1955a, 1963a, and 1963b (natural history). Bembidion (Notaphus) operosum Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion (Plataphus) oppressum Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion (Bembidion) oregonense Hatch, 1953 Natural History. Unknown. Bembidion (Lionepha) osculans Casey, 1918 Ecology. Lowlands and mountains. Margins of cold, small streams. Shaded ground; moist soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: March-December. Tenerals: March-August. Dispersal power. Macropterous, probably capable of flight. Moderate runner.

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Bembidion (Cillenus) palosverdes Kavanaugh and Erwin, 1992 Ecology. Coastal lowlands. Seashore: intertidal zone. Open ground; wet, rocky soil. Nocturnal; sheltering during the day in burrows dug in the soil and under stones. Biology. Seasonality: June. Tenerals: June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Reference. Kavanaugh and Erwin, 1992 (natural history). Bembidion (Notaphus) patruele Dejean, 1831 Ecology. Lowlands and mountains. Borders of pools (often small temporary ones), ponds, lakes, reservoirs, marshes, slow rivers, brooks, irrigation canals, and roadside ditches; swamps, mud flats, wet pastures, and meadows. Open or shaded ground; wet, often clayish or muddy soil, rich in organic debris, covered with sparse vegetation (e.g., Agrostis, Carex). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in leaf litter, under mats of dead grass, small stones, pieces of wood, mud cakes, in cracks in the soil, and at the base of plants. Biology. Seasonality: JanuaryDecember. Copulating pairs: mid-July (PE). Tenerals: late June-September. Adults found overwintering on higher and drier ground, at the edge of marshes and woods, in swampforests, woods, and in the upper zone of sand pits, on hills, in pastures and vacant lots; in leaf litter (mostly), under small stones, in clumps of moss and grass, under the loose bark of fallen trees, and in the soil. Predators: frogs and toads. Dispersal power. Macropterous. Frequent flier (at sunset; to artificial lights at night; to atificial lights in captivity). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants, e.g., Equisetum, Epilobium, Gramineae). Strongly favored by human activities. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping; turning drift material; using an aspirator; sweeping plants. References. Lindroth, 1955a (as posticum Haldeman), 1963a, and 1963 b (natural history); Kirk, 1969 (dispersal power) and 1975c (ecology, dispersal power); Will et al. 1995 (dispersal power). Bembidion (Bembidion) pedicellatum LeConte, 1857 Ecology. Lowlands. Open forests and their vicinity: roadsides, adjoining pastures, and borders of small streams. Open ground; moderately dry, sandy soil. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under dead leaves, other organic debris, stones, and on the top of ant nests. Biology. Seasonality: FebruaryNovember. Tenerals: early July (KY). Adults found overwintering in woods; under the loose bark of logs and tree-stumps. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. Reference. Erwin, 1981 (natural history). Bembidion perbrevicolle Casey, 1924 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight.

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Bembidion pernotum Casey, 1918 Ecology. Uplands. Biology and Dispersal power. Unknown. Bembidion (Peryphus) petrosum attuense Lindroth, 1963 Ecology. Coastal lowlands. Banks of small brooks (mostly) and margins of pools close to the sea. Open ground; moist, stony-gravelly, bare soil. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Bembidion (Peryphus) petrosum petrosum Gebler, 1833 Ecology. Lowlands and mountains. Banks of large rivers (mostly); shores of large lakes; seashore. Open ground; moist or wet soil consisting of sand, often mixed with stones, sometimes saline, bare or covered with sparse vegetation. Nocturnal; sheltering during the day under stones, pieces of wood, and dead leaves. Biology. Seasonality: AprilOctober. Copulating pairs: late May to late July (NB, QU). Tenerals: late June to late August; mostly from mid-July on. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; to artifical lights in captivity). Moderate runner. Collecting techniques. Turning stones; light trapping. References. Lindroth, 1955a, 1963a, 1963b and 1985 (natural history). Bembidion pilatei Chaudoir, 1868 Natural History. Unknown. Bembidion (Notaphus) pimanum Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion (Plataphodes) placeranum Casey, 1924 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion plagiatum (Zimmermann, 1869) Ecology. Lowlands and mountains. Banks of rivers and brooks. Open ground; wet or moist, gravelly, clayish or sandy, bare soil. Nocturnal; sheltering during the day among gravel, under stones and dead leaves. Biology. Seasonality: April-August. Tenerals: early to mid-July (OH, VA, WV). Dispersal power. Macropterous. Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting technique. Raking the soil and the leaf litter. Reference. Hoffman, 1981 (ecology).

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Bembidion (Plataphus) planatum (LeConte, 1848) Ecology. Lowlands and mountains. Banks of cold rivers and brooks. Close to water. Open ground; moist or wet, gravelly-stony, bare soil. Nocturnal; sheltering during the day under stones. Gregarious. Biology. Seasonality: February, April-October, December. Copulating pairs: early and mid-July (MT, QU). Tenerals: late June to late July. Defense-mechanism: when alarmed, the adult may escape by flying. Dispersal power. Macropterous. Frequent flier (when alarmed; to artificial lights in captivity). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Turning stones and drift material. References. Lindroth, 1955a, 1963a, and 1963b (natural history). Bembidion (Plataphus) planiusculum Mannerheim, 1843 Ecology. Lowlands, mountains, subalpine and alpine zones. Banks of running waters (e.g., small streams). Close to water. Open (mostly) or shaded ground; moist, stony and gravelly soil. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under stones. Biology. Seasonality: January, April-August. Tenerals: early August (AK). Dispersal power. Macropterous. Occasional flier (in the sunshine). Moderate runner. Collecting technique. Turning stones. References. Lindroth, 1963b (natural history); Kavanaugh, 1992 (natural history). Bembidion (Trichoplataphus) planum (Haldeman, 1843) Ecology. Lowlands and mountains. Banks of rivers and brooks. Island beaches (Lake Erie region). Close to water. Open ground; moist or wet, bare, gravelly soil, sometimes mixed with clay or sand. Nocturnal; sheltering during the day among gravel and under stones. Gregarious. Biology. Seasonality: April-October. Tenerals: early June to September. Adults found overwintering on higher and drier ground, in the upper zone of river banks and in woods; under dead leaves. Predators: frogs and toads. Defense-mechanism: when alarmed, the adult may escape by flying, hiding under stones or diving into the water (where it hides under stones). Difficult to capture when moving readily in the interstitial spaces of dolomitic gravel at the margin of the wave splash. Dispersal power. Macropterous. Frequent flier (when alarmed; to artificial lights at night; to artifical lights in captivity). Moderate runner. Collecting techniques. Pouring water over the ground; raking the soil. References. Lindroth, 1963b (natural history); Landry, 1976 (biology); Will et al. 1995 (ecology). Bembidion (Peryphanes) platynoides Hayward, 1897 Ecology. Mountains. Borders of running waters (e.g., small streams). Close to water. Slightly shaded ground; moist, sandy-gravelly, bare soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: January-December. Tenerals: early June (BC) to early August (WY). Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous (probably capable of flight). Moderate runner. Collecting technique. Turning stones. Reference. Lindroth, 1963b (natural history).

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Bembidion (Cyclolopha) poculare Bates, 1884 Ecology. Mountains. Borders of a dam (Mexico). Biology. Seasonality: July. Dispersal power. Unknown. Bembidion (Trepanedoris) politum (Motschulsky, 1845) Natural History. Unknown. Bembidion (Peryphus) poppii schalleri Lindroth, 1963 Ecology. Mountains. Inundation area of a river bank, on open ground with slightly moist, bare soil composed of very fine sand. Biology. Seasonality: June-August. Tenerals: early August (AK). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Reference. Lindroth, 1963b (natural history). Bembidion postremum Say, 1830 Ecology. Lowlands and low mountains. River banks. At some distance from water. Ground often half-shaded by Salix-bushes; steep, moderately moist, bare, sandy soil, mixed with silt. Nocturnal; sheltering during the day in cracks in the soil, under clods of sand, leaf litter, and pieces of wood. Biology. Seasonality: June-October. Tenerals: early June (QU) to mid-July NH); mostly June. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pouring water over the ground. Reference. Lindroth, 1963b (natural history). Bembidion (Bembidion) praecinctum LeConte, 1879 Ecology. Uplands. Upper banks of large pools, lakes, and sloughs. At some distance from water. Open ground; rather dry, gravelly or clayish soil. Biology. Seasonality: MayOctober. Dispersal power. Wing-dimorphic: macropterous (probably capable of flight) or brachypterous (incapable of flight). Moderate runner. Collecting technique. Pouring water over the ground. Bembidion (Semicampa) praticola Lindroth, 1963 Ecology. Lowlands and mountains. Moist deciduous forests (often close to small lakes and pools); swamps; wooded borders of eutrophic marshes and ponds. Shaded ground; moist, clayish or muddy soil covered with some vegetation (e.g., Carex, grass, moss) and leaf litter. Nocturnal; sheltering during the day under dead leaves and in moss. Biology. Seasonality: January, April-December. Tenerals: May (rarely; QU), July-September, November. Adults found overwintering on higher and drier ground, in swampy areas, in woods, at the edge of woods, and on hillocks; under dead leaves (mostly) and in clumps of moss and grass. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous (probably capable of flight). Moderate runner.

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Collecting techniques. Raking the leaf litter from the soil. Reference. Lindroth, 1963b (natural history). Bembidion (Metallina) properans (Stephens, 1827) Ecology. Lowlands. Meadows, grasslands, cultivated fields (e.g., hay), vacant lots, edges of fields, roadsides, sand pits, and open forests. Open ground; moderately moist, preferably clay soil, sometimes gravelly or sandy, covered with sparse or moderate vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under small stones, dead leaves, and pieces of wood. Semi-gregarious. Biology. Seasonality: May-August. Tenerals: mid-July (PE), early August (QU). Adult food, in the field: small insects. Larval food, in captivity (Europe): Drosophila-flies. Dispersal power. Macropterous. Flight records (Europe). Frequent in seashore drift material, which indicates flight ability. Fast runner. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning small stones; raking the leaf litter; turning drift material. References. Lindroth, 1945 and 1963b (natural history); Sharova, 1981 (ecology, biology); Lindroth, 1985 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology); Ball and Bousquet, 2001 (taxonomy). Bembidion (Trepanedoris) pseudocautum Lindroth, 1963 Ecology. Lowlands and mountains. Borders of eutrophic marshes and swamps (e.g., Typha, Salix); wooded margins of slow brooks and rivers; low forests. Ground shaded by shrubs and trees; wet, clay soil covered with dense vegetation (e.g., Typha, Solanum, Lysimachia). Nocturnal; sheltering during the day under dead leaves and dead fallen plants (Typha). Biology. Seasonality: March-December. Tenerals: April (rarely; ON), June (NY, QU), August (mostly). Adults found overwintering on higher and drier ground, at the edge of woods and in the upper zone of brook borders; in leaf litter. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Treading the vegetation into the water; pouring water over the ground. Reference. Lindroth, 1963b (natural history). Bembidion (Lionepha) pseudoerasum Lindroth, 1963 Ecology. Mountains. Biology. Seasonality: May-August. Dispersal power. Macropterous, probbly capable of flight. Bembidion (Bracteon) punctatostriatum Say, 1823 Ecology. Lowlands and uplands. Banks of large rivers (mostly) and lakes. Open ground; moist bare soil often composed of sand-mixed clay. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days in soil cracks, burrows dug in the ground, and under pieces of wood. Biology. Seasonality: March-October. Tenerals: August (QU, VT). Adults found overwintering on higher and drier ground, in the upper zone of river banks; in the sandy soil. Predators: toads. Defense-mechanism: when alarmed, the adult

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escapes by running and flying readily like a tiny Cicindela; the beetle is difficult to pursue; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; in the bright sunshine). Fast runner. Collecting techniques. Pouring water over the ground; hunting with a butterfly net. References. Lindroth, 1963b (natural history); Maddison, 1993 (ecology); Will et al. 1995 (dispersal power). Bembidion (Hydriomicrus) quadratulum Notman, 1919 Ecology. Lowlands and mountains. Sphagnum-bogs: along ponds and lakes. Close to water. Open ground; soft, wet, floating peaty soil covered with some vegetation (e.g., Sphagnum, Kalmia, Saracenia, Oxycoccus). Mostly nocturnal; sometimes active in the sunshine in Sphagnum-carpets and on bare patches of peat; usually sheltering during the day in moss carpets. Gregarious. Biology. Seasonality: May-August. Tenerals: late July to late August (NF). Dispersal power. Macropterous. Occasional flier (to artificial lights in captivity). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Treading floating Sphagnum-carpets and the peaty soil with the feet; pouring water over the ground. References. Lindroth, 1955a, 1963a, and 1963b (natural history); Landry, 1978 (ecology). Bembidion (Plataphodes) quadrifoveolatum Mannerheim, 1843 Ecology. Lowlands, mountains, subalpine and alpine zones. Stream banks, lake shores, pond margins, meadows, and vicinity of snowfields. Open (mostly) or shaded ground; moist, sandy-gravelly, bare soil. Nocturnal; sheltering during the day under stones and among gravel. Gregarious. Biology. Seasonality: May-August. Tenerals: early June (BC), late August (AK). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Raking the soil; pouring water over the ground. References. Lindroth, 1963b (natural history); Papp, 1978 (ecology); Kavanaugh, 1992 (natural history). Bembidion (Bembidion) quadrimaculatum dubitans (LeConte, 1852) Ecology. Lowlands and mountains. Cultivated fields, roadsides, and gardens; upper zone of lake shores, river banks, and pool margins. Open ground; moderately dry soil composed of fine sand or clay, bare or almost so. Biology. Seasonality: February, MayAugust. Adult food, in the field: dipterous eggs. Dispersal power. Wing-polymorphic: macropterous (probably capable of flight) to brachypterous (incapable of flight). Moderate runner. Collecting technique. Pitfall trapping. Bembidion (Bembidion) quadrimaculatum oppositum Say, 1823 Ecology. Lowlands and mountains. Cultivated fields (e.g., corn, wheat, barley, soybean, sugar beet, oat, flax, asparagus, dry bean, onion, alfalfa, potato, cabbage, carrot, raspberry, turnip), gardens, meadows, pastures, vacant lots, lawns, roadsides, sand pits, gravel pits, ski fields; upper zone of lake shores, river banks, and irrigation canals; or-

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chards (e.g., apple) and open forests. Open ground; moderately dry soil, usually composed of fine sand or clay, bare or almost so. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under dead leaves, small stones, pieces of wood, heaps of hay, dead plants, pieces of cardboard or paper, and in cracks in the soil. Biology. Seasonality: January-December. Copulating pairs: May, early July. Gravids: May to mid-July, mostly before July (AB, ON). Tenerals: late June to August, November. Adults found overwintering on higher and drier ground, in vacant lots, on roadsides, in gravel pits, on sandy or gravelly hillocks, in hawthorn hedges, at the edge of woods, and in woods; under dead leaves and small stones (mostly), in ant nests, and in the soil (as deep as 15 cm). Active at low temperatures under snow in October-November. Adult food: all lepidopterous and dipterous immature stages (in the field); aphid nymphs, dipterous eggs and larvae, lepidopterous eggs, chrysomelid eggs, immature stages of weevils, larval chrysomelids (in captivity). Economic importance: may play an important role in the control of the Onion Maggot (Hylemya antiqua (Meigen)). Predators: toads, starlings, and mice. Dispersal power. Wing-polymorphic: macropterous (capable of flight) to brachypterous (incapable of flight). Frequent flier (in late afternoon; to artificial lights at night; to artifical lights in captivity). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants and trees). Strongly favored by human activities. Collecting techniques. Pitfall trapping; using an aspirator; sweeping plants. References. Schwarz, 1890 (biology; as quadrimaculatum (Linné)); Lindroth, 1963a and 1963b (natural history); Rivard, 1964c (biology); Frank, 1971b (biology); Kirk, 1975c (ecology, dispersal power; as quadrimaculatum (Linné)); Hsin et al. 1979 (biology); Tyler and Ellis, 1979 (biology); Erwin, 1981 (natural history; as quadrimaculatum (Linné)); Los and Allen, 1983 (biology); Boivin and Hance, 1994 (biology); Levesque and Levesque, 1994 (biology). Bembidion (Hirmoplataphus) quadrulum LeConte, 1861 Ecology. Lowlands and mountains. Banks of rivers and brooks; lake shores. Open ground; moist or wet, at least partially sandy soil. Nocturnal; sheltering during the day under stones or among gravel. Biology. Seasonality: April, June-August. Copulating pairs: early July (MT). Tenerals: mid-June (WA), early July (ID, MT). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Raking the soil; pouring water over the ground. Reference. Lindroth, 1963b (natural history). Bembidion (Notaphus) rapidum (LeConte, 1848) Ecology. Lowlands, mountains, subalpine and alpine zones. Lake shores, banks of rivers and brooks, cattle water holes; wet meadows, pastures, cultivated fields (e.g., corn, tobacco, cabbage, alfalfa, barley, soybean, oat), orchards (e.g., apple), and swamp-forests. Open ground; wet or moist, clayish, muddy or sandy soil covered with some vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in cracks in the soil (mostly), under dead leaves, dead plants, and stones. Biology. Seasonality: February-October, December. Tenerals: July-August. Adult food, in the field: lepidopterous immatures stages, dipterous larvae and pupae. Dispersal pow-

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er. Macropterous. Frequent flier (to artificial lights at night). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; pouring water over the ground; treading the soil with the feet; using an aspirator; turning drift material. References. Lindroth, 1963b (natural history); Bell, 1971 (ecology); Kirk, 1975c (ecology, dispersal power); Hsin et al. 1979 (biology); Erwin, 1981 (natural history); Los and Allen, 1983 (biology); Will et al. 1995 (dispersal power). Bembidion (Hirmoplataphus) recticolle LeConte, 1863 Ecology. Mountains. Banks of cold swift rivers (mostly) and lake shores. Open ground; moist, bare, gravelly soil mixed with sand or clay. Nocturnal; sheltering during the day under stones. Biology. Seasonality: March-August. Tenerals: mid-July (WA). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Raking the soil; pouring water over the ground. Reference. Lindroth, 1963b (natural history). Bembidion renoanum Casey, 1918 Natural History. Unknown. Bembidion (Odontium) robusticolle Hayward, 1897 Ecology. Lowlands and mountains. Stream banks. Open ground; moist, sandy bare soil. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in soil cracks and burrows dug in the ground. Biology. Seasonality: June-August. Defense-mechanism: when alarmed, the adult escapes by flying readily. Dispersal power. Macropterous. Frequent flier (when alarmed). Fast runner. Collecting techniques. Pouring water over the ground; hunting with a butterfly net. Bembidion (Trichoplataphus) rolandi Fall, 1922 Ecology. Lowlands and mountains. River banks; lake shores. Open ground; wet, bare, gravelly soil, sometimes mixed with clay or sand. Nocturnal; sheltering during the day among gravel and under stones. Gregarious. Biology. Seasonality: April-October. Tenerals: July-August (QU, WV). Defense-mechanism: when alarmed, the adult may escape by dashing under water or remain clinging under small stones. Dispersal power. Macropterous. Frequent flier (to artificial lights in captivity). Moderate runner. Collecting techniques. Raking the loose gravel; pouring water over the ground. References. Larochelle, 1972h (ecology); Davidson, 1978 (ecology). Bembidion (Semicampa) roosevelti Pic, 1902 Ecology. Uplands. Vicinity of alkaline lakes, pools, and sloughs. At some considerable distance from water. Open ground; moderately moist, alkaline soil covered with rather

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dense vegetation (e.g., grass, Triglochin). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day among the vegetation. Biology. Seasonality: May-September. Teneral: mid-July (SA). Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous (probably capable of flight). Moderate runner. Collecting technique. Pouring water over the ground. Reference. Lindroth, 1963b (natural history). Bembidion (Plataphodes) rosslandicum Lindroth, 1963 Ecology. Mountains and subalpine zone. Margins of forest snowfields and small streams with cold water. Open ground; moist soil. Biology. Seasonality: May, July-August. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Lindroth, 1963b (natural history). Bembidion (Semicampa) rubiginosum LeConte, 1879 Ecology. Uplands. Borders of alkali ponds. Open ground; firm, moist soil covered with some vegetation. Biology. Seasonality: March-May. Dispersal power. Wing-condition unknown. Moderate runner. Bembidion (Plataphus) rufinum Lindroth, 1963 Ecology. Mountains. Banks of mostly small streams. Bare, gravelly soil. Biology. Seasonality: June-August. Tenerals: late July (AB, BC). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Lindroth, 1963b (natural history). Bembidion (Pseudoperyphus) rufotinctum Chaudoir, 1868 Ecology. Lowlands and mountains. In or near the spray zone of narrow rocky gorges, dams and waterfalls of wooded rivers and large streams. Open or shaded ground; wet, rocky soil, almost bare or covered with algal carpets. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in rock cracks and under debris. Gregarious. Associated with saldid bugs (Pentacora). Biology. Seasonality: April-September. Tenerals: mid-September (VT). Adults found overwintering in woods; in leaf litter. Dispersal power. Macropterous. Occasional flier. Moderate runner. Collecting techniques. Turning debris; pouring water over the ground. References. Cooper, 1976 (natural history); Davidson, 1981 (ecology). Bembidion (Peryphus) rupicola (Kirby, 1837) Ecology. Cultivated fields (e.g., corn, wheat, barley, oat, flax), meadows, sand dunes, gravel pits; upper zone of river banks, lake shores, pond margins, and saline sloughs. Open ground; very dry, clayish, loamy, gravelly or sandy soil covered with some vegetation. Nocturnal; sheltering during the day under stones and vegetal debris. Semigregarious. Biology. Seasonality: April-November. Copulating pairs: early May (AB).

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Tenerals: mid-July to September. Adult food: dipterous larvae (in the field); lepidopterous eggs (in captivity). Defense-mechanism: when alarmed, the adult may escape by flying. Dispersal power. Macropterous. Frequent flier (when alarmed; to artificial lights at night). Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; light trapping. References. Lindroth, 1963b (natural history); Frank, 1971b (biology); Will et al. 1995 (dispersal power). Bembidion (Plataphus) rusticum lenensoides Lindroth, 1963 Ecology. Lowlands and mountains. Margins of running waters. Open ground; moist, bare, gravelly, sometimes clayish soil. Nocturnal; sheltering during the day among gravel. Biology. Seasonality: June-September. Defense-mechanism: when alarmed, the adult often escapes by flying. Dispersal power. Macropterous. Frequent flier (when alarmed). Moderate runner. Collecting techniques. Pouring water over the ground; raking the loose gravel. Bembidion (Plataphus) rusticum rusticum Casey, 1918 Ecology. Lowlands and mountains. Banks of cold, swift brooks and rills. Open ground; wet, gravelly bare soil, sometimes mixed with clay. Nocturnal; sheltering during the day among gravel. Gregarious. Biology. Seasonality: April-October. Copulating pairs: mid-July (QU). Tenerals: late June (NB) to mid-August (ME). Defense-mechanism: when alarmed, the adult may escape by flying. Dispersal power. Macropterous. Frequent flier (when alarmed; to artificial lights in captivity). Moderate runner. Collecting techniques. Pouring water over the ground; raking the loose gravel. References. Lindroth, 1955a, 1963a and 1963b (natural history). Bembidion (Hirmoplataphus) salebratum (LeConte, 1848) Ecology. Lowlands and mountains. Banks of rivers, brooks, and rills; lake shores. Open ground; moist or wet, gravelly, sandy or clayish, bare soil. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day among gravel, under stones, and in cracks in the soil. Gregarious. Biology. Seasonality: April-November. Copulating pairs: early June (QU). Tenerals: late June (NB) to late August (NF). Adults found overwintering on higher and drier ground, in the upper zone of river banks and on hillocks; under small stones. Defense-mechanism: when alarmed, the adult may escape by flying or digging into the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; to artificial lights in captivity). Moderate runner. Collecting techniques. Raking the loose gravel; pouring water over the ground. References. Lindroth, 1955a, 1963a, and 1963b (natural history). Bembidion (Peryphodes) salinarium Casey, 1918 Ecology. Uplands. Borders of saline lakes and ponds, alkali sloughs, and semi-alkaline lakes. Open ground; wet, saline or alkaline soil, bare or covered with sparse vegetation. Noctur-

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nal; sheltering during the day in cracks in the soil (mostly) and under pieces of wood. Biology. Seasonality: February-April, June-July, September. Tenerals: mid-July (MB). Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous (capable of flight). Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Reference. Lindroth, 1963b (natural history). Bembidion satelles Casey, 1918 Ecology. Mountains. Lake shores. Biology. Seasonality: June-September. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Bembidion (Trepanedoris) scenicum Casey, 1918 Natural History. Unknown. Bembidion scopulinum (Kirby, 1837) Ecology. Lowlands, mountains, and subalpine zone. Margins of rivers, brooks, ponds, lakes, and pools; clay pits, gravel pits, and cultivated fields (e.g., corn). Open or slightly shaded ground; moist or wet, clayish or gravelly soil, bare or covered with sparse vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under stones, among gravel, under logs, in moss, and in cracks in the soil. Gregarious. Biology. Seasonality: January, April-November. Copulating pairs: late May (NY) to mid-July (QU). Tenerals: late May to early October; mostly from mid-July on. Adults found overwintering on higher and drier ground, at the edge of woods and in woods; in leaf litter. Dispersal power. Wing-polymorphic: macropterous (capable of flight) to brachypterous (incapable of flight). Frequent flier (to artificial lights at night; to artifical lights in captivity). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Turning stones and drift material. References. Lindroth, 1955a, 1963a, and 1963b (natural history); Dearborne and Donahue, 1993 (dispersal power). Bembidion scudderi LeConte, 1878 Ecology. Uplands. Borders of saline lakes, ponds, and sloughs. Open ground; moist, clayish saline soil, bare or covered with sparse vegetation (e.g., Distichlis, Atriplex). Attracted to metabolites emitted by blue-green algae. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in cracks in the soil. Biology. Seasonality: AprilAugust. Tenerals: June (NV), July-August (BC, SA). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Reference. Lindroth, 1963b (natural history). Bembidion (Odontium) sculpturatum (Motschulsky, 1859) Ecology. River margins. Open ground; moist soil. Diurnal; active in the sunshine. Biology. Seasonality: unknown. Defense-mechanism: when alarmed, the adult escapes by

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flying readily. Dispersal power. Macropterous. Frequent flier (when alarmed; in the bright sunshine). Fast runner. Collecting technique. Hunting with a butterfly net. Bembidion (Peryphus) sejunctum sejunctum Casey, 1918 Ecology. Coastal lowlands. Sea beaches; sand dunes (just above the tidal zone). Open ground; moderately moist bare saline soil consisting of fine sand. Nocturnal; sheltering during the day deep under logs, fallen branches, and boards. Gregarious. Biology. Seasonality: April-October. Copulating pairs: late June (QU). Tenerals: early July (PE) to early August (NF, QU). Defense-mechanism: when alarmed, the adult may feign death for 45 seconds before running away. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous (capable of flight). Occasional flier (to artificial lights in captivity). Moderate runner. Strong burrower. Collecting technique. Turning logs. References. Lindroth, 1955a, 1963a, and 1963b (natural history); Larochelle, 1974a (ecology). Bembidion (Peryphus) sejunctum semiaureum Fall, 1922 Ecology. Coastal lowlands. Sea beaches; dunes (just above the tidemark). Open ground; moderately moist, bare saline soil composed of fine sand. Nocturnal; sheltering during the day deep under pieces of wood and other debris. Gregarious. Biology. Seasonality: June-November. Tenerals: mid-July (WA). Adults found overwintering in woods; in leaf litter and moss. Dispersal power. Wing-dimorphic: macropterous (mostly; probably capable of flight) or brachypterous (incapable of flight). Moderate runner. Strong burrower. Collecting technique. Lifting pieces of wood. Reference. Kavanaugh, 1992 (natural history). Bembidion (Semicampa) semicinctum Notman, 1919 Ecology. Lowlands. Borders of brooks passing through swamps. Ground shaded by bushes (e.g., Salix, Alnus, Cornus); firm, moist, clayish or muddy soil covered with grass, ferns (Onoclea), dead leaves, and moss. Nocturnal; sheltering during the day among dead leaves (mostly), under dead plants and stones. Gregarious. Biology. Seasonality: January, March-November. Tenerals: early May to early June (rarely; QU), August (mostly), October. Adults found overwintering in woods; in leaf litter (mostly) and in clumps of moss and grass. Defense-mechanism: body color-pattern matching the soil. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; pouring water over the ground. Reference. Lindroth, 1963b (natural history). Bembidion (Notaphus) semiopacum Casey, 1924 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight.

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Bembidion (Notaphus) semipunctatum (Donovan, 1806) Ecology. Banks of large rivers. At some distance from water. Shaded ground; moderately moist soil composed of sand-mixed clay, bare or covered with sparse vegetation (e.g., Equisetum). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under vegetal debris. Biology. Seasonality: June-August. Tenerals: early August (SA). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; raking vegetal debris. References. Lindroth, 1945, 1963b, and 1985 (natural history); Koch, 1989 (ecology). Bembidion (Hydriomicrus) semistriatum (Haldeman, 1843) Ecology. Lowlands and mountains. Margins of rivers and brooks crossing forests. Slightly shaded ground; moist or wet, gravelly-sandy, bare soil. Nocturnal; sheltering during the day among gravel and under stones. Gregarious. Biology. Seasonality: February, April-October. Tenerals: late June to September; mostly from mid-July on. Adults found overwintering on higher and drier ground, in woods; under dead leaves. Defense-mechanism: when alarmed, the adult may escape by digging into the soil. Dispersal power. Macropterous. Occasional flier (to artificial lights in captivity). Moderate runner. Collecting techniques. Pouring water over the ground; raking the soil. References. Lindroth, 1963b (natural history); Erwin, 1981 (natural history). Bembidion (Lionepha) sequoiae Lindroth, 1963 Ecology. Mountains. Brook edges. Biology. Seasonality: March, June-August. Tenerals: July (CA). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Bembidion (Plataphus) sierricola Casey, 1924 Ecology. Margins of cold, swift streams. Shaded ground; moist, stony soil. Nocturnal; sheltering during the day under stones, dead leaves, and moss. Biology. Seasonality: April-July. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Lindroth, 1963b (natural history). Bembidion simplex Hayward, 1897 Ecology. Lowlands and mountains. Borders of rivers and brooks crossing forests. Close to water. Shaded ground; moist, bare, gravelly soil, sometimes mixed with sand. Nocturnal; sheltering during the day among gravel and under stones. Gregarious. Biology. Seasonality: April-October. Tenerals: early July to September; mostly from mid-July on. Defense-mechanism: when alarmed, the adult may escape by digging. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pouring water over the ground; raking the soil. Reference. Lindroth, 1963b (natural history).

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Bembidion (Trepanedoris) siticum Casey, 1918 Ecology. Lowlands and mountains. Marshy borders of slow rivers and brooks; edges of eutrophic marshes; mud flats. Open ground; wet, clayish or muddy soil, bare or covered with sparse vegetation (e.g., Equisetum). Nocturnal; sheltering during the day in cracks in the soil and under stones. Gregarious. Biology. Seasonality: May-July, September, December. Tenerals: early July (ID,WA). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pouring water over the ground. Reference. Lindroth, 1963b (natural history). Bembidion sordidum (Kirby, 1837) Ecology. Lowlands and mountains. Banks of large rivers. At some distance from water. Shaded ground; moist, clay soil covered with sparse vegetation. Nocturnal; sheltering during the day under pieces of wood, clods of earth, and stones. Biology. Seasonality: June-August. Tenerals: mid-May (BC) to late July (MB); mostly before July. Adult food, in the field: dipterous larvae. Dispersal power. Wing-dimorphic: macropterous (probably capable of flight) or brachypterous (incapable of flight). Occasional flier (to artificial lights at night). Frequent in seashore drift material, which indicates flight ability. Moderate runner. Collecting technique. Turning pieces of wood and drift material. References. Lindroth, 1963b (natural history); Kirk, 1975c (dispersal power). Bembidion (Amerizus) spectabile (Mannerheim, 1852) Ecology. Forests (coniferous, mixed). Sometimes above the tree line. Shaded ground; moderately moist soil covered with leaf litter. Nocturnal; sheltering during the day deep under logs, stones, and under the loose bark of trees. Biology. Seasonality: May-August. Tenerals: July (WA). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Strong burrower. Collecting technique. Turning debris. References. Lindroth, 1963b (natural history); Kavanaugh, 1992 (natural history). Bembidion (Cyclolopha) sphaeroderum Bates, 1882 Ecology. Mountains. Mexico: vicinity of a pond; border of a brook; a prairie, at forest edge. Biology and Dispersal power. Unknown. Bembidion (Peryphanes) stephensii Crotch, 1866 Ecology. Lowlands. Steep slopes with trickling water: often beside brooks, rivulets, pools and ponds; also in gravel pits, sand pits, clay pits, vacant lots, and roadsides. More or less shaded ground; wet soil consisting of clay or clay-mixed sand, bare or covered with sparse vegetation. Mostly nocturnal; usually sheltering during the day under small embedded stones and in cracks in the soil. In tunnels of worms (Lumbricus; Europe). Biology. Seasonality: March-October. Tenerals: mid-June to October; mostly from midJuly on. Adults found overwintering on higher and drier ground, in the upper zone of

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gravel pits, brooks, and vacant lots; under stones. Adult food, in captivity (Europe): dead flies and small carabids. Adults are cannibalistic. Predators: toads. Dispersal power. Macropterous. Occasional flier (during the day; at sunset; to artificial lights in captivity). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strong burrower. Strongly favored by human activities. Collecting techniques. Lifting small embedded stones; pouring water over the ground. References. Lindroth, 1945, 1955a, 1963a, 1963b, and 1985 (natural history); Koch, 1989 (ecology). Bembidion (Plataphus) stillaguamish Hatch, 1950 Ecology. Lowlands and mountains. Stream margins. Open ground; moist, stony soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: May, July. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Turning stones. Reference. Hatch, 1953 (ecology). Bembidion striola (LeConte, 1852) Ecology. Coastal lowlands. Stream margins. Open ground; moist soil. Biology. Seasonality: January, June, August, December. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Hatch, 1953 (ecology). Bembidion (Hirmoplataphus) subaerarium Casey, 1924 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion (Peryphanes) subangustatum Hayward, 1897 Ecology. Mountains. Biology. Seasonality: June-August. Dispersal power. Unknown. Bembidion (Plataphus) sulcipenne hyperboroides Lindroth, 1963 Ecology. Banks of large streams. Open ground; moist, bare, gravelly soil. Nocturnal; sheltering during the day under stones and leaf litter. Biology. Seasonality: June-August. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Turning stones; raking the leaf litter. Bembidion (Plataphus) sulcipenne prasinoides Lindroth, 1963 Ecology. Lowlands and mountains. Banks and bars: along cold swift rivers and brooks. Open ground; moist or wet, gravelly, bare soil. Mostly nocturnal; sheltering during the day among gravel and under stones. Gregarious. Biology. Seasonality: June-October. Copulating pairs: early July (QU). Tenerals: late June (NB) to late August (NF). Defensemechanism: when alarmed, the adult may escape by flying or digging. Dispersal power. Macropterous. Frequent flier (when alarmed; to artificial lights in captivity). Moder-

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ate runner. Collecting techniques. Pouring water over the ground; raking the soil. References. Lindroth, 1963a (as lenense Poppius) and 1963b (natural history). Bembidion (Peryphus) tetracolum Say, 1823 Ecology. Lowlands and mountains. River banks, lake shores, and marsh borders. Cultivated fields (e.g., corn, oat, soybean), meadows, pastures, gardens, vacant lots, courtyards, wharfs, clay pits, gravel pits, sand pits, and vicinity of human habitations. Open ground; moderately moist, clay soil (sometimes sand- or gravel-mixed) covered with sparse vegetation (e.g., weeds). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under stones, vegetal debris, and in cracks in the soil. Biology. Seasonality: February-December. Copulating pairs: late May (QU). Tenerals: July-September. Adults found overwintering on higher and drier ground, in vacant lots, in the upper zone of river banks, at the edge of woods, and in woods; under stones, pieces of wood, and dead leaves. Adult food: springtails and insect eggs (in the field; Europe); earthworms (in captivity). Predators: toads. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous (capable of flight). Occasional flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants, e.g., Solidago). Clearly effective as a colonist: quick to establish in newly modified habitats. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; sweeping plants. References. Lindroth, 1945 and 1955a (as ustulatum Linné), 1963a, 1963b and 1985 (natural history); Kirk, 1975c (ecology, dispersal power); Koch, 1989 (ecology); Luff, 1993 (biology). Bembidion (Peryphanes) texanum Chaudoir, 1868 Ecology. Lowlands and mountains. Banks of rivers and brooks crossing forests. Caves (occasionally). Open or shaded ground; moist or wet, gravelly soil (sometimes claymixed) covered with some vegetation. Nocturnal; sheltering during the day among gravel (mostly) and vegetal debris. Biology. Seasonality: April-July, October. Tenerals: late May to mid June (AR). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; raking the loose gravel. Reference. Holeski and Graves, 1978 (ecology). Bembidion (Leuchydrium) tigrinum LeConte, 1879 Ecology. Coastal lowlands. Seashore: intertidal zone. Open ground; wet soil. Biology and Dispersal power. Unknown. Bembidion timidum (LeConte, 1848) Ecology. Lowlands, mountains, subalpine and alpine zones. Borders of ponds, cattle water holes, and streams; alkali pastures, meadows, cultivated fields (e.g., wheat, corn), and sand dunes. Open ground; moist, sandy, clayish, muddy or peaty soil, mixed with organic debris, with bare spots surrounded by sparse vegetation. Mostly nocturnal; some-

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times active in the sunshine; usually sheltering during the day in mud cracks, under pieces of wood, and stones. Biology. Seasonality: March-September. Tenerals: mid-July to mid-August (AB, SA). Adult food, in the field: dipterous larvae. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; pitfall trapping. References. Lindroth, 1963b (natural history); Bell, 1971 (ecology); Kirk, 1975c (ecology, dispersal power). Bembidion (Diplocampa) transparens (Gebler, 1829) Ecology. Lowlands and mountains. Marshy borders of pools, ponds, lakes, and irrigation canals; borders of marshes and bogs; swamps and wet meadows. Open ground; wet, clayish, muddy or peaty, sometimes saline soil, rich in organic debris, covered with dense vegetation (e.g., Carex, Typha, Juncus, moss carpets). Nocturnal; sheltering during the day under dead leaves and mats of dead plants, under fallen branches and moss. Gregarious. Biology. Seasonality: January, March-December. Tenerals: early June (NY) to mid-August (MN); mostly from mid-July on. Adults found overwintering on higher and drier ground, on sand dunes, at the edge of woods, and in woods; in leaf litter (mostly) and in rotten logs. Active at low temperatures under snow in October-November. Dispersal power. Wing-dimorphic: macropterous (capable of flight) or brachypterous (incapable of flight). Frequent flier (to artificial lights at night; to artifical lights in captivity). Moderate runner. Frequent in lake- and seashore drift material, which indicates flight ability. Favored by spring floods along large rivers. Collecting techniques. Treading the vegetation into the water; light trapping; turning drift material. References. Lindroth, 1945, 1955a, 1963a, 1963b and 1985 (natural history); Kirk, 1975c (dispersal power). Bembidion transversale Dejean, 1831 Ecology. Lowlands and mountains. Banks of large rivers; lake shores; sea beaches (including the tidal zone). Close to water. Open or slightly shaded ground; moist or wet, bare soil consisting of gravel or coarse sand. Nocturnal; sheltering during the day under stones, logs, and fallen branches. Gregarious. Biology. Seasonality: February-December. Copulating pairs: early July (QU). Tenerals: late June to late August; mostly from mid-July on. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Frequent flier (to artificial lights in captivity). Moderate runner. Collecting technique. Turning stones. References. Lindroth, 1955a, 1963a, and 1963b (natural history); Kavanaugh, 1992 (natural history). Bembidion (Trechonepha) trechiforme (LeConte, 1852) Ecology. Mountains. Biology. Seasonality: January-July, December. Dispersal power. Macropterous, probably capable of flight. Bembidion triviale Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight.

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Bembidion ulkei Lindroth, 1963 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion (Notaphus) umbratum (LeConte, 1848) Ecology. Lowlands and mountains. Banks of large rivers. Ground shaded by scattered small bushes (e.g., Salix); sloping, wet bare soil consisting of clay-mixed sand. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in cracks in the soil. Biology. Seasonality: May-August, November. Tenerals: early July (WY) to late August (AB). Dispersal power. Macropterous. Frequent flier (swarming to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Reference. Lindroth, 1963b (natural history). Bembidion umiatense Lindroth, 1963 Ecology. Vicinity of ponds and brooks (in the tundra). Open ground; moist, gravelly soil covered with some vegetation (e.g., moss, sedges, grass). Biology. Seasonality: JulyAugust. Dispersal power. Wing-dimorphic: macropterous (probably capable of flight) or brachypterous (incapable of flight). Moderate runner. Collecting technique. Pouring water over the ground. Reference. Lindroth, 1963b (natural history). Bembidion (Lymneops) utahense Van Dyke, 1925 Natural History. Unknown. Bembidion (Plataphus) vandykei Blaisdell, 1902 Natural History. Unknown. Bembidion (Notaphus) variegatum Say, 1823 Ecology. Lowlands. River banks. Ground shaded by bushes (e.g., Salix); wet, clayish or sandy-clayish soil, bare or covered with depressed vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under logs, pieces of wood, and other debris. Biology. Seasonality: January-December. Tenerals: early July (KY) to late August (QU). Adults found overwintering under the loose bark of fallen trees and tree-stumps (e.g., Platanus). Predators: frogs and swallows. Defense-mechanism: body color-pattern matching the soil. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; turning debris. References. Blatchley, 1910 (biology, dispersal power); Kirk, 1975c (dispersal power); Holeski and Graves, 1978 (ecology); Erwin, 1981 (natural history).

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Bembidion versicolor (LeConte, 1848) Ecology. Lowlands, mountains, subalpine and alpine zones. Margins of ponds, eutrophic marshes, pools (in gravel pits and sand pits), lakes, and slow rivers, brooks, and irrigation canals; swamps, flood-plain forests, moist meadows, mud flats, cultivated fields (e.g., corn, barley, cabbage), and lawns. Open or half-shaded ground; moist, sandy, clayish or peaty soil, mixed with organic debris, with bare spots surrounded by sparse vegetation (e.g., Carex). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under dead leaves, pieces of wood, heaps of hay, mats of dead plants (Gramineae), small stones, at the base of plants (Carex), and in cracks in the soil. Gregarious. Biology. Seasonality: January-December. Gravids: late July (AB). Tenerals: early May to mid-September; mostly from mid-July on. Adults found overwintering on higher and drier ground, on hills and hillocks, in the upper zone of sand pits and gravel pits, in field hedges, at the edge of woods, and in woods; in leaf litter (mostly), under small stones, and in rotten logs. Adult food, in captivity: lepidopterous eggs. Predators: frogs, toads, ducks, starlings, and swallows. Defense-mechanism: when alarmed, the adult escapes by flying readily; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; in late afternoon; to artificial lights at night). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Moderately favored by human activities. Collecting techniques. Pouring water over the ground; treading the soil with the feet. References. Lindroth, 1955a, 1963a, and 1963b (natural history); Frank, 1971b (biology); Kirk, 1975c (ecology, dispersal power); Kavanaugh, 1992 (natural history). Bembidion (Notaphus) versutum LeConte, 1878 Ecology. Lowlands and mountains. Margins of slow rivers and brooks (mostly), ponds and, sometimes quite small pools. Close to water. Open ground; wet, bare sandy soil, often mixed with clay. Nocturnal; sheltering during the day in cracks in the soil (mostly) and under vegetal debris. Biology. Seasonality: April-September. Tenerals: August (MN, NY). Dispersal power. Macropterous. Occasional flier (to artificial lights at night; to artifical lights in captivity). Moderate runner. Occasional climber (on plants). Collecting technique. Pouring water over the ground. References. Lindroth, 1963b (natural history); Dearborne and Donahue, 1993 (dispersal power). Bembidion (Plataphus) vespertinum Casey, 1918 Ecology. Stream margins. Moist soil. Biology. Seasonality: June. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Bembidion (Plataphodes) viator Casey, 1918 Ecology. Lowlands and mountains. Borders of marshes. Open ground; soft, wet soil covered with some vegetation (e.g., Carex). Nocturnal; sheltering during the day in vegetal debris. Biology. Seasonality: June-August. Dispersal power. Macropterous, proba-

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bly capable of flight. Moderate runner. Collecting technique. Pouring water over the ground. Reference. Kavanaugh, 1992 (natural history). Bembidion (Emphanes) vile (LeConte, 1852) Ecology. Coastal lowlands. A lagoon, with wet, clay soil covered with some vegetation (Salicornia, grass; BC). Biology. Seasonality: May-July. Dispersal power. Macropterous, probably capable of flight. References. Lindroth, 1963b (natural history); Ball and Bousquet, 2001 (taxonomy). Bembidion viridicolle (LaFerté-Sénectère, 1841) Ecology. Lowlands and mountains. Borders of lakes, ponds, and sloughs; salt meadows and salt flats. Open ground; wet, clayish, alkaline or saline soil covered with sparse vegetation (e.g., Salicornia, Spergularia, Atriplex, Distichlis). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in cracks in the soil. Biology. Seasonality: May-August. Tenerals: late July (MB) and early August (NE, SA). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Occasional climber (on plants). Collecting techniques. Pouring water over the ground; treading the soil with the feet; sweeping. References. Lindroth, 1963b (natural history); Kirk, 1975c (ecology, dispersal power). Bembidion vividum Casey, 1884 Natural History. Unknown. Bembidion (Notaphus) vulpecula Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Bembidion wickhami Hayward, 1897 Ecology. Mountains. Stream banks. Open ground; stony soil. Biology. Seasonality: JuneJuly. Tenerals: July-August (CA). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Erwin and Kavanaugh, 1981 (natural history; as carlhi Erwin and Kavanaugh, 1981). Bembidion (Amerizus) wingatei Bland, 1864 Ecology. Lowlands, mountains, subalpine and alpine zones. Forests (deciduous, mixed, coniferous); subalpine and alpine meadows; open grasslands (NF); caves (regular; southern Appalachians). Shaded or open ground; moderately moist soil covered with thick leaf litter (in forests) or scattered stones. Nocturnal; sheltering during the day under large well-embedded stones, dead leaves, in decaying logs, in and under moss. Biology.

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Seasonality: March-November. Tenerals: late June to early September; mostly from midJuly on. Adults found overwintering in woods; under well-embedded stones. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting techniques. Raking the leaf litter; turning stones; pitfall trapping. References. Lindroth, 1963a and 1963b (natural history); Barr, 1969 (ecology). Bembidion (Peryphanes) yukonum Fall, 1926 Ecology. Lowlands and mountains. River banks, pool margins, and wastelands. At some distance from water. Open ground; moist, clayish, bare soil. Nocturnal; sheltering during the day under stones and fallen dead branches. Biology. Seasonality: May-August. Dispersal power. Wing-dimorphic: macropterous (capable of flight) or brachypterous (incapable of flight). Moderate runner. Collecting technique. Turning stones. References. Lindroth, 1963b and 1985 (natural history). Bembidion (Bracteon) zephyrium Fall, 1910 Ecology. Coastal lowlands. Salt marshes, sea beaches (above the tidemark), and estuaries. Open ground; moist, often saline, bare sandy soil, frequently mixed with clay. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days in soil cracks and burrows dug in the ground. Biology. Seasonality: May-August. Tenerals: late July (BC). Defense-mechanism: when alarmed, the adult runs and flies readily like a tiny Cicindela; the beetle is difficult to pursue; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; in the bright sunshine). Fast runner. Collecting techniques. Pouring water over the ground; hunting with a butterfly net. References. Lindroth, 1963b (natural history); Maddison, 1993 (ecology). Blemus discus (Fabricius, 1792) Ecology. Mostly in the vicinity of rivers, brooks, pools, ponds, and lakes, in swamps, Sphagnum-bogs, and beaver houses. Also in gravel pits, roadside ditches, cultivated fields (e.g., corn, cabbage, raspberry), and vacant lots. Often near water. Open ground; slightly moist, usually clayish or peaty soil covered with dense vegetation (e.g., grass, sedges). Nocturnal; sheltering during the day under well-embedded stones (mostly), pieces of wood, and heaps of hay. In the burrows of rodents and moles (Europe). Biology. Seasonality: May-September. Tenerals: mid-June to late September; mostly before mid-July. Adult food, in the field: staphylinids (Japan). Predators: frogs and toads. Defense-mechanism: when alarmed, the adult may emit a musky smell. Dispersal power. Macropterous. Frequent flier (sometimes swarming by the hundreds to artificial lights at night; up to 1 km from its habitat). Occasional in seashore drift material, which indicates flight ability. Fast runner. Strong burrower. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Light trapping; turning stones; pouring water over the ground. References. Lindroth, 1945, 1961, and 1985 (natural history); Liebherr and Ward, 1978 (ecology, dispersal power); Koch, 1989 (ecology); Dearborne and Donahue, 1993 (dispersal power); Levesque and Levesque, 1994 (biology).

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Blethisa catenaria Brown, 1944 Ecology. Margins of cold, small pools (in the tundra). Open ground; soft, wet, muddy soil covered with dense vegetation (Eriophorum- and Scorpidium-carpets). Biology. Seasonality: June-August. Tenerals: August (AK). Defense-mechanism: body color-pattern matching its surroundings. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Treading the vegetation into the water. References. Lindroth, 1954 (dispersal power) and 1961 (natural history); Nelson, 1983 (ecology). Blethisa inexspectata Goulet and Smetana, 1983 Ecology. A small swampy area adjacent to a stream, with wet soil covered with debris and moss (AK). Nocturnal. Biology. Seasonality: June. Dispersal power. Macropterous, probably capable of flight. Blethisa julii LeConte, 1863 Ecology. Lowlands and mountains. Margins of cold ponds and pools. Open or shaded ground; soft, wet, muddy soil, with bare spots surrounded by moss-carpets. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days in moss. Biology. Seasonality: May-August. Adult food, in the field: small insects. Defense-mechanism: when alarmed, the adult may stridulate; body color-pattern matching its surroundings. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Treading the vegetation into the water. References. Lindroth, 1954, 1955a, 1961, and 1963a (natural history). Blethisa multipunctata aurata Fischer von Waldheim, 1828 Ecology. Lowlands and mountains. Margins of eutrophic marshes, ponds, lakes, and pools; swamps and marshy roadside ditches. Associated with cold water. Open or shaded ground; soft, wet, muddy or clayish soil, rich in organic debris, covered with dense vegetation (e.g., Typha, Carex, moss (not Sphagnum)). Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under dead fallen plants, dead leaves, and pieces of wood. Biology. Seasonality: March-October. Copulating pairs: late June (QU). Gravids: late June (VT). Tenerals: early August (QU). Adults found overwintering on higher and drier ground, in vacant lots (up to 100 m away from marshes) and in woods; under dead leaves. Adult and larval food, in captivity: mealworms and shrimps. Parasites: Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult may stridulate or escape by swimming and diving into the water (may remain submerged for a few minutes); body color-pattern matching its surroundings. Dispersal power. Macropterous. Regular flier (in the sunshine; to artificial lights in captivity). Occasional in drift material, which indicates flight ability. Fast runner. Excellent swimmer and diver. Collecting technique. Treading the vegetation into the water. References. Lindroth, 1954, 1955a, 1961, and 1963a (natural history).

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Blethisa oregonensis LeConte, 1853 Ecology. Lowlands and mountains. Borders of small ponds, marshes, and marshy banks of slow rivers. Associated with cold water.Open ground; soft, wet soil, rich in organic debris, covered with dense vegetation (e.g., Eleocharis, Carex). Mostly diurnal; active in the sunshine or sheltering on cloudy or cool days under dead plants and dead leaves. Biology. Seasonality: February-November. Tenerals: mid-September (WA). Adults found overwintering under the loose bark of trees, over 1 km from its breeding habitat. Defense-mechanism: when alarmed, the adult may stridulate or escape by diving into the water; body color-pattern matching its surroundings. Dispersal power. Macropterous, probably capable of flight. Fast runner. Excellent swimmer and diver. Collecting technique. Treading the soil with the feet. References. Lindroth, 1961 (natural history); Nelson, 1988 (biology); LaBonte and Johnson, 1989 (ecology). Blethisa quadricollis Haldeman, 1847 Ecology. Lowlands and mountains. Margins of ponds and pools (in acid bogs); margins of Typha-marshes and swamps. Associated with cold water. Open ground; soft, wet soil, with floating moss carpets (Sphagnum, Oxycoccus, scattered Myrica), or rich in organic debris and densely vegetated (Typha). Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days in moss (mostly) or among plant debris. Biology. Seasonality: April-September. Copulating pairs: mid-July (QU). Gravids: mid-June to early July (QU). Tenerals: mid-July (QU) to late August (NF); mostly from mid-July on. Adult and larval food, in captivity: mealworms. Parasites: Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult may stridulate, escape by diving into the water (may remain submerged for 3 minutes), or emit a strong repugnant odor that clings strongly to fingers when the beetle is seized; body color-pattern matching its surroundings. Dispersal power. Macropterous. Occasional flier (to artificial lights at night; to artifical lights in captivity). Occasional in lake- and seashore drift material, which indicates flight ability. Fast runner. Excellent swimmer and diver. Collecting technique. Treading the vegetation into the water. References. Frost, 1932 (ecology); Lindroth, 1954, 1955a, 1961, and 1963a (natural history); Larochelle, 1973f (biology); Landry and Anctil, 1975 (dispersal power). Brachinus (Neobrachinus) aabaaba Erwin, 1970 Ecology. Unknown. Biology. Seasonality: April-October. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Brachinus (Neobrachinus) adustipennis Erwin, 1969 Ecology. Lowlands. River banks; marsh borders (e.g., Scirpus, Typha). Wet, muddy, soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day in clumps of grass, and under stones and pieces of wood. Biology. Seasonality: January, March-October. Tenerals: May (FL), August (TN). Dispersal power. Macropterous. Fre-

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quent flier (to artificial lights at night). Fast runner. Collecting techniques. Pitfall trapping; light trapping. Reference. Erwin, 1970b (natural history). Brachinus (Neobrachinus) alexiguus Erwin, 1970 Ecology. Unknown. Biology. Seasonality: March-April. Dispersal power. Brachypterous, incapable of flight. Brachinus (Neobrachinus) alternans Dejean, 1825 Ecology. Lowlands. River banks, swamps, low forests, salt meadows, and cultivated fields (e.g., corn). Shaded or open ground; moist, sometimes saline soil. Nocturnal; sheltering during the day under pieces of wood and logs. Gregarious. Biology. Seasonality: January-December. Tenerals: April (rarely; IL), August (TN), September (NE), November (NC). Predators: raccoons and skunks. Parasites: Laboulbeniales (Fungi). Defensemechanism: when alarmed, the adult may emit a repugnant audible spray from the pygidial glands, which can cause great pain to the fingers if the beetle is seized. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Collecting techniques. Pitfall trapping; lifting pieces of wood; baiting traps with banana. Reference. Erwin, 1970b (natural history). Brachinus (Neobrachinus) americanus (LeConte, 1844) Ecology. Pond margins, flood-plain forests, and cultivated fields (e.g., corn). Open or shaded ground; moist or wet soil. Nocturnal; sheltering during the day under pieces of wood. Gregarious. Biology. Seasonality: February-October. Tenerals: September (AK). Defensemechanism: when alarmed, the adult may emit a repugnant audible spray from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting techniques. Pitfall trapping; lifting pieces of wood. Reference. Erwin, 1970b (natural history). Brachinus (Neobrachinus) azureipennis Chaudoir, 1876 Ecology. Grasslands and pastures (Mexico). Nocturnal; sheltering during the day. Biology. Seasonality: January-February, July-August, October. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Reference. Erwin, 1970b (natural history). Brachinus (Neobrachinus) capnicus Erwin, 1970 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Brachinus (Neobrachinus) cibolensis Erwin, 1970 Ecology. Pond margins. Moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: June-September. Adult food, in the field: flies and other arthropods.

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Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Brachinus (Neobrachinus) conformis Dejean, 1831 Ecology. Lowlands. Biology. Seasonality: May, June, November. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Collecting technique. Light trapping. Brachinus (Neobrachinus) cordicollis Dejean, 1826 Ecology. Lowlands and mountains. River banks and lake shores. Open ground; wet, sandy, gravelly or clayish soil covered with sparse vegetation. Nocturnal; sheltering during the day under stones. Gregarious. Biology. Seasonality: April-October. Gravids: mid-June and late August (QU), early July (ON). Tenerals: April (NY), May (ON), July (PA), September (WI). Adult food, in captivity: meat. Defense-mechanism: when alarmed, the adult may emit a repugnant audible spray from the pygidial glands. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Collecting techniques. Pitfall trapping; turning stones. References. Larochelle, 1974a (ecology); Erwin, 1970b and 1981 (natural history). Brachinus (Neobrachinus) costipennis Motschulsky, 1859 Ecology. Stream margins. Moist soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: February-December. Tenerals: July (CA). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Collecting technique. Turning stones. Reference. Erwin, 1970b (natural history). Brachinus (Neobrachinus) cyanipennis Say, 1823 Ecology. Lowlands. Banks of rivers and brooks; lake shores; flood-plain forests and borders of eutrophic marshes; beaver houses. Open or shaded ground; moist or wet soil consisting of sand, gravel, clay, or organic mud. Nocturnal; sheltering during the day under stones (mostly), pieces of wood, dead leaves, dead grasses, and in cracks in the soil. Gregarious. Biology. Seasonality: March-October. Gravids: late June (once; VT), late August to late September (QU). Eggs are laid singly or in clusters of 3-25, in mud or clay cells, on the underface of loosely set stones, on dead twigs and plant stems. Tenerals: May (KS), August (mostly; QU, VT). Adults found overwintering on higher and drier ground, in the upper zone of sand pits and gravel pits; under stones. Adult food, in the field: dead or dying small arthropods. The larva is an ectoparasite of beetle pupae (Gyrinus: gyrinids). Predators: scelionid wasps (egg-parasites). Defense-mechanism: when alarmed, the adult may emit a repugnant audible spray from the pygidial glands. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Collecting techniques. Pitfall trapping; turning stones. References. King, 1919 (biology); Lindroth, 1969a (ecology); Erwin, 1970b (natural history); Larochelle, 1974a (ecology); Kirk, 1975c (dispersal power); Erwin, 1981 (natural history).

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Brachinus (Neobrachinus) cyanochroaticus Erwin, 1969 Ecology. Lowlands and mountains. Margins of ponds, lakes, and eutrophic marshes; borders of rivers (in lows deciduous forests), swamps, and beaver houses. Open or half-shaded ground; wet, sandy, gravelly, clayish, or muddy soil. Nocturnal; sheltering during the day under stones (mostly), dead leaves, and logs. Gregarious. Biology. Seasonality: April-December. Gravids: May-June and August-September (QU, VT). Tenerals: April-May (QU), August (QU, ME. Adults found overwintering in colonies on higher and drier ground, at the edge of woods, in woods, in the upper zone of gravel pits and sand pits; in sandy or gravelly soil, under large, well-embedded stones (mostly; 17.5-25.5 cm deep) and in leaf litter. Defense-mechanism: when alarmed, the adult may emit a repugnant audible spray from the pygidial glands. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Strong burrower. Collecting techniques. Pitfall trapping; turning stones. References. Lindroth, 1969a (ecology); Erwin, 1970b (natural history); Will et al. 1995 (dispersal power). Brachinus (Neobrachinus) elongatulus Chaudoir, 1876 Ecology. Stream and pond margins. Moist, loamy soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: January-December. Tenerals: August (AZ). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Fast runner. Collecting techniques. Pitfall trapping; light trapping. Reference. Erwin, 1970b (natural history). Brachinus (Neobrachinus) favicollis Erwin, 1965 Ecology. Mountains. Stream margins. Moist, sandy soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: January-October. Tenerals: May (AZ, CA), July (AZ). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Collecting techniques. Pitfall trapping; turning stones. Reference. Erwin, 1970b (natural history). Brachinus (Neobrachinus) fulminatus Erwin, 1969 Ecology. Lowlands. Sphagnum-bogs. Open ground; wet, peaty soil covered with moss. Nocturnal; sheltering during the day. Biology. Seasonality: April-November. Tenerals: April (IN). Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Pitfall trapping. Reference. Erwin, 1970b (natural history). Brachinus (Neobrachinus) fumans (Fabricius, 1781) Ecology. Lowlands and mountains. Banks of rivers and brooks; lake shores; edges of reservoirs and irrigation canals; wet meadows. Open or half-shaded ground; wet, gravelly, clayish, or muddy, sometimes saline soil. Nocturnal; sheltering during the day

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under stones, pieces of wood, and dead fallen grass. Gregarious. Biology. Seasonality: March-October. Gravids: late August (QU). Tenerals: mid-August to early September (BC, QU). Adults found overwintering on higher and drier ground, in woods and on sandy-gravelly hillocks; in the soil (7.5-17.5 cm deep) under well-embedded stones. Adult food: dead or dying insects, and animal matter (in the field); earthworms and mealworms (in captivity). Parasites: Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult may emit a repugnant audible spray from the pygidial glands, which can cause great pain to the fingers if the beetle is seized. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Fast runner. Strong burrower. Collecting techniques. Pitfall trapping; turning debris; light trapping. References. Erwin, 1970b and 1981 (natural history); Kirk, 1975c (dispersal power); Will et al. 1995 (dispersal power). Brachinus (Neobrachinus) gebhardis Erwin, 1965 Ecology. Lowlands and mountains. Stream margins. Open ground; moist, rocky soil with underlying gravel, covered with sparse low vegetation. Biology. Seasonality: January-November. Tenerals: September (CA). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Collecting technique. Pitfall trapping. Reference. Erwin, 1970b (natural history). Brachinus (Neobrachinus) geniculatus Dejean, 1831 Ecology. Mexico: palm forests at the edge of a grassy swamp; margins of a stream; a roadside. Biology. Seasonality: March-December. Tenerals: October (Mexico). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Fast runner. Reference. Erwin, 1970b (natural history; as rhytiderus Chaudoir, 1876). Brachinus (Neobrachinus) hirsutus Bates, 1884 Ecology. Stream and pond margins. Moist soil composed of coarse gravel. Nocturnal; sheltering during the day. Biology. Seasonality: January, March-October, December. Tenerals: June-July (AZ). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Collecting technique. Pitfall trapping. Reference. Erwin, 1970b (natural history). Brachinus (Neobrachinus) ichabodopsis Erwin, 1970 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Brachinus (Neobrachinus) imperialensis Erwin, 1965 Ecology. Uplands. Biology. Seasonality: March-April, June-September, November. Dispersal power. Macropterous. Frequent flier (to artificial lights at light).

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Brachinus (Neobrachinus) imporcitis Erwin, 1970 Ecology. Mountains. Biology. Seasonality: January, May-September. Tenerals: May (AZ). Dispersal power. Brachypterous, incapable of flight. Fast runner. Reference. Erwin, 1970b (biology, dispersal power). Brachinus (Neobrachinus) janthinipennis (Dejean, 1831) Ecology. Lowlands and mountains. Lake shores and river banks; borders of pools and eutrophic marshes. Open or half-shaded ground; moist or wet, muddy, clayish, sandy, or gravelly, almost bare soil. Nocturnal; sheltering during the day under stones and pieces of wood. Gregarious. Biology. Seasonality: April-October. Tenerals: May, JulyOctober; mostly from July on. The larva is an ectoparasite of gyrinid (Dineutes) and hydrophilid (Tropisternus) pupae, and pupates within the mud cocoon of its host. Defense-mechanism: when alarmed, the adult may emit a repugnant audible spray from the pygidial glands; when alarmed, the larva lifts the anterior part of its body so that feet are in the air and maintains this position for some minutes. Dispersal power. Submacropterous, probably incapable of flight. Fast runner. Collecting techniques. Pitfall trapping; turning debris. References. Wickham, 1893 and 1894 (biology); Dimmock and Knab, 1904 (biology); Erwin, 1970b (natural history); Kirk, 1975c (ecology, dispersal power). Brachinus (Neobrachinus) javalinopsis Erwin, 1970 Ecology. Pond margins. Moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: January, May-September. Oviposition: mid to late July (AZ). Tenerals: July (NM) to November (AZ). Adult food, in captivity: flies and other small arthropods. The larva is an ectoparasite of hydrophilid beetle (Tropisternus) pupae. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Fast runner. Collecting techniques. Pitfall trapping; light trapping. References. Erwin, 1970b (natural history); Juliano, 1985a (biology). Brachinus (Neobrachinus) kansanus LeConte, 1862 Ecology. River and brook margins (mostly); sand dunes. Open or shaded ground; moist, sandy soil. Nocturnal; sheltering during the day. Biology. Seasonality: June-October. Tenerals: July (OK). Adult food, in the field: beetles and other small arthropods. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Pitfall trapping. Reference. Erwin, 1970b (natural history). Brachinus (Neobrachinus) kavanaughi Erwin, 1969 Ecology. Mountains. Lake shores and stream margins. Moist, gravelly-clayish soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: February-October Tenerals: May (TX), September (IL). Dispersal power. Macropterous. Occasional flier

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(to artificial lights at night). Fast runner. Collecting techniques. Pitfall trapping; turning stones. References. Erwin, 1970b (natural history); Kirk, 1975c (ecology). Brachinus (Neobrachinus) lateralis Dejean, 1831 Ecology. Lake shores; margins of ponds and streams; wet meadows. Wet, gravelly, sometimes saline soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: January-December. Oviposition: May-September (AZ). Eggs are laid within one meter from water. Tenerals: August-November (CA, AZ). Adult food, in captivity: larval and adult flies, other small arthropods, and dead salamander larvae. The larva is an ectoparasite of dytiscid (Eretes) and hydrophilid (Tropisternus, Berosus) pupae. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Fast runner. Collecting techniques. Pitfall trapping; turning stones; light trapping. References. Erwin, 1970b (natural history); Juliano, 1984, 1985a, 1985b, 1986a, and 1986b (biology). Brachinus (Neobrachinus) medius T.W. Harris, 1828 Ecology. Lowlands and mountains. Lake shores; margins of ponds, reservoirs, marshes, pools, rivers, and brooks; flood-plain forests, swamps, bogs, and wet pastures. Open or shaded ground; moist or wet, clayish or muddy soil (sometimes sand- or gravel-mixed) covered with sparse vegetation. Nocturnal; sheltering during the day under dead leaves, stones, dead plants, pieces of wood, and logs. Gregarious. Biology. Seasonality: January-December. Gravids: mid-May (once), late August to early October (QU). Tenerals: May (BC), August (QU), September (MI). Adults found overwintering on higher and drier ground, at the edge of woods, in woods, and in the upper zone of sand pits and gravel pits; in the soil (17.5-37.5 cm deep) and under stones. Defense-mechanism: when alarmed, the adult may emit a repugnant audible spray from the pygidial glands. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Fast runner. Favored by spring floods along large rivers. Collecting techniques. Pitfall trapping; raking the leaf litter from the soil; light trapping. References. Lindroth, 1969a (natural history); Erwin, 1970b (natural history); Kirk, 1975c (dispersal power). Brachinus (Neobrachinus) mexicanus Dejean, 1831 Ecology. Mountains. Borders of ponds (even temporary) and streams. Moist, gravelly soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day under stones and logs. Biology. Seasonality: January-December. Oviposition: May and July-August (AZ). Eggs are laid within 1 m from water. Tenerals: November (AZ). Adult food, in captivity: adult and larval flies, injured insects, other small arthropods, dead salamander larvae, and meat. The larva is an ectoparasite of dytiscid (Eretes) and hydrophilid (Tropisternus) pupae. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Fast runner. Collecting techniques. Pitfall trapping; turning logs and stones; light trapping. References. Erwin, 1970b (biology, dispersal power) and 1972 (biology); Juliano, 1985a, 1986a, and 1986b (biology).

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Brachinus (Neobrachinus) microamericanus Erwin, 1969 Ecology. Unknown. Biology. Seasonality: May, August. Dispersal power. Brachypterous, incapable of flight. Brachinus (Neobrachinus) mobilis Erwin, 1970 Ecology. Unknown. Biology. Seasonality: March. Dispersal power. Macropterous, probably capable of flight. Brachinus (Neobrachinus) neglectus LeConte, 1844 Ecology. Lowlands. Biology. Seasonality: March-October. Tenerals: May (GA). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Reference. Erwin, 1970b (biology, dispersal power). Brachinus (Neobrachinus) ovipennis LeConte, 1862 Ecology. Lowlands. Pond margins (mostly) and cultivated fields (e.g., corn, wheat, oat). Open ground; moist soil. Nocturnal; sheltering during the day under debris. Gregarious. Biology. Seasonality: March-October. Tenerals: April (NY), September (IN, KS, NY). Adults found overwintering on higher and drier ground, in vacant lots; under plant rosettes (Verbascum). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Collecting techniques. Pitfall trapping; turning debris. References. Erwin, 1970b (natural history); Kirk, 1975c (ecology, dispersal power). Brachinus (Neobrachinus) oxygonus Chaudoir, 1843 Ecology. Lowlands. Biology. Seasonality: January-July. Defense-mechanism: when alarmed, the adult may emit a repugnant audible spray from the pygidial glands. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Collecting technique. Light trapping. Brachinus (Neobrachinus) pallidus Erwin, 1965 Ecology. Margins of intermittent streams and permanent rivers. Wet soil consisting of coarse gravel. Nocturnal; sheltering during the day. Biology. Seasonality: May-September, December. Copulating pairs: May-September (CA); coitus lasting a few seconds; copulation position lasting an hour or more. Each egg is deposited onto a damp rock, near a stream or pond. The larva is an ectoparasite of hydrophilid pupae (Tropisternus, Berosus). Dispersal power. Macropterous. Occasional flier (in captivity). Fast runner. Collecting technique. Pitfall trapping. References. Erwin, 1966, 1967 (biology), and 1970b (natural history).

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Brachinus (Neobrachinus) patruelis LeConte, 1844 Ecology. Lowlands. Biology. Seasonality: April-September. Tenerals: May (MA). Dispersal power. Submacropterous, probably incapable of flight. Reference. Erwin, 1970b (biology, dispersal power). Brachinus (Neobrachinus) perplexus Dejean, 1831 Ecology. Lowlands. Biology. Seasonality: February, May-August, October. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Brachinus (Neobrachinus) phaeocerus Chaudoir, 1868 Ecology. Mountains. Margins of streams and eutrophic marshes (Carex). Nocturnal; sheltering during the day. Biology. Seasonality: February-September. Tenerals: May (TX), June (OK). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Collecting technique. Pitfall trapping. Reference. Erwin, 1970b (natural history). Brachinus (Neobrachinus) puberulus Chaudoir, 1868 Ecology. Unknown. Biology. Seasonality: May. Dispersal power. Macropterous, probably capable of flight. Brachinus (Neobrachinus) quadripennis Dejean, 1825 Ecology. Lowlands and mountains. Mostly lake shores and pond margins (near rivers); also cultivated fields (e.g., corn). Moist soil. Nocturnal; sheltering during the day. Gregarious. Biology. Seasonality: January-September. Tenerals: March (FL), August (MI, WA). Defense-mechanism: when alarmed, the adult may emit a repugnant audible spray from the pygidial glands. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Fast runner. Collecting techniques. Pitfall trapping; light trapping. References. Erwin, 1970b (natural history); Kirk, 1975c (ecology, dispersal power). Brachinus (Neobrachinus) rugipennis Chaudoir, 1868 Ecology. Lowlands and mountains. Margins of lakes, small streams, and irrigation canals. Moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: January-July, October, December. Tenerals: March (FL), October (MA), December (FL). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Fast runner. Collecting techniques. Pitfall trapping; light trapping. Reference. Erwin, 1970b (natural history). Brachinus (Neobrachinus) sublaevis Chaudoir, 1868 Ecology. Lowlands. Borders of eutrophic marshes (Scirpus, Typha); flood-plain forests. Open or half-shaded ground; wet, clayish or muddy soil. Nocturnal; sheltering during

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the day in clumps of grass. Gregarious. Biology. Seasonality: January-August. Adults found overwintering in woods; in standing rotten tree-stumps and under the loose bark of fallen trees. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Fast runner. Collecting techniques. Pitfall trapping; light trapping. References. Erwin, 1970b (natural history); Kirk, 1975c (dispersal power). Brachinus (Neobrachinus) tenuicollis LeConte, 1844 Ecology. Lowlands and mountains. Margins of rivers, ponds, reservoirs, pools (in sand pits and gravel pits), and marshes; flood-plain forests and swamps. Open or half-shaded ground; wet, clayish or muddy soil. Nocturnal; sheltering during the day under stones and vegetal debris. Gregarious. Biology. Seasonality: March-November. Tenerals: August (ON). Adults found overwintering on higher and drier ground, in woods and on sandy or gravelly hillocks; in the soil (16-25 cm deep) and under well-embedded stones. Adult food, in captivity: mealworms. The larva is an ectoparasite of hydrophilid (Hydrophilus) pupae. Parasites: Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult may emit a repugnant audible spray from the pygidial glands; the beetle is able to produce over 20 consecutive discharges. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Fast runner. Strong burrower. Collecting techniques. Pitfall trapping; turning stones; light trapping. References. Eisner, 1958 (biology; as ballistarius LeConte); Lindroth, 1969a (biology); Erwin, 1970b (natural history); Larochelle, 1974a (ecology). Brachinus (Neobrachinus) texanus Chaudoir, 1868 Ecology. Lowlands. Biology. Seasonality: March-November. Tenerals: July (TX). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Collecting technique. Light trapping. Reference. Erwin, 1970b (biology, dispersal power). Brachinus (Neobrachinus) velutinus Erwin, 1965 Ecology. A marsh (Typha, Scirpus). Wet soil. Biology. Seasonality: March-May, September. Dispersal power. Macropterous, probably capable of flight. Reference. Erwin, 1970b (natural history). Brachinus (Neobrachinus) viridipennis Dejean, 1831 Ecology. Lowlands. Wet meadows. Nocturnal; sheltering during the day under logs. Biology. Seasonality: February-April, June-July, September-December. Dispersal power. Submacropterous, probably incapable of flight. Fast runner. Collecting techniques. Lifting logs; baiting traps with banana.

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Brachinus (Neobrachinus) vulcanoides Erwin, 1969 Ecology. Lowlands. Biology. Seasonality: March-May, July-November. Defense-mechanism: when alarmed, the adult may emit a repugnant audible spray from the pygidial glands. Dispersal power. Macropterous, probably capable of flight. Bradycellus (Stenocellus) ardelio (Casey, 1914) Natural History. Unknown. Bradycellus (Stenocellus) aridus (Casey, 1914) Natural History. Unknown Bradycellus (Triliarthrus) atrimedeus (Say, 1823) Ecology. Lowlands. Borders of brooks and small slow rivers; margins of pools and ponds (e.g., in sand pits). Open ground; soft, wet, sandy soil, slightly mixed or covered with silt, somewhat mixed with dead leaves, covered with sparse vegetation (e.g., Equisetum). Mostly nocturnal; active in the spring sunshine; usually sheltering during the day in the soil (mostly; as deep as 8 cm), dead leaves, clumps of grass, and under stones. Biology. Seasonality: January, March-December. Tenerals: mid-August (VT). Adults found overwintering on higher and drier ground, in the upper zone of river banks and sand pits, and at the edge of woods; under dead leaves, depressed grasses, logs, and in the soil (as deep as 27 cm). Predators: flycatchers. Defense-mechanism: body color matching the soil. Dispersal power. Macropterous. Frequent flier (in the spring sunshine; to artificial lights in captivity). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Favored by spring floods along large rivers. Collecting techniques. Pouring water over the ground; treading the soil with the feet. References. Larochelle, 1972b (ecology); Erwin, 1981 (natural history). Bradycellus (Triliarthrus) badipennis (Haldeman, 1843) Ecology. Lowlands, mountains, and subalpine zone. Deciduous woodland swamps. Close to water (e.g., pools). Ground shaded by bushes (e.g., Alnus, Salix) and trees (e.g., Acer); wet, often humus-rich soil covered with thick leaf litter. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under dead leaves and heaps of grass. Gregarious. Biology. Seasonality: January-August, October-December. Gravids: early May (QU). Tenerals: early July (WV), August and November (QU). Adults found overwintering on higher and drier ground, in woods (mostly), at the edge of woods, in the upper zone of sand pits, gravel pits, sandy hillocks, and roadsides; under dead leaves (mostly), heaps of hay, in moss, under logs, and other debris. Adult food, in captivity: corn kernels, earthworms, and pieces of meat. Dispersal power. Macropterous. Frequent flier (in the sunshine in the spring; at sunset; to artificial lights in captivity; into rotary traps). Frequent in seashore drift material, which indicates flight ability.

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Moderate runner. Occasional climber (on plants and trees). Collecting techniques. Raking the leaf litter; turning drift material; sweeping; using rotary traps. Reference. Erwin, 1981 (natural history). Bradycellus (Stenocellus) californicus (LeConte, 1857) Ecology. Lowlands and mountains. Borders of pools, ponds, eutrophic marshes, lakes, reservoirs, slow rivers, and irrigation canals; mud flats, wet meadows, and vacant lots. Open ground; wet, muddy, sandy, or gravelly soil covered with some vegetation (e.g., grass, Juncus, Equisetum, Typha). Nocturnal; sheltering during the day under dead grass and stones. Gregarious. Biology. Seasonality: February-April, June-July. Tenerals: July (BC, ID, MT, OR, WA). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Treading the soil with the feet; pouring water over the ground; hanging yellow sticky boards in vineyards. Reference. Lindroth, 1968 (natural history). Bradycellus (Stenocellus) carolinensis (Casey, 1924) Natural History. Unknown Bradycellus (Triliarthrus) conformis (Fall, 1905) Ecology. Lowlands and mountains. Borders of ponds, lakes, reservoirs, and slow rivers; mud flats (in fields) and open low forests. Frequently near water. Open ground; rather dry, sandy soil covered with discontinuous vegetation (e.g., grass). Nocturnal; sheltering during the day under plant mats (e.g., grass, Ludwigia) and stones. Biology. Seasonality: July-August. Tenerals: late July (WA). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pitfall trapping. References. Lindroth, 1968 (natural history); LaBonte and Johnson, 1989 (ecology). Bradycellus (Stenocellus) congener (LeConte, 1848) Ecology. Lowlands and mountains. Borders of marshes, ponds (e.g., in sand pits and gravel pits), lakes, reservoirs, cattle water holes, slow rivers and brooks; swamps; pastures, vacant lots, cultivated fields and their borders (e.g., wheat, barley), and roadsides. Open ground; somewhat wet, clayish, often saline soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day in cracks in the soil, under dead leaves and dead plants, pieces of wood, dry cow dung, and stones. Biology. Seasonality: February-September, November. Gravids: late July (AB). Tenerals: April (rarely; QU), June to August; mostly from mid-June on. Adults found overwintering on higher and drier ground, on hillocks and hills, in pastures, and at the edge of woods; under stones and dead leaves. Active at low temperatures under snow in October-November. Adult food, in the field: coccinellids. Dispersal power. Macropterous. Frequent flier (at sunset; to artificial lights in captivity). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner.

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Moderately favored by human activities. Collecting techniques. Treading the soil with the feet; pitfall trapping; light trapping; turning drift material. References. Lindroth, 1968 (natural history); Bell, 1971 (ecology); Kirk, 1975c (dispersal power); Will et al. 1995 (dispersal power). Bradycellus (Liocellus) curticollis (Casey, 1924) Natural History. Unknown. Bradycellus (Stenocellus) decorus (Casey, 1914) Natural History. Unknown. Bradycellus (Stenocellus) discipulus (Casey, 1914) Natural History. Unknown. Bradycellus (Stenocellus) exstans (Casey, 1914) Natural History. Unknown. Bradycellus (Bradycellus) fenderi Hatch, 1951 Ecology. Unknown. Biology. Seasonality : January-May, October-December. Adults found overwintering in duff (Acer, Alnus) and moss. Dispersal power. Brachypterous, incapable of flight. Collecting technique. Using berlese funnels. Reference. Nelson, 1988 (biology). Bradycellus (Stenocellus) festinans (Casey, 1914) Ecology. Lowlands and mountains. Biology. Seasonality: May-August. Dispersal power. Unknown. Bradycellus (Triliarthrus) georgei Lindroth, 1968 Ecology. Borders of a lake, among Typha and Carex, with wet soil. Biology. Seasonality: May, July. Dispersal power. Macropterous, probably capable of flight. Bradycellus (Bradycellus) harpalinus (Audinet-Serville, 1821) Ecology. Lowlands. Disturbed habitats: gravel pits, vacant lots, and roadsides. Open ground; moderately dry, sandy or gravelly soil covered with weeds. Diurnal and nocturnal; active during the day on plants or sheltering under dead leaves. Biology. Seasonality: June-July. Tenerals: June (BC). Adult food, in the field: grass seeds (Europe). Dispersal power. Wing-dimorphic: macropterous (mostly; capable of flight) or brac-

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hypterous (incapable of flight). Mass flights observed in late summer (Europe). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Regular climber (on plants; Europe). Clearly effective as a colonist; quick to establish in recently modified habitats. Strongly favored by human activities. Collecting techniques. Pitfall trapping; raking the leaf litter; sweeping plants; using window traps. References. Lindroth, 1945 and 1968 (natural history); Boer, 1980 (biology); Lindroth, 1986 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology). Bradycellus (Stenocellus) humboldtianus (Casey, 1924) Natural History. Unknown. Bradycellus (Stenocellus) insulsus (Casey, 1914) Ecology. Lowlands. Borders of ponds and slow rivers. Open ground; moist, clayish or muddy soil covered with some vegetation. Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: February, June-July. Adults found overwintering in vacant lots; under dead leaves. Dispersal power. Macropterous. Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pouring water over the ground; raking the leaf litter. Bradycellus (Liocellus) intermedius (Fall, 1905) Ecology. Mountains. Biology. Seasonality: August. Dispersal power. Unknown. Bradycellus (Triliarthrus) kirbyi (G.H. Horn, 1883) Ecology. Lowlands. Borders of eutrophic marshes, slow rivers, pools and ponds (e.g., in sand pits and gravel pits); swamps, low deciduous forests, flood-plain forests, and moist meadows. Open or shaded ground; moist or wet, clayish or muddy soil covered with some vegetation. Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering during the day in the soil (mostly; as deep as 8 cm), under dead leaves, stones, and in moss. Defense-mechanism: body color-pattern matching the soil. Biology. Seasonality: February, April-December. Tenerals: September-October (QU). Adults found overwintering on higher and drier ground, at the edge of woods and in woods; under dead leaves, pieces of wood, stones, in moss and rotten logs, and under the loose bark of logs. Predators: frogs. Dispersal power. Macropterous. Regular flier (to artificial lights in captivity; into rotary traps and malaise traps). Moderate runner. Occasional climber (on plants). Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet; using rotary traps; using malaise traps; sweeping plants. References. Lindroth, 1968 (natural history); Larochelle, 1974a (ecology). Bradycellus (Stenocellus) larvatus (Casey, 1914) Natural History. Unknown.

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Bradycellus (Liocellus) laticollis (Casey, 1924) Natural History. Unknown. Bradycellus (Catharellus) lecontei Csiki, 1932 Ecology. Lowlands and mountains. Borders of marshes, pools (even temporary), ponds, lakes, irrigation canals, and roadside ditches; swamps, marshy meadows, and cultivated fields (e.g., barley). Open ground; somewhat moist, mostly clay soil covered with discontinuous cover of depressed plants (e.g., grass, Carex, Juncus) and sometimes polluted by refuse. Nocturnal; sheltering during the day under dead leaves, other organic debris, and stones. Gregarious (in winter). Biology. Seasonality: January, March-November. Tenerals: late July (NF) to late September (QU). Adults found overwintering on higher and drier ground, in the upper zone of brook and marsh borders, at the edge of woods, and in woods; under dead leaves and stones. Active at low temperatures under snow in October-November. Dispersal power. Macropterous. Frequent flier (at sunset; to artificial lights at night; to artifical lights in captivity; into rotary traps). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Moderately favored by human activities and spring floods along large rivers. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping; using rotary traps; sweeping plants. References. Lindroth, 1955a, 1963a, and 1968 (natural history); Kirk, 1975c (ecology, dispersal power). Bradycellus (Stenocellus) lineatus (Casey, 1914) Natural History. Unknown. Bradycellus (Triliarthrus) lugubris (LeConte, 1848) Ecology. Lowlands, mountains, and subalpine zone. Deciduous woodland swamps (close to water); low deciduous forests; borders of eutrophic marshes, brooks, and roadside ditches; moist meadows. Ground shaded by bushes and trees (e.g., Alnus, Viburnum, Acer); moderately moist or wet soil, often rich in humus. Nocturnal; sheltering during the day under dead leaves (mostly), in humus, heaps of grass, and under stones. Biology. Seasonality: March-November. Copulating pairs: mid-May (QU). Gravids: early May (QU). Tenerals: early July (WV), November (QU). Adults found overwintering on higher and drier ground, in woods, at the edge of woods, and in the upper zone of roadside ditches; under dead leaves and stones. Adult and larval food, in captivity: mealworms. Predators: spiders. Dispersal power. Macropterous. Occasional flier (at sunset; to artificial lights in captivity). Frequent in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Collecting techniques. Raking the leaf littter; turning drift material; sweeping plants. References. Lindroth, 1955a (as protractus Casey), 1963a, and 1968 (natural history).

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Bradycellus (Stenocellus) lustrellus (Casey, 1914) Natural History. Unknown. Bradycellus (Stenocellus) montanus (Casey, 1914) Natural History. Unknown. Bradycellus (Stenocellus) nebulosus LeConte, 1853 Ecology. Unknown. Biology. Seasonality: June. Dispersal power. Unknown. Bradycellus (Stenocellus) neglectus (LeConte, 1848) Ecology. Lowlands, mountains, and subalpine zone. Moraines, gravel pits, vacant lots, ski fields, sand pits, roadsides, and cultivated fields (e.g., alfalfa, cabbage). Open ground; dry, gravelly soil covered with sparse low vegetation. Nocturnal; sheltering during the day under stones, dead leaves and dead plants (Gramineae), logs, and at the base of plants. Gregarious (in winter). Biology. Seasonality: March-December. Tenerals: April (rarely; QU), late June to late September; mostly from mid-July on). Adults found overwintering on higher and drier ground, at the edge of woods and in woods; under stones, pieces of wood, and in the soil (as deep as 2.5 cm). Dispersal power. Macropterous. Regular flier (at sunset; to artificial lights at night; to artifical lights in captivity). Moderate runner. Occasional climber (on plants). Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; light trapping; sweeping plants. References. Lindroth, 1955a, 1963a, and 1968 (natural history); Kirk, 1975c (dispersal power). Bradycellus (Stenocellus) nigerrimus Lindroth, 1968 Ecology. Lowlands and mountains.Vicinity of sloughs, marshes, ponds, lakes, slow rivers and brooks; moist pastures, cultivated fields and their borders (e.g., wheat). Open ground; moist or wet, clayish, often alkaline soil covered with some vegetation (e.g., Eleocharis, Carex, Juncus, grass). Nocturnal; sheltering during the day under vegetal debris and dry cow dung. Biology. Seasonality: April-August. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping; turning vegetal debris. Reference. Lindroth, 1968 (natural history). Bradycellus (Stenocellus) nigriceps LeConte, 1868 Ecology. Lowlands. Wet meadows, salt meadows; borders of marshes, slow brooks and rivers; swamps, roadside ditches, and cultivated fields (e.g., carrot). Open ground; moist or wet, clayish or muddy, often saline soil covered with dense vegetation (e.g., grass, Scirpus). Mostly nocturnal; active in the spring sunshine; usually sheltering during the

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day under vegetal debris (mostly; e.g., dead plants, heaps of hay), in moss, under stones, and at the base of plants. Gregarious (in winter). Biology. Seasonality: February-December. Adults found overwintering on higher and drier ground, in the upper zone of sand dunes and roadsides, in vacant lots, at the edge of woods, and in woods; clinging upside down under dead leaves, stones, pieces of wood, the loose bark of fallen treestumps and trees, in rotten logs, under plant rosettes (Verbascum), and buried in the soil (as deep as 7.5 cm). Adult food, in captivity: meat. Predators: frogs and toads. Dispersal power. Macropterous. Frequent flier (in the spring sunshine; to artificial lights at night; to artifical lights in captivity; into rotary traps). Frequent in lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants and trees). Moderately favored by human activities and spring floods along large rivers. Collecting techniques. Treading the vegetation into the water; pouring water over the ground; light trapping; turning drift material; sweeping plants; using rotary traps. References. Lindroth, 1968 (natural history); Larochelle, 1974a (ecology); Will et al. 1995 (dispersal power). Bradycellus (Lipalocellus) nigrinus (Dejean, 1829) Ecology. Lowlands, mountains, subalpine and alpine zones. Wet meadows, borders of marshes; swamps; marshy borders of lakes, ponds, pools, slow rivers and brooks, and roadside ditches; mud flats in sand pits, low forests or thickets bordering water bodies. Open or shaded ground; soil rather moist (continental parts of the area) or dry (Pacific Coast grasslands), often rich in humus, covered with discontinuous vegetation (e.g., Carex, Juncus, grass). Mostly nocturnal; active in the spring sunshine; usually sheltering during the day under dead leaves (mostly) and dead plants, fallen branches, and stones. Biology. Seasonality: January, March-November. Gravids: late June and early July (QU). Tenerals: April (rarely; QU), early July to October; mostly July and August. Adults found overwintering on higher and drier ground, in swamps, in woods, at the edge of woods, in hawthorn hedges, and in vacant lots; under dead leaves (mostly), stones, in moss, and under the loose bark of fallen trees. Adult and larval food, in captivity: mealworms. Predators: spiders. Dispersal power. Macropterous. Frequent flier (in the spring sunshine; at sunset; to artificial lights at night; to artifical lights in captivity; into rotary traps). Moderate runner. Occasional climber (on plants). Collecting techniques. Treading the vegetation into the water; pouring water over the ground; light trapping; using rotary traps; sweeping plants. References. Lindroth, 1955a, 1963a, and 1968 (natural history); Kavanaugh, 1992 (natural history); Levesque and Levesque, 1994 (biology); Ball and Bousquet, 2001 (taxonomy). Bradycellus (Liocellus) nitidus (Dejean, 1829) Ecology. Lowlands and mountains. On the gravelly banks of a stream (CA). On the moist bank of an arroyo, among grasses (CA). Pastures, forests (e.g., Pinus, Quercus), and disturbed places (Mexico). Nocturnal; sheltering during the day. Biology. Seasonality: February-December. Dispersal power. Macropterous. Occasional in drift material, which indicates flight ability. Moderate runner.

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Bradycellus (Stenocellus) nubifer LeConte, 1858 Ecology. Lowlands and mountains. Borders of slow rivers and brooks, lakes, reservoirs, and marshes. Open ground; moist or wet, sandy, gravelly, or muddy soil covered with sparse vegetation. Nocturnal; sheltering during the day in cracks in the soil. Biology. Seasonality: June-September. Tenerals: early July to mid-September; mostly July. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with feet. Reference. Lindroth, 1968 (natural history). Bradycellus (Liocellus) obtusus (Fall, 1905) Natural History. Unknown. Bradycellus (Stenocellus) picipes (Casey, 1914) Natural History. Unknown. Bradycellus (Liocellus) politus (Fall, 1905) Ecology. Mountains. Biology. Seasonality: April, June-September. Dispersal power. Macropterous, probably capable of flight. Bradycellus (Stenocellus) provoensis (Casey, 1914) Natural History. Unknown. Bradycellus (Stenocellus) puncticollis (Casey, 1914) Ecology. Unknown. Biology. Seasonality: January, May. Dispersal power. Unknown. Bradycellus (Stenocellus) purgatus (Casey, 1914) Ecology. Lowlands and mountains. Biology. Seasonality: spring-autumn. Dispersal power. Unknown. Bradycellus (Stenocellus) rivalis LeConte, 1858 Natural History. Unknown. Bradycellus (Stenocellus) rupestris (Say, 1823) Ecology. Lowlands and mountains. Vacant lots, meadows, pastures, cultivated fields (e.g., cabbage, cotton, tobacco, alfalfa), ski fields, sand pits, roadsides, and orchards (e.g., peach, apple). Open ground; rather dry, gravelly soil covered with sparse vegetation. Mostly noc-

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turnal; active in the spring sunshine; usually sheltering during the day under stones and dead leaves (mostly), pieces of wood and cardboard, heaps of hay and grass, and at the base of plants (Gramineae). Gregarious (in winter). Larval habitat: in the soil (IN). Biology. Seasonality: January-December. Tenerals: April-May (rarely; QU), late June to early September. Adults found overwintering on higher and drier ground, in fields, in the upper zone of hills, hillocks, gravel pits, dunes, and roadsides, at the edge of woods, and in woods; under stones and dead leaves (mostly), logs, in fallen cotton balls, rotten logs, and in the soil (as deep as 50 cm). Adult food, in the field: small worms. Predators: toads, blackbirds, flycatchers, swallows, and towhees. Parasites: tapeworms and Laboulbeniales (Fungi). Phoretic mites found on adult body. Dispersal power. Macropterous. Frequent flier (in the spring sunshine; to artificial lights at night; into rotary traps). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants and trees). Strongly favored by human activities and spring flood along large rivers. Collecting techniques. Pitfall trapping; turning debris; using an aspirator; light trapping; using rotary traps; sweeping; baiting traps with molasses and fermented sugar. References. Blatchley, 1910 (dispersal power); Lindroth, 1968 (natural history); Kirk, 1969 (ecology), 1970 (dispersal power) and 1975c (ecology, dispersal power); Erwin, 1981 (natural history). Bradycellus (Stenocellus) sejunctus (Casey, 1914) Natural History. Unknown. Bradycellus (Lipalocellus) semipubescens Lindroth, 1968 Ecology. Lowlands, mountains, and subalpine zone. Deciduous woodland swamps (often along brooks and pools); borders of eutrophic marshes; low forests, wet meadows, and roadside ditches. Ground shaded by bushes (e.g., Alnus, Salix, Acer); wet soil, often rich in humus. Mostly nocturnal; quite active in the spring sunshine; usually sheltering during the day under dead leaves, clumps of grass, and heaps of hay. Gregarious. Biology. Seasonality: March-November. Gravids: early June (QU). Tenerals: late July (QU). Adults found overwintering on higher and drier ground, in woods, at the edge of woods, in vacant lots, in the upper zone of hills, hillocks, and roadsides; under stones and dead leaves (mostly), and in moss. Adult food, in captivity: corn kernels. Predators: toads. Dispersal power. Macropterous. Frequent flier (in the spring sunshine; at sunset; to artificial lights at night; to artifical lights in captivity). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Raking the leaf litter; light trapping; sweeping. References. Larochelle, 1973b (ecology); Ball and Bousquet, 2001 (taxonomy). Bradycellus (Stenocellus) suavis (Casey, 1914) Natural History. Unknown. Bradycellus (Stenocellus) subcordatus Chaudoir, 1868 Natural History. Unknown.

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Bradycellus (Stenocellus) supplex (Casey, 1914) Natural History. Unknown. Bradycellus (Stenocellus) symetricus (Motschulsky, 1850) Natural History. Unknown. Bradycellus (Liocellus) tahoensis (Casey, 1924) Natural History. Unknown. Bradycellus (Stenocellus) tantillus (Dejean, 1829) Ecology. Lowlands. Borders of eutrophic marshes, ponds, lakes, and slow brooks; mud flats. Open ground; wet, clayish or muddy soil covered with dense vegetation (e.g., Juncus). Mostly nocturnal; usually sheltering during the day under debris and in the soil. Biology. Seasonality: January-October, December. Tenerals: February (rarely; AL), July (AL, WV). Adults found overwintering on higher and drier ground, in sand pits and vacant lots; under dead leaves and in the soil (as deep as 3.5cm) at the base of plants. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; sometimes during the day). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on trees). Strong burrower. Collecting techniques. Treading the vegetation into the water; light trapping; baiting traps with molasses and fermented sugar; beating trees. References. Lindroth, 1968 (dispersal power); Kirk, 1970 (biology); Will et al. 1995 (dispersal power). Bradycellus (Stenocellus) veronianus (Casey, 1924) Natural History. Unknown. Broscodera insignis (Mannerheim, 1852) Ecology. Lowlands, mountains, subalpine and alpine zones. Below tree line: margins of small streams, with wet, loose stony or rocky soil. Above tree line: barren slopes and edges of snowfields. Open ground. Nocturnal; sheltering during the day in loose rocky substrate (below tree line) or under stones (above tree line). Gregarious. Biology. Seasonality: July-August. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pitfall trapping. References. Lindroth, 1961 (natural history); Kavanaugh, 1992 (natural history). Broscus cephalotes cephalotes (Linné, 1758) Ecology. Coastal lowlands. Sand dunes and rear beaches. Open ground; well-drained, dry, sandy soil, bare or soil covered with sparse vegetation. Nocturnal; sheltering dur-

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ing the day in burrows (as deep as 12 cm) dug in the soil under logs and pieces of wood. Gregarious. Larval habitat: same as adult. Biology. Seasonality: July. Copulating pairs: July (NS). Eggs are laid in deep shafts into the soil (Europe). Tenerals: mid- and late July (NS, PE). Adult food, in the field: flies, beetles, amphipods (Talitrus), and other arthropods (Europe); the opening of a beetle burrow is frequently strewn with remains of his prey. The larva is also carnivorous. Defense-mechanism: when alarmed, the adult may bite strongly or discharge a defensive fluid from the pygidial glands. The females protects her brood. Dispersal power. Macropterous; probably incapable of flight (with elytra fused anteriorly). Slow runner. Strong burrower. Clearly effective as a colonist. Collecting techniques. Lifting logs; pitfall trapping. References. Lindroth, 1945 (natural history); Kempf, 1955 (biology); Sharova, 1981 (ecology, biology); Lindroth, 1985 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology). Calathus (Procalathus) advena (LeConte, 1848) Ecology. Lowlands, mountains, subalpine and alpine zones. Forests (preferably mixed and deciduous; also coniferous); forest edges and adjoining meadows (subalpine and alpine zones); vicinity of snowfields, lakes and streams. Shaded (mostly) or open ground; moderately moist soil, rich in organic debris or devoid of vegetation. Nocturnal; sheltering during the day deep among moss or in leaf litter (mostly), under logs, stones, and the loose bark of fallen trees and logs. Gregarious. Biology. Seasonality: May-October. Gravids: late June to mid-July (QU). Tenerals: June-October; mostly July-August. Adult and larval food, in captivity: mealworms. Predators: toads. Dispersal power. Macropterous. Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strong burrower. Occasional climber (on trees; up to 2-3 m above ground). Collecting techniques. Pitfall trapping; beating trees. Reference. Lindroth, 1955a, 1963a, and 1966 (natural history). Calathus (Neocalathus) calceus Ball and Nègre, 1972 Ecology. Uplands. A dry forest (Populus-Abies; ID). Vicinity of a small stream, on moist soil (UT). Biology. Seasonality: March, May-September. Dispersal power. Wing-dimorphic: macropterous (probably capable of flight) or brachypterous (incapable of flight). Moderate runner. Calathus (Calathus) fuscipes (Goeze, 1777) Ecology. Lowlands. Cultivated fields, greasslands, meadows, vacant lots, roadsides, and vicinity of human habitations. Greenhouses (occasionally). Open (mostly) or slightly shaded ground; rather dry, sandy or clayish soil covered with sparse vegetation (e.g., grass, weeds). Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: April-October. Tenerals: May (BC); early summer (Europe). Adult food, in the field (Europe): aphids, lepidopterous caterpillars, ants, cereals, and forest seeds. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Moderate runner. Clearly effective as a colonist. Strongly favored by human activ-

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ities. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Lindroth, ° 1945 and 1966 (natural history); Greenslade,1965 (ecology, biology); Kurka, 1972 (biology); Finlayson and Campbell, 1976 (biology); Sharova, 1981 (ecology, biology); Lindroth, 1986 (natural history); Hurkå, 1986b (biology); Spence and Spence, 1988 (dispersal power); Koch, 1989 (ecology); Luff, 1993 (biology). Calathus (Neocalathus) gregarius (Say, 1823) Ecology. Forests (mostly decidous; mixed, coniferous), tree plantations (e.g., Pinus), thickets, forest clearings and edges, hedges, orchards, cultivated fields (e.g., corn, oat, wheat, hay, tobacco, raspberry), grasslands, meadows, old fields, vacant lots, fencerows, sand pits, gravel pits, and roadsides. Shaded (mostly) or open ground; dry, often sandy soil covered with thick leaf litter or sparse vegetation (e.g., grass, herbs). Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering during the day in leaf litter and under stones (mostly), under pieces of wood and logs. Gregarious. Biology. Seasonality: January-December. Copulating pairs: July (QU). Tenerals: May-August; mostly before mid-summer. Adults found overwintering on higher and drier ground, in woods, at the edge of woods, on hills, in vacant lots and pastures, and in the upper zone of gravel pits; under stones and dead leaves (mostly), logs, plant rosettes (Verbascum), in rotten tree-stumps, under dead plants, and deep in the soil (as deep as 35 cm). Adult food: sawfly cocoons, lepidopterous caterpillars, and grass pollen (e.g., Phleum) (in the field); mealworms and meat (in captivity). Larval food, in captivity: mealworms. Predators: toads (frequently), frogs, flycatchers, and robins. Defense-mechanism: when alarmed, the adult may emit a repugnant smoky spray from the pygidial glands. Dispersal power. Wingdimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (to artificial lights in captivity). Moderate runner. Occasional climber (on trees (e.g., Tsuga, Abies) and plants). Strongly favored by human activities. Collecting techniques. Pitfall trapping; raking the leaf litter; beating trees; night collecting with a headlamp. References. Lindroth, 1966 (natural history); Larochelle, 1972g (dispersal power); Esau and Peters, 1975 (biology); Kirk, 1975c (ecology); Liebherr and Mahar, 1979 (biology); Levesque et al. 1979 (biology); Erwin, 1981 (natural history); Epstein and Kulman, 1990 (ecology, biology); Dearborne and Donahue, 1993 (ecology, biology); Will et al. 1995 (dispersal power). Calathus (Neocalathus) ingratus Dejean, 1828 Ecology. Lowlands, mountains, subalpine and alpine zones. Forests (mostly coniferous; mixed, deciduous), forest clearings and edges, thickets, hedges, and adjacent roadsides, cultivated fields (e.g., wheat, barley, raspberry), vacant lots, hills, meadows, moraines, sand pits, gravel pits; spruce bogs, open grasslands and coastal tundra (e.g., NF, Aleutian Islands). Vicinity of human habitations (urban and rural sites). In the shade of decidous trees or bushes (mostly) or on open ground; moderately moist or rather dry, often gravelly soil, often covered with thick leaf litter or sparse vegetation (e.g., grass). Mostly nocturnal; usually sheltering during the day in leaf litter, moss and under stones (mostly), under reindeer moss (Cladonia), logs, fallen tree-stumps, branches, boards, and heaps of hay, in rotten logs, under the loose bark of standing rotten tree-stumps, on

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trees and plants. Gregarious. Biology. Seasonality: March-November. Gravids: late June to late August (AB, QU). Tenerals: May-November; mostly before mid-summer. Adults found overwintering on higher and drier ground, in woods; in the soil (as deep as 15 cm), in leaf litter, or in rotten tree trunks. Adult food, in captivity: lepidopterous caterpillars, mealworms, and shrimps. Larval food, in captivity: mealworms. Predators: toads (regularly), frogs, woodpeckers, and shrews. Parasites: mites. Dispersal power. Wingdimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional in seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (on trees, e.g., Picea, Abies; also on plants). Strongly favored by human activities. Collecting techniques. Pitfall trapping; raking the leaf litter; beating or sweeping the vegetation. References. Lindroth, 1955a, 1963a, and 1966 (natural history); Frank, 1971b (biology); Larochelle, 1972d (dispersal power); Ostaff and Freitag, 1973 (biology); Bousquet and Pilon, 1977 (biology); Reeves et al. 1983 (biology); Holliday, 1991 (biology); Niemelä and Spence, 1991 (ecology); Niemelä et al. 1992 (ecology, biology). Calathus (Neocalathus) melanocephalus (Linné, 1758) Ecology. Lowlands. Meadows, grasslands, dunes, heaths, and light forests (Europe). Moderately dry soil. Nocturnal; sheltering during the day. Biology. Seasonality: August. Adult food, in the field (Europe): insect eggs, spiders, and plant seeds. Dispersal power. Brachypterous, incapable of flight (North America). Wing-dimorphic: brachypterous (mostly) or macropterous (rarely; Europe). Fast runner. References. Lindroth, 1945 (natural history); Forsskåhl, 1972 (biology); Thiele, 1977 (natural history); Sharova, 1981 (ecology, biology); Lindroth, 1986 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology). Comments. Doubtfully established in North America. Calathus (Neocalathus) opaculus LeConte, 1854 Ecology. Lowlands, mountains, subalpine and alpine zones. Meadows, old fields, pastures, vacant lots, cultivated fields (e.g., corn, tobacco, soybean, alfalfa), lawns, hedges, sandhills, sand pits, orchards (e.g., peach, plum), forest edges, and open forests (e.g., Quercus). Open (mostly) or shaded ground; dry, sandy soil covered with sparse or patchy low vegetation (e.g., grass, herbs). Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering during the day under embedded stones, dead leaves, reindeer moss (Cladonia), in the soil at the base of plants (Solidago), under pieces of wood, in clumps of broomsedges, or on trees and plants. Semi-gregarious (summer); gregarious (winter). Biology. Seasonality: January-December. Tenerals: June-July (ON, QU). Adults found overwintering in fields, hedges, and woods; under well-embedded stones (mostly), plant rosettes (Verbascum), mats of dead fallen low vegetation, and in wood debris. Adult food, in captivity: ant eggs. Predators: toads. Dispersal power. Macropterous. Occasional flier (to lights at night). Fast runner. Occasional climber. Strong burrower. Strongly favored by human activities. Collecting techniques. Pitfall trapping; baiting traps with malt; turning embedded stones; night collecting with a headlamp. References. Lindroth, 1966 (ecology, dispersal power); Kirk, 1969 (biology); Los and Allen, 1983 (biology); Anderson et al. 1995 (ecology); Will et al. 1995 (dispersal power).

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Calathus (Neocalathus) peropacus Casey, 1920 Ecology. Mountains. Forests (deciduous, mixed, and coniferous). Shaded ground; moderately moist soil. Nocturnal; sheltering during the day in thick layers of dead leaves. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Strong runner. Collecting technique. Raking the leaf litter. Calathus (Neocalathus) ruficollis grandicollis Casey, 1920 Ecology. Lowlands. A dry, sandy ridge covered with some Artemisia and other xerophytes (BC). Biology. Seasonality: May, July. Tenerals: late May (BC). Dispersal power. Brachypterous, incapable of flight. Reference. Lindroth, 1966 (natural history). Calathus (Neocalathus) ruficollis ignicollis Casey, 1920 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Calathus (Neocalathus) ruficollis ruficollis Dejean, 1828 Ecology. Lowlands and mountains. Vicinity of forests, orchards (e.g., apple), streams, and lakes. Open ground; moderately dry soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day under stones. Biology. Seasonality: February, April, June-July, October. Adult food, in the field: lepidopterous caterpillars. Predators: flickers (woodpeckers) and spiders. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning stones; pitfall trapping. Calleida (Philophuga) caerulea (Casey, 1913) Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Calleida (Calleida) circumcincta Bates, 1883 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Calleida (Calleida) decora (Fabricius, 1801) Ecology. Lowlands. Grasslands, cultivated fields (e.g., cotton, soybean, peanut, okra, corn), pastures, old fields, vacant lots, fencerows, and open forests. Open ground; more or less vegetated soil. Mostly diurnal; active in the sunshine on plants (flowers, grass, weeds), shrubs (e.g., Sambucus), and trees; sheltering on cloudy or cool days in moss and under tree bark (Pinus). Biology. Seasonality: January-December. Each egg is laid singly into a silk-like purse attached to a leaf by a silken thread. Adults found overwintering in woods; under the loose bark of trees (Quercus, Pinus) and decayed tree-stumps, also in moss. Adult food, in the field and captivity: lepidopterous eggs and larvae, na-

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bid nymphs. Larval food, in the field and in captivity: lepidopterous eggs and small caterpillars. Larvae are cannibalistic. Economic importance: may play an important role in the control of lepidopterous pests (e.g., Anticarsia gemmatalis Hübner). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Frequent climber (on plants, shrubs, and trees). Strongly favored by human activities. Collecting techniques. Sweeping or beating the vegetation; pitfall trapping. References. Leng, 1915 (ecology, biology); Löding, 1945 (ecology); Whitcomb and Bell, 1964 (biology); Kirk, 1969 (natural history), 1970 (biology) and 1975c (ecology); McWorter et al. 1984 (biology); Harris et al. 1985 (biology); Fuller, 1988 (biology). Calleida (Calleida) fimbriata Bates, 1883 Natural History. Unknown. Calleida (Calleida) fulgida Dejean, 1831 Ecology. Lowlands. On low scrub oaks (Quercus) and bushes. Diurnal; active in the sunshine. Biology. Seasonality: March-June, August, November. Adults found overwintering under loose bark. Adult food, in the field: chrysomelid larvae. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Frequent climber (on trees and shrubs). Collecting techniques. Sweeping shrubs. References. Leng, 1915 (ecology); Löding, 1945 (ecology). Calleida (Calleida) obrieni Mateu, 1994 Ecology. Lowlands. Biology. Seasonality: May. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Reference. Mateu, 1994 (taxonomy). Calleida (Calleida) planulata LeConte, 1858 Ecology. A tropical jungle, on herbage (Mexico). Diurnal; active in the sunshine Biology. Seasonality: June-July. Dispersal power. Unknown. Calleida (Calleida) platynoides G.H. Horn, 1882 Ecology. Unknown. Biology. Seasonality: February-March. Dispersal power. Unknown. Calleida (Calleida) punctata LeConte, 1848 Ecology. Lowlands and mountains. Abandoned fields, meadows, pastures, cultivated fields and their borders (e.g., alfalfa), wastelands, marsh borders, swamps, roadsides, sand pits, dunes, forest clearings, and open forests. Open ground; moderately moist soil covered with some vegetation. Mostly diurnal; active in the sunshine on flowering plants, shrubs, and trees (mostly Solidago and Salix; also Acer, Alnus, Ambrosia, Artemisia, Asclepias, Barbarea, Cornus, Eupatorium, Oenethera, Prunus, Ranuncula, Spiraea, Taraxacum, Trifolium, Tsuga, Vibur-

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num, Vicia); sheltering on cloudy or cool days under stones. Biology. Seasonality: January, March-November. Copulating pairs: mid-June (QU). Gravids: late May and mid-June (QU). Tenerals: early August to mid-September. Adults found overwintering in fields, on hills, in pastures and hawthorn hedges, at the edge of woods, and in woods; under stones, dead leaves, plant rosettes (Verbascum), in the soil at the base of plants, and under the loose bark of standing dead trees (e.g., Pinus). Adult food, in the field: lepidopterous caterpillars (mostly) and chrysomelid nymphs. Predators: frogs, toads, and robins. Dispersal power. Macropterous. Occasional flier (in the bright sunshine; into rotary traps). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (on plants, shrubs, and trees). Collecting techniques. Sweeping or beating the vegetation; using rotary traps. References. Blatchley, 1910 (ecology); Larochelle, 1974a (ecology). Calleida (Calleida) punctulata Chaudoir, 1848 Ecology. Macrohabitat unknown. Diurnal; active in the sunshine on plants. Biology. Seasonality: July. Dispersal power. Macropterous. Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Sweeping or beating the vegetation. Calleida (Calleida) purpurea (Say, 1823) Ecology. Lowlands. Deciduous forests (e.g., Quercus) and dry tall-grass prairies. Diurnal; active in the sunshine on trees, shrubs, and grass (e.g., Anemone). Biology. Seasonality: January-September. Adults found overwintering in dead oaks (Quercus). Larval food, in captivity: lepidopterous caterpillars. Dispersal power. Macropterous. Occasional in seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (on foliage). Collecting techniques. Sweeping or beating the vegetation; pitfall trapping. Reference. Tinerella, 2000 (ecology). Calleida (Calleida) striata Casey, 1913 Natural History. Unknown. Calleida (Philophuga) viridicollis (LeConte, 1848) Ecology. Lowlands and mountains. Cultivated fields (alfalfa and other legumes). Open ground; soil covered with sparse or moderate vegetation. Diurnal; active in the sunshine. Biology. Seasonality: April-July (TX). Tenerals: April-July. Larval food, in the field: lepidopterous caterpillars. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pitfall trapping. Calleida (Calleida) viridipennis (Say, 1823) Ecology. Lowlands. Open forests and cypress-swamps. Mostly diurnal; active in the sunshine on trees; sheltering on cloudy or cool days under the loose bark and bark

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scales of trees (e.g., Pinus, Juglans, Acer, Salix). Biology. Seasonality: January-August, October. Adults found overwintering in woods; under the loose bark and bark scales of trees (e.g., Quercus), also in moss. Adult and larval food, in the field: lepidopterous caterpillars. Economic importance: may play an important role in the control of lepidopterous caterpillars (e.g., the Fruittree Leafroller, Archips argyrospila (Walker)). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Frequent climber (on trees). Collecting techniques. Beating trees; using tanglefoot screens. References. Leng, 1915 (ecology); Löding, 1945 (biology); Kirk, 1969 (ecology); Erwin, 1981 (natural history); Braun et al. 1990 (biology); Zhou and Goyer, 1993 (ecology, biology); Davidson, 1995 (dispersal power). Calleida (Philophuga) viridis amoena (LeConte, 1848) Ecology. Lowlands and mountains. Short-grass prairies; mixed-grass rangelands, and dry fields. Open ground; dry, clay soil covered with some vegetation (e.g., grass). Diurnal; active in the sunshine on plants (Artemisia, Purshia) and at their base (yuccas); sheltering on cloudy and cool days under stones and other debris. Biology. Seasonality: March-October. Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Regular climber (on plants). Collecting techniques. Sweeping plants; pitfall trapping. References. Larson, 1969 (natural history); Lavigne, 1977 and 1978 (ecology). Calleida (Philophuga) viridis horni (Chaudoir, 1877) Ecology. Uplands. Desert scrubs and shrub-steppes. Open ground; dry soil. Diurnal; active in the sunshine. Biology. Seasonality: April-May, July. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Larson, 1969 (ecology, dispersal power). Calleida (Philophuga) viridis klamathea (Larson, 1969) Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Reference. Larson, 1969 (dispersal power). Calleida (Philophuga) viridis viridis (Dejean, 1831) Ecology. Unknown. Biology. Seasonality: December. Dispersal power. Brachypterous, incapable of flight. Reference. Larson, 1969 (dispersal power). Calosoma (Chrysostigma) affine Chaudoir, 1843 Ecology. Lowlands and mountains. Pastures, cultivated fields (e.g., alfalfa), and sand dunes. Open ground; sandy soil covered with some vegetation. Diurnal and nocturnal; sometimes sheltering during the day in the soil. Gregarious (sometimes swarming). Biology. Seasonality: April, June-September. Gravids: July-August (CA). Adult food, in the field

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and in captivity: lepidopterous caterpillars. Economic importance: may play an important role in the control of lepidopterous caterpillars in cultivated fields. Defense-mechanism: when alarmed, the adult may emit a defensive fluid from the pygidial glands. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Occasional climber (on plants). Collecting techniques. Pitfall trapping; sweeping plants. References. McCullough, 1966b (biology); Wallin, 1991 (biology, dispersal power). Calosoma (Carabosoma) angulatum Chevrolat, 1834 Ecology. Unknown. Biology. Seasonality: May-October. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Reference. Ball and Bousquet, 2001 (taxonomy). Coillecting technique. Light trapping. Calosoma (Calodrepa) aurocinctum Chaudoir, 1850 Ecology. Lowlands. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of fligh. Calosoma (Chrysostigma) calidum (Fabricius, 1775) (Common names: The Fiery Hunter (adult); Cutworm Lion (larva)) Ecology. Lowlands and mountains. Meadows, pastures, rangelands, cultivated fields (e.g., corn, soybean, oat, cotton, wheat, barley, alfalfa, tobacco, hop, flax, clover, potato), gardens, vacant lots, roadsides, roads, sidewalks, sand pits, gravel pits, dumps, dunes, sandhills, orchards (e.g., apple), forest clearings, and open forests. Open ground; rather dry, sandy soil, bare or covered with low vegetation. Diurnal and nocturnal; active in the sunshine (running in the grass) or sheltering under plant rosettes (Verbascum), dead fallen plants, dead leaves, heaps of hay, logs, boards, and stones. Biology. Seasonality: April-December. Copulating pairs: May-June (MA, QU). Gravids: late May to early July (AB, MA, QU). Eggs are laid in the ground. Tenerals: late July to August. Adults found overwintering in the soil (mostly; as deep as 5 cm) and in decaying logs. Life span: 1-3 years. Adult food, in the field: lepidopterous caterpillars, chrysomelids, scarabaeids, and acridids. Larval food, in the field: lepidopterous caterpillars. Adults and larvae are cannibalistic in captivity. Economic importance: may play an important role in the control of cutworms and armyworms. Predators: toads, snakes, crows, woodpeckers, grackles, flycatchers, starlings, swallows, foxes, raccoons, and skunks. Parasites: tachinids (Sitophaga). Defense-mechanism: when alarmed, the adult may emit a fiery burning acid or a very disagreeable odor from the pygidial glands, or escape by diving into the water (may remain submerged for 2 minutes); when disturbed, the larva may bite hard. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in drift material, which indicates flight ability. Moderate runner. Poor climber (adult and larva). Good diver. Strongly favored by human activities. Collecting techniques. Pitfall trapping; light trapping; tying burlap sacks around lower tree trunks; sugaring tree trunks. References. Blatchley, 1910 (ecology, biology); Burgess and Collins, 1917 (biology); Moore, 1933 (dispersal power); Lindroth, 1961 and 1963a (natural history);

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Frank, 1971b (biology); Kirk, 1975c (ecology); Landry and Chamberland, 1976 (biology); Monteith, 1976 and 1977 (biology); Erwin, 1981 (natural history). Calosoma (Tapinosthenes) cancellatum Eschscholtz, 1833 Ecology. Lowlands and mountains. Cultivated fields (e.g., sugar beet). Open ground; dry soil. Diurnal; active in the sunshine on the ground or low shrubbery. Biology. Seasonality: April-September. Gravids: May-June (MA; in captivity). Adult food, in the field: lepidopterous caterpillars, elaterids, flies, and orthopterans. Larval food, in the field: lepidopterous caterpillars. Predators: crows. Dispersal power. Submacropterous, probably incapable of flight. Moderate runner. Occasional climber (on shrubs). The larva lives on or near the ground. Collecting technique. Pitfall trapping. References. Burgess and Collins, 1917 (biology); Lindroth, 1961 (natural history); Ball and Bousquet, 2001 (taxonomy). Calosoma (Microcallisthenes) dietzii Schaeffer, 1904 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Calosoma (Microcallisthenes) discors LeConte, 1857 Ecology. Uplands. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Calosoma (Carabosoma) eremicola Fall, 1910 Ecology. Fields and roads. Biology. Seasonality: March-June. Dispersal power. Macropterous, probably capable of flight. Calosoma (Callitropa) externum (Say, 1823) Ecology. Lowlands and mountains. Cultivated fields (e.g., oats, corn, alfalfa, cotton), pastures, vacant lots, and open forests. Open ground. Mostly nocturnal; usually sheltering during the day under stones (adult, larva) and logs (larva). Solitary or in pairs. Biology. Seasonality: April-November. Copulating pairs: early July (MA). Oviposition: early June to mid July (MA). Adults found overwintering in the soil (2.5-7.5 cm deep). Life span: 2 years. Adult food: lepidopterous caterpillars, grasshoppers, and scarabaeids (in the field); meat (in captivity). Larval food, in captivity: lepidopterous caterpillars. Predators: crows, grackles, meadowlarks, foxes, and raccoons. Parasites: mites. Defensemechanism: when alarmed, the adult may emit an acrid defensive fluid from the pygidial glands, at a range of up to 30 cm. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in drift material, which indicates flight ability. Moderate runner. Occasional climber (on trees). The larva is usually ground-dwelling. Collecting techniques. Pitfall trapping; light trapping. References. Blatchley, 1910 (ecolo-

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gy); Smith, 1910 (ecology, biology); Burgess and Collins, 1917 (biology); Löding, 1945 (dispersal power); Lindroth, 1961 (natural history); McCullough and Weinheimer, 1966 (biology); Kirk, 1969 (dispersal power). Calosoma (Calosoma) frigidum Kirby, 1837 Ecology. Lowlands and mountains. Deciduous, mixed or coniferous forests, tree plantations (e.g., Pinus), and orchards. Shaded ground; moderately dry soil covered with thick leaf litter. Diurnal and nocturnal; active on trees (e.g., Abies, Picea, Populus, Quercus) and shrubs, or sheltering under the loose bark of dead trees and in leaf litter. Gregarious (during caterpillar outbreaks). Biology. Seasonality: May-November. Copulating pairs: May-June (MA, NY, QU). Gravids: mid-May to late June (AB, MA). Each egg is laid singly into the soil. Some females lay eggs for two successive years. Tenerals: late May (QU) to mid-September (NS). Adults found overwintering after pupation in their own pupal cavity in the soil (about 5 cm deep on average) and in rotten tree-stumps. Life span: 2-5 years. Adult food: lepidopterous caterpillars, cicadas, and locusts (in the field); lepidopterous caterpillars (in captivity). Larval food, in captivity: lepidopterous pupae and caterpillars. Economic importance: plays an important role in the control of lepidopterous caterpillar (e.g., Heterocampa, Choristoneura) outbreaks. Predators: toads and grouses. Frequently crushed by cars. Parasites: tachinids. Defense-mechanism: when alarmed, the adult may emit a strong fetid smell. Dispersal power. Macropterous. Frequent flier (often swarming; during the day; to artificial lights at night). Frequent in drift material, which indicates flight ability. Moderate runner. Frequent climber (on trees, shrubs, and in malaise traps). The larva is ground-dwelling. Favored by the presence of large numbers of forest defoliators. Collecting techniques. Sweeping or beating trees and shrubs infested by caterpillar outbreaks; raking the leaf litter; light trapping; turning drift material; following the scent of the foraging adult. References. Blatchley, 1910 (dispersal power); Burgess, 1910 (biology); Burgess and Collins, 1917 (biology); Lindroth, 1961 (natural history); Larochelle, 1974a (ecology) and 1976f (biology); Landry, 1975a (biology); Loesch, 1977 (biology); Sanders and Frankenhuyzen, 1979 (biology); Reeves et al. 1983 (biology); Dearborne and Donahue, 1993 (ecology, biology); Snider and Snider, 1997 (biology). Calosoma (Blaptosoma) haydeni haydeni G.H. Horn, 1870 Ecology. Fields (mostly) and beds of dry brooks. Open ground; sheltering during the day under debris. Biology. Seasonality: April-November. Gravids: early June and midAugust (MA; in captivity). Tenerals: July (MA; in captivity). Adults found overwintering in the soil. Adult and larval food, in the field: lepidopterous caterpillars and pupae. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Burgess and Collins, 1917 (biology). Calosoma (Blaptosoma) haydeni punctulicolle Bates, 1891 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight.

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Calosoma (Microcallisthenes) lariversi Van Dyke, 1943 Ecology. Unknown. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Calosoma (Microcallisthenes) latipenne G.H. Horn, 1870 Ecology. Mountains. Biology. Seasonality: May-June. Adult food, in the field: grasshoppers. Dispersal power. Brachypterous, incapable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Calosoma (Chrysostigma) lepidum LeConte, 1845 Ecology. Cultivated fields and their borders (e.g., wheat); prairies. Open ground; rather dry soil covered with some vegetation (e.g., grass). Biology. Seasonality: mostly spring and early summer. Dispersal power. Submacropterous, probably incapable of flight. Moderate runner. Strongly favored by human activities. Collecting technique. Pitfall trapping. Reference. Lindroth, 1961 (ecology, dispersal power). Calosoma (Microcallisthenes) luxatum Say, 1823 Ecology. Uplands. Prairies, rangelands (short- and mid-grass), shrub-steppes, dry riverbeds, and pine forests. Open ground; very dry, sandy (mostly), clayish or rocky soil covered with some vegetation (e.g., Bromus, Poa). Biology. Seasonality: April-July. Tenerals: July-August (SA). Adult food: lepidopterous caterpillars, grasshoppers, and tenebrionids (in the field); lepidopterous caterpillars (in captivity). Predators, in captivity: grasshopper mice, deer mice, and ground squirrels. Defense-mechanism: when alarmed, the adult may emit a smelling defensive fluid from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. References. Lindroth, 1961 (natural history); McCullough, 1972a (biology); Lavigne, 1977 (ecology, biology); Ball and Bousquet, 2001 (taxonomy). Calosoma (Callitropa) macrum LeConte, 1853 Ecology. Unknown. Biology. Seasonality: June-August, October. Defense-mechanism: when alarmed, the adult may emit a defensive fluid from the pygidial glands. Dispersal power. Macropterous, probably capable of flight. Reference. McCullough, 1966b (biology). Calosoma (Camegonia) marginale Casey, 1897 Ecology. Uplands. Prairies, pastures, cultivated fields (e.g., cotton), and wastelands. Open ground; dry soil covered with some vegetation. Crepuscular; sheltering during the day. Biology. Seasonality: April-December. Oviposition: late July and August (MA; in captivity). Adults found overwintering in the soil (1.2-15 cm deep). Life span: 3 years. Adult food: lepidopterous caterpillars and scarabaeids (in the field); lepidopterous pu-

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pae and larvae, and mealworms (in captivity). Larval food, in captivity: lepidopterous pupae and larvae. Predators: toads. Parasites: nematodes. Defense-mechanism: when alarmed, the adult may emit a defensive fluid from the pygidial glands. Dispersal power. Macropterous. Frequent flier (sometimes swarming on warm nights to artificial lights). Moderate runner. Ground-dweller (adult and larva). Collecting techniques. Pitfall trapping; light trapping. References. Burgess and Collins, 1917 (biology; as lugubre LeConte); McCullough and Weinheimer, 1966 (biology). Calosoma (Microcallisthenes) moniliatum (LeConte, 1852) Ecology. Uplands. Dry forests. Biology. Seasonality: May-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. References. Lindroth, 1961 (natural history); Ball and Bousquet, 2001 (taxonomy). Calosoma (Microcallisthenes) monticola Casey, 1897 Ecology. Mountains. Sagebrush (Artemisia) areas. Open ground; dry soil. Biology. Seasonality: February-June, September. Adult food, in the field: cutworms, wireworms, scarabaeids, coccinellids, meloids, and staphylinids. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Ball and Bousquet, 2001 (taxonomy). Calosoma (Chrysostigma) morrisonii G.H. Horn, 1885 Ecology. Lowlands. Sand dunes. Open ground; dry soil. Biology. Seasonality: April. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Calosoma (Chrysostigma) obsoletum Say, 1823 Ecology. Uplands. Prairies, rangelands, cultivated fields (e.g., alfalfa, corn, wheat), fencerows, and roadsides. Open ground; soil covered with some vegetation (e.g., grass or weeds). Crepuscular; sheltering during the day under pieces of wood, dry cow dung, and dead fallen weeds; sometimes active by the hundreds on cloudy days. Biology. Seasonality: April-November. Oviposition: early June to early August (MA; in captivity). Adults found overwintering in cavities in the soil (as deep as 15 cm). Life span: 2-3 years. Adult and larval food, in the field: lepidopterous caterpillars and grasshoppers. Larval food, in captivity: lepidopterous caterpillars. Predators: kites. Dispersal power. Macropterous. Occasional flier. Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning debris. References. Burgess and Collins, 1917 (biology); Bell, 1971 (ecology, dispersal power); Kirk, 1975c (ecology). Calosoma (Microcallisthenes) oregonum (Gidaspow, 1959) Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Reference. Ball and Bousquet, 2001 (taxonomy).

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Calosoma (Camegonia) parvicolle Fall, 1910 Ecology. Uplands. Sand dunes. Open ground; dry soil covered with sparse vegetation. Diurnal and nocturnal. Gregarious. Biology. Seasonality: March-July. Adult food, in the field: lepidopterous caterpillars (mostly), orthopterans, and spiders. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Reference. McCullough, 1966b (biology). Calosoma (Camedula) peregrinator Guérin-Méneville, 1844 Ecology. Uplands. Grasslands, pastures, and golf courses. Open ground; soil covered with some vegetation. Diurnal and nocturnal. Biology. Seasonality: March-December. Copulating pairs: June (AZ). Oviposition: late June and early July (AZ). Adults found overwintering in cavities in the soil. Adult food, in the field: lepidopterous caterpillars. Adult and larval food, in captivity: lepidopterous caterpillars and pupae. Parasites: tachinids. Defense-mechanism: when alarmed, the adult may emit a defensive fluid from the pygidial glands. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Occasional climber (on trees). The larva is grounddwelling. Collecting technique. Pitfall trapping. References. Burgess and Collins, 1917 (biology); Ball and Bousquet, 2001 (taxonomy). Calosoma (Microcallisthenes) placerum (Gidaspow, 1959) Ecology. Uplands. Biology. Seasonality: April-June. Dispersal power. Brachypterous, incapable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Calosoma (Camegonia) prominens LeConte, 1853 Ecology. Unknown. Biology. Seasonality: August-September. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Calosoma (Callitropa) protractum LeConte, 1862 Ecology. Uplands. Biology. Defense-mechanism: when alarmed, the adult may discharge a spray from the pygidial glands. Dispersal power. Macropterous, probably capable of flight. Calosoma (Castrida) sayi Dejean, 1826 Ecology. Lowlands and mountains. Cultivated fields (e.g., corn, soybean, peanut, cotton, tobaccco, millet), grasslands, pastures, old fields, roadsides, low shrubbery, orchards (e.g., plum, peach, citrus), and open low forests (e.g., Quercus, Pinus). Open ground. Mostly active at sunset and sunrise; sheltering during the day under pieces of wood. Larvae are mostly diurnal. Biology. Seasonality: January-November. Adults found overwintering in cavities in the soil. Adult food, in the field: lepidopterous caterpillars and pupae (mostly), chrysomelids, and scarabaeid larvae. Larval food, in the field: insect

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larvae. Economic importance: plays an important role in the control of noctuid caterpillars in soybean fields. Predators: starlings. Defense-mechanism: when alarmed, the adult may emit a strong smoke-like odor from the pygidial glands. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in drift material, which indicates flight ability. Moderate runner. Occasional climber (on trees, shrubs, and plants). Strongly favored by human activities. Collecting techniques. Pitfall trapping; lifting pieces of wood; light trapping. References. Leng, 1915 (ecology, dispersal power); Burgess and Collins, 1917 (biology); Löding, 1945 (dispersal power); McCullough, 1966b (biology); Kirk, 1969 (biology, dispersal power) and 1970 (ecology); Price and Shepard, 1978a and 1978b (biology); Lesiewicz et al. 1983 (ecology, biology); Young, 1985a and 1985b (biology); Young and Hamm, 1985 and 1986 (biology). Calosoma (Microcallisthenes) schaefferi Breuning, 1928 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Calosoma (Calodrepa) scrutator (Fabricius, 1775) Ecology. Lowlands and mountains. Forests and their edges (mostly; deciduous), neighboring gardens, cultivated fields (e.g., cotton, oat), meadows, and wastelands; orchards (e.g., peach, plum, lime). Shaded ground. Mostly nocturnal; usually sheltering during the day under stones, sticks, and the loose bark of trees; active at night on trees and plants. Biology. Seasonality: March-November; rarer from August on. Oviposition: early June to mid-July (MA). Adults found overwintering in cavities in the soil (2.5-22.5 cm deep). Life span: 2 years. Adult food, in the field: lepidopterous caterpillars and pupae (mostly), cicadids, grasshoppers, scarabaeids; observed attacking a young bird and sucking its blood. Larval food, in captivity: lepidopterous caterpillars and mealworms. Economic importance: plays an important role in the control of the Gypsy Moth and the Forest Tent Caterpillar. Predators: frogs, bats, foxes, raccoons, cardinals, crows, cuckoos, grackles, swallows, and woodpeckers. Defense-mechanism: when alarmed, the adult may emit a burning defensive fluid or a pungent odor from the pygidial glands. Dispersal power. Macropterous. Frequent flier (sometimes swarming to artificial lights at night). Frequent in drift material, which indicates flight ability. Moderate runner. Frequent climber (on trees and plants). The larva is ground-dwelling. Collecting techniques. Light trapping; beating or sweeping the vegetation; turning drift material; locating the rustling sound of the foraging adult. References. Blatchley, 1910 (natural history); Burgess and Collins, 1917 (biology); Lindroth, 1961 (natural history); McCullough, 1966a (biology); McCullough and Weinheimer, 1966 (biology); Kirk, 1969 (dispersal power); Erwin, 1981 (natural history). Calosoma (Chrysostigma) semilaeve LeConte, 1852 Ecology. Lowlands and uplands. Cultivated fields (e.g., sugar beet, clover), roads, low shrubberies, and pea-vines. Diurnal and nocturnal. Biology. Seasonality: April-August.

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Copulating pairs: April (CA). Gravids: early May to early June (CA). Adults found overwintering in cavities in the soil (7.5-10 cm deep). Adult food, in the field: lepidopterous caterpillars (mostly), tenebrionids, elaterids, and carabids. Larval food, in captivity: lepidopterous caterpillars. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Occasional climber (adult and larva). Strongly favored by human activities. Collecting technique. Pitfall trapping. References. Burgess and Collins, 1917 (biology); Garner, 1954 (biology). Calosoma (Chrysostigma) simplex LeConte, 1878 Ecology. Unknown. Biology. Seasonality: May. Dispersal power. Macropterous, probably capable of flight. Calosoma (Calodrepa) splendidum Dejean, 1831 Ecology. Lowlands. Biology. Unknown. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Calosoma (Carabosoma) sponsum Casey, 1897 Ecology. Uplands. Biology. Seasonality: March, August. Dispersal power. Macropterous, probably capable of flight. Calosoma (Microcallisthenes) striatium Hatch, 1953 Ecology. Unknown. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Calosoma (Microcallisthenes) subaeneum Chaudoir, 1869 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Calosoma (Microcallisthenes) subasperatum Schaeffer, 1915 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Calosoma (Calosoma) sycophanta (Linné, 1758) Ecology. Lowlands and mountains. Deciduous forests (mostly) and orchards (e.g., apple). Shaded ground; moderately moist soil. Diurnal and nocturnal (adult and larva); the adult is active on trees and shrubs or on the ground. The female spends more time eating and staying on the ground than does the male. Gregarious (adult and larva). Biology. Seasonality: January-December; rarely after July. Copulating pairs: June-July.

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Gravids: from mid-May. Eggs are laid in the soil. Adults found overwintering in cavities in the soil (5-50 cm deep); some individuals remain dormant for almost 2 years. Life span: 2-4 years. Adult food, in the field: lepidopterous caterpillars and pupae (mostly; e.g., Lymantriidae, Thaumatopoeidae) and coccinellids. Larval food: lepidopterous caterpillars and pupae (in the field); beef liver and chicken meat (in captivity). Economic importance: plays an important role, as a specific predator, in the control of the Gypsy Moth. Predators: foxes, raccoons, skunks, crows, woodpeckers, and pentatomids. Parasites: tachinids, mites, and nematodes. Defense-mechanism: when alarmed, the adult may emit a pungent smell from the pygidial glands. Dispersal power. Macropterous. Frequent flier (to artificial lights at night, particularly in the spring). Moderate runner. Frequent climber (adult and larva; on trees and shrubs). Favored by Gypsy Moth outbreaks, human activities, and spring floods along large rivers. Collecting techniques. Light trapping; using tree-trunk barrier traps; beating or sweeping the vegetation. References. Burgess, 1911a and 1911b (natural history); Burgess and Collins, 1915 and 1917 (natural history); Lindroth, 1945 (natural history); Smith, 1959 (biology); Doane and Schaefer, 1971 (dispersal power); Sharova, 1981 (ecology; biology); Lindroth, 1985 (natural history); Weseloh, 1985 (biology, dispersal power), 1988, 1989a, 1989b, 1990 and 1993 (biology); Koch, 1989 (ecology); Luff, 1993 (biology). Calosoma (Chrysostigma) tepidum LeConte, 1852 Ecology. Uplands. Cultivated fields (e.g, pea), rangelands, and roadsides. Open ground; dry soil covered with sparse low vegetation. Biology. Seasonality: May-August. Tenerals: early May (BC). Adult food, in the field: lepidopterous caterpillars and grasshoppers. Predators: crows. Dispersal power. Submacropterous, probably incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Lindroth, 1961 (natural history). Calosoma (Calodrepa) wilcoxi LeConte, 1848 Ecology. Lowlands and mountains. Deciduous forests (mostly; e.g., Quercus, Acer), their edges, adjacent gardens and cultivated fields (e.g., pea, cotton); orchards (e.g., plum, peach) and trees in cities. Shaded ground. Nocturnal and diurnal; active in the sunshine on trees or sheltering during the day under stones and fallen trees. Biology. Seasonality: January-December. Copulating pairs: June (MA). Gravids: June (MA). Adults found overwintering in cavities in the soil (2.5-15 cm deep); able to hibernate through 2 winters. Life span: 2-3 years. Adult food, in the field: lepidopterous caterpillars and grasshoppers. Larval food, in captivity: lepidopterous caterpillars. Economic importance: plays an important role in the control of caterpillar (e.g., cankerworms) outbreaks. Predators: crows and woodpeckers. Parasites: tachinids (Sitophaga). Defense-mechanism: when alarmed, the adult may emit a pungent odor from the pygidial glands. Dispersal power. Macropterous. Frequent flier (sometimes swarming by the hundreds to artificial lights at night). Frequent in drift material, which indicates flight ability. Moderate runner. Frequent climber (on trees and plants). The larva is ground-dwelling. Collecting techniques. Light trapping; beating or sweeping the vegetation; turning drift material.

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References. Burgess and Collins, 1917 (biology); Lindroth, 1961 (natural history); Kirk, 1969 (dispersal power). Calosoma (Microcallisthenes) wilkesii (LeConte, 1852) Ecology. Fields; meadows (sometimes surrounded by light forests). Open ground; dry soil covered with low vegetation. Biology. Seasonality: May-September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. References. Lindroth, 1961 (natural history); Ball and Bousquet, 2001 (taxonomy). Calybe (Ega) laetula (LeConte, 1851) Ecology. Uplands. Margins of slow-running rivers and brooks; borders of pools (near streams). Open ground; wet, muddy soil covered with some sparse vegetation (e.g., grass). Diurnal; active in the sunshine. Biology. Seasonality: March. Dispersal power. Wing-condition unknown. Moderate runner. Collecting technique. Treading the soil with the feet. Comments. The adult resembles and moves like an ant. Calybe (Ega) sallei (Chevrolat, 1839) Ecology. Lowlands. Borders of lakes, ponds, sloughs, ditches, artesian wells, slow-running rivers and brooks. Open ground; wet soil consisting of mud or clay (sometimes mixed with sand), covered with some sparse vegetation. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under algal mats and small sticks. Gregarious. Associated with ants. Larval habitat: margin of a farm pond, under algal mats and small sticks. Biology. Seasonality: January-November. Larvae: April (TX). Tenerals: late July (TX). Adults found overwintering (AL). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Treading the soil with the feet. References. Leng, 1915 (ecology); Kirk, 1969 (ecology, dispersal power) and 1970 (dispersal power); Liebherr, 1983b (ecology, biology). Comments. The adult resembles and moves like an ant. Carabus (Autocarabus) auratus Linné, 1761 Ecology. Lowlands and mountains. Gardens, yards, and cultivated fields (e.g., hay, bean). Europe: warm, preferably loamy areas, such as cultivated fields (preferably), meadows, pastures, gardens, parks, roadsides. Mostly diurnal; active in the sunshine. Biology. Seasonality: April-July, November. Adults found overwintering in the soil (Europe). Life span: several years, in captivity. Adult food, in the field: earthworms and lepidopterous caterpillars (North America), insects, molluscs, earthworms, carrion, and vegetal matter (Europe). Defense-mechanism: when alarmed, the adult may emit a repugnant Brachinus-like smoke or burning defensive fluid from the pygidial glands, discharged towards the eyes of the aggressor. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Clearly effective as a colonist. Collecting technique. Pitfall trapping. References. Lindroth, 1945 and 1985 (natural history); Smith, 1959 (biology);

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° H urka, 1973 (biology); Nelson and Reynolds, 1987 (ecology, dispersal power); Koch, 1989 (ecology); Dearborne and Donahue, 1993 (biology). Comments. Introduced into North America as a potential predator of the Gypsy Moth.

Carabus (Autocarabus) cancellatus Illiger, 1798 Ecology. Open country (Europe). Biology. Seasonality: unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. References. Lindroth, 1945 and 1985 (natural history); Koch, 1989 (ecology). Comments. Doubtfully established in North America. Carabus (Tomocarabus) chamissonis Fischer von Waldheim, 1820 Ecology. Lowlands, mountains, and alpine zone. Moraines and alpine meadows. Coniferous (e.g., Pinus) and mixed forests (Populus, Picea). Vicinity of human habitations (urban and rural sites). Open or shaded ground; dry or moderately moist, stony, rocky or gravelly soil covered with discontinuous vegetation (e.g., Empetrum), or, rich soil covered with leaf litter. Nocturnal; sheltering during the day under stones or among dead leaves. Biology. Seasonality: May-September. Copulating pairs: early July (QU); coitus lasting 5-8 minutes. Tenerals: mid-August (AK), mid-September (NH). Adults found overwintering in their own pupal cavity. Adult food, in captivity: mealworms, shrimps, and a spider. Larval food, in captivity: mealworms. Eaten by insectivorous plants (Sarracenia). Defense-mechanism: when alarmed, the adult may emit a strong odor from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Favored by human activities. Collecting technique. Pitfall trapping. References. Lindroth, 1961 (natural history); Niemelä and Spence, 1991 (ecology); Niemelä et al. 1992 (ecology, biology); Ball and Bousquet, 2001 (taxonomy). Carabus (Tanaocarabus) finitimus Haldeman, 1852 Ecology. Forests. Nocturnal; sheltering during the day. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Carabus (Tanaocarabus) forreri Bates, 1882 Ecology. Mountains. Forests. Biology. Seasonality: June-July, September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Carabus (Carabus) goryi Dejean, 1831 (Carabus limbatus Say, 1823) Ecology. Lowlands and mountains. Deciduous (mostly; e.g., Quercus, Acer) and coniferous forests (e.g., Picea, Abies), orchards; cultivated fields (e.g., soybean) neighboring forests; cranberry bogs. Shaded ground; somewhat moist soil covered with thick leaf litter. Nocturnal; sheltering during the day among dead leaves and under stones (most-

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ly), under logs, boards, and in tree-stumps. Gregarious (in habitats providing few opportunities for shelter). Biology. Seasonality: February-November. Copulating pairs: April-May (NC). Gravids: mid-April (IL). Tenerals: April (rarely; MD), June-July, September. Adults found overwintering in woods; under stones and in rotten logs. Adult food: immature stages of the Gypsy Moth and soft-bodied insect larvae (in the field); mealworms, a Carabus, a dead salamander, and meat (in captivity). Larval food, in captivity: mealworms. Predators: owls. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on tree trunks; up to 20 m above ground). Foraging wider distances on warmer nights. Collecting techniques. Pitfall trapping; raking the leaf litter; baiting traps with molasses or beer. References. Blatchley, 1910 (ecology; as limbatus Say); Gilbert, 1957 (biology; as limbatus Say); Barr, 1969 (ecology; as limbatus Say); Liebherr and Mahar, 1979 (biology; as limbatus Say); Erwin, 1981 (natural history); Lenski, 1982 and 1984 (biology; as limbatus Say); Anderson et al. 1995 (ecology). Carabus (Carabus) granulatus granulatus Linné, 1758 Ecology. Lowlands. Low deciduous forests, swamps; borders of marshes, pools, and roadside ditches; cultivated fields, gardens, and orchards (e.g., apple). Greenhouses (occasionally). Preferably close to water. Slightly shaded or open ground; moderately dry clay soil. Mostly nocturnal; usually sheltering during the day under the loose bark of tree-stumps. Gregarious (in winter). Biology. Seasonality: May-July, September-November. Eggs are laid in cavities in the soil. Adults found overwintering in woods; under the loose bark of standing tree-stumps (e.g., Ulmus, Acer), in rotten tree-stumps, and in fallen timber. Adult food: snails (mostly), insects, and earthworms (Europe) (in the field); meat and a banana skin (in captivity). Larval food, in the field (Europe): mostly snails. Defense-mechanism: when alarmed, the adult may emit a strong aromatic vapor. Dispersal power. Brachypterous, incapable of flight (North America). Wing-dimorphic: brachypterous (mostly) or macropterous (rarely; capable of flight). Flight records (Europe). Moderate runner. Frequent climber (on trees, treestumps, low herbage). Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Lifting the loose bark of trees; pitfall trapping; beating or sweeping the vegetation. References. Lindroth, 1945 and 1961 (natural histo° ry); Sharova, 1981 (ecology); Hurka, 1973 (biology); Lindroth, 1985 (natural history); Luff, 1993 (biology). Carabus (Carabus) granulatus hibernicus Lindroth, 1956 Ecology. Lowlands. Field margins, cultivated fields (e.g., hay, potato); borders of marshes and temporary pools; mud flats, lake shores, and orchards. Open ground; moist or wet, clay soil covered with some vegetation. Nocturnal; sheltering during the day under stones, pieces of wood or cardboard, and dead fallen plants. Biology. Seasonality: AprilSeptember, November. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Clerly effective as a colonist. Strongly favored by human activities. Collecting technique. Pitfall trapping. Reference. Lindroth, 1961 (natural history).

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Carabus (Homoeocarabus) maeander Fischer von Waldheim, 1820 Ecology. Lowlands and mountains. Edges of eutrophic marshes, ponds, Carex-bogs, fens, sloughs, and pools; swamps; edges of small lakes, slow rivers, brooks, and irrigation canals; wet meadows, low open forests, and cultivated fields (e.g., carrot). Open ground; somewhat moist, clayish, muddy or peaty soil covered with moderate or dense vegetation (e.g., Carex, Typha, Sphagnum, grass). Diurnal and nocturnal; active in the sunshine among the vegetation (especially in spring) or sheltering on cloudy or cool days under dead plants, in and under moss, under stones, logs, and pieces of cardboard. Solitary. Biology. March-November. Gravids: late-May (QU). Tenerals: mid-July (MI) to early August (QU). Adults found overwintering from October on higher and drier ground, in the upper zone of brook borders, roadsides, and gravel pits, in vacant lots, on hills, and at the edge of woods; under well-embedded stones (mostly), in the soil at the base of trees, under the loose bark of fallen trees, and under logs. Adult food, in captivity: mealworms, shrimps, lepidopterous caterpillars, earthworms, spiders, and meat. Larval food, in captivity: mealworms. Predators: frogs, toads, bluebirds, crows, and spiders. Parasites: tachinids. Sometimes crushed by cars on the roads in spring. Defense-mechanism: when alarmed, the adult may emit a strong fetid odor from the pygidial glands or escape by swimming. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous (possibly capable of flight). Moderate runner. Occasional climber (on plants). Strong burrower. Good swimmer. Moderately favored by human activities. Collecting techniques. Treading the edges of Carex-marshes into the water; turning large stones; tearing the sod away from large tree roots from a sparsely wooded pasture (Blatchley); baiting traps with fish; sweeping plants. References. Blatchley, 1910 (ecology); Lindroth, 1955a, 1961, and 1963a (natural history); Larochelle, 1973k and 1976g (dispersal power). Carabus (Archicarabus) nemoralis O.F. Müller, 1764 Ecology. Lowlands and mountains. Parks, gardens, lawns, pastures, cultivated fields (e.g., potato, corn, hay, bean) and theirs borders; vacant lots, backyards, docks, ski fields, gravel pits, sand pits, roadside ditches, dumps, grenhouses, basements, orchards, thickets, hedges, decidous and mixed forests. Open or shaded ground; moderately moist, often clay soil covered with some vegetation. Mostly nocturnal; usually sheltering during the day under stones, boards, stacks of lumber, dead leaves, heaps of hay, and other debris; sometimes active on cloudy days on the ground. Biology. Seasonality: March-December. Copulating pairs: April-June (QU); coitus lasting 10-54 minutes. Gravids: May (QU). Eggs are laid singly in cavities in the soil. Tenerals: mid-July (NS) to September (VT). Adults found overwintering on higher and drier ground, in hedges, at the edge of woods, and in woods; in cavities in the soil, under stones, in leaf litter, and under the loose bark of fallen treestumps; able to hibernate through 2 winters. Adult food: slugs, earthworms, lepidopterous caterpillars, and a small snake (in the field); slugs (mostly), a crushed snail, earthworms, lepidopterous caterpillars, carabids, chrysomelid eggs, mealworms, shrimps, and meat (in captivity). Larval food: lepidopterous caterpillars (in the field); mealworms and meat (in captivity). Economic importance: plays an important role in the control of slugs.

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Predators: frogs, toads, shrews (in captivity), starlings, and spiders. Parasites: tachinids and mites. Frequently crushed by cars on roads in the spring and by pedestrians on sidewalks. Defense-mechanism: when alarmed, the adult may emit a pungent smell from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Sometimes swarming on the ground under artificial lights at night. Moderate runner. Clearly effective as a colonist. Strongly favored by human activities (e.g., cultivation and transportation of agricultural and horticultural produce). Collecting techniques. Pitfall trapping; turning stones and pieces of wood; collecting along city sidewalks at night. References. Delkeskamp, 1930 (biology); Lindroth, 1945, 1955a, 1961, and 1963a (natural history); Glendenning, 1952 (biology); Smith, 1959 (biology); Poulin and O’Neill, 1969 (biology); Barlow, ° 1973 (biology); Hurka, 1973 (biology); Ostaff and Freitag, 1973 (biology); Randolph et al. 1976 (biology); Sharova, 1981 (ecology); Lindroth, 1985 (natural history); Wallin and Ekblom, 1988 (dispersal power); Koch, 1989 (ecology); Kavanaugh, 1992 (natural history); Luff, 1993 (biology); Dearborne and Donahue, 1993 (biology); Digweed, 1994 (biology); Johnson and Reeves, 1995 (biology). Carabus (Hemicarabus) serratus Say, 1823 Ecology. Lowlands and mountains. Moraines, gravel pits, sand pits, meadows, pastures, cultivated fields and their borders (e.g., wheat, fescue, barley), vacant lots, roadsides, foot of rock-falls, railway embankments, hills, forest clearings, and open forests. Open ground; rather dry, gravelly (mostly) or sandy soil covered with sparse vegetation. Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering during the day under stones (mostly), logs, pieces of cardboard, heaps of hay, and dead leaves. Biology. Seasonality: April-November; rarely in July. Copulating pairs: mid-June (NF, QU). Gravids: late May (QU). Tenerals: mid-July to September. Adults found overwintering on higher and drier ground, in the upper zone of sand pits and gravel pits; under plant rosettes (Verbascum). Adult food, in captivity: mealworms, shrimps, earthworms, lepidopterous caterpillars, and apple. Predators: toads, starlings, and skunks. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous (possibly capable of flight). Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; baiting traps with molasses; turning large stones from sand pits and gravel pits; light trapping. References. Weeks, 1888 (biology); Darlington, 1936 (dispersal power); Lindroth, 1955a, 1961, and 1963a (natural history); Kirk, 1975c (ecology); Erwin, 1981 (natural history). Carabus (Tanaocarabus) sylvosus Say, 1823 Ecology. Lowlands and mountains. Deciduous (mostly; e.g., Quercus), mixed, and coniferous forests (e.g., Pinus), orchards (e.g., peach), and adjacent cultivated fields (e.g., soybean, potato). Shaded ground; more or less moist soil. Nocturnal; sheltering during the day under logs, stones, and boards. Biology. Seasonality: January-December. Gravids: autumn (NC). Tenerals: late spring and early summer (NC), early July (NJ, WI). Adults found overwintering in woods; under logs and in the soil. Adult food: lepidopterous pupae and scarabaeids (in the field); mealworms, soft-bodied insect larvae, and dead

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salamanders (in captivity). Larval food, in captivity: mealworms. Predators: crows, starlings, and raccoons. Defense-mechanism: when alarmed, the adult may emit a Brachinus-like smoke from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning logs and stones; baiting traps with molasses. References. Lindroth, 1961 (natural history); Barr, 1969 (ecology); Kirk, 1970 (biology); Erwin, 1981 (natural history); Lenski, 1982 and 1984 (biology); Anderson et al. 1995 (ecology). Carabus (Tomocarabus) taedatus agassii LeConte, 1850 Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, meadows, shrubsteppes, canyons, mountain slopes and cirques, cultivated fields (e.g., wheat, barley), sagebrush-steppes, rangelands, open deciduous (e.g., Populus) or coniferous forests (e.g., Abies, Picea), and burnt forests. Open ground; rather dry, mostly gravelly soil covered with thin low vegetation or dead leaves. Nocturnal; sheltering during the day under stones and logs. Biology. Seasonality: May-October. Adults found overwintering in woods; in rotten tree-stumps. Adult food, in the field: lepidopterous caterpillars. Adult and larval food, in captivity: lepidopterous eggs and caterpillars, mealworms, shrimps, and meat. Economic importance: may play an important role in the control of the Forest Tent Caterpillar. Predators: frogs and coyotes. Defense-mechanism: when alarmed, the adult may emit a defensive fluid from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Quick to colonize recently burnt woods. Collecting techniques. Pitfall trapping; turning stones and logs; baiting traps with molasses; smoking tree-stumps; light trapping. References. Lindroth, 1961 and 1963a (natural history; as taedatus Fabricius); Benn et al. 1973 (biology; as taedatus Fabricius); Parmenter and MacMahon, 1988 (biology); Holliday, 1991 (biology); Ball and Bousquet, 2001 (taxonomy). Carabus (Tomocarabus) taedatus taedatus Fabricius, 1787 Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, meadows, edges of snowfields, and vicinity of forests. Open ground. Mostly nocturnal; usually sheltering during the day under stones, logs, and dead leaves; sometimes active during the day on snow patches. Biology. Seasonality: June-July. Adult food, in captivity: scarabaeid larvae. Predators: spiders. Parasites: tachinids. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones and logs. Reference. Ball and Bousquet, 2001 (taxonomy). Carabus (Aulonocarabus) truncaticollis Eschscholtz, 1833 Ecology. Uplands. Meadow slopes (in tundra). Open ground; moderately moist, stony or muddy soil covered with meadow-type vegetation, dwarf birches or willows. Larval habitat: under moss carpets. Biology. Seasonality: June-August (AK). Larvae: July-August (AK). Adult food, in the field: a dipterous larva. Predators: jaegers. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. References. Lindroth, 1961 (natural history); Ball and Bousquet, 2001 (taxonomy).

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Carabus (Megodontus) vietinghoffii vietinghoffii Adams, 1812 Ecology. Lowlands. Forests (e.g., Populus, Betula, Picea) and barren country. Shaded or open ground. Nocturnal; sheltering during the day under logs, stones, and other debris. Biology. Seasonality: May-October. Tenerals: early August (NT). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones and logs. References. Hicks, 1953 (ecology); Lindroth, 1961 (natural history). Carabus (Carabus) vinctus (Weber, 1801) Ecology. Lowlands and mountains. Flood-plain forests (e.g., Liriodendron, Liquidambar), low deciduous forests, and swamp-forests. Close to water. Shaded or half-shaded ground; wet, clayish or muddy soil covered with dead leaves. Nocturnal; sheltering during the day under logs, fallen branches, the loose bark of fallen trees, stones, and boards. Biology. Seasonality: January-December. Tenerals: late July (VA). Adults found overwintering in swamp-forests; in rotten tree-stumps and under the loose bark of fallen trees. Able to survive winter inundations. Adult food: dead fish, flesh, and fowl (in the field); lepidopterous caterpillars (in captivity). Predators: frogs, bluebirds, crows, ducks, starlings, and skunks. Defense-mechanism: when alarmed, the adult may emit an acid secretion from the pygidial glands at a distance of 20 cm or more, striking the pursuer’s face and causing an intense burning. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; baiting traps with molasses or bananas; light trapping. References. Blatchley, 1910 (ecology); Frost, 1910a and 1929a (biology); Lindroth, 1961 (natural history); Kirk, 1969 (ecology); Erwin, 1981 (natural history); Lesiewicz et al. 1983 (ecology, biology); Anderson et al. 1995 (ecology). Chlaenius (Chlaenius) aestivus Say, 1823 Ecology. Lowlands and mountains. Flood-plain forests along large rivers, small brooks, and lakes. Moist habitats, well away from open water (Lake Erie islands). Shaded ground; moist or wet, clayish, muddy or sand-mixed clay soil. Nocturnal; sheltering during the day under stones and logs (mostly), dead leaves, and fallen tree bark. Gregarious (in winter). Biology. Seasonality: March-December. Eggs are laid in mud or clay cells stuck on dead twigs, leaves, plant stems, and on the bark of trees and shrubs (sometimes up to 3 m above ground). Tenerals: late July (NJ), mid-August (MA). Adults found overwintering on higher and drier ground, in woods and vacant lots; in rotten standing treestumps, under the loose bark of fallen trees, under fallen trees and dead leaves. Adult food, in captivity: meat. Predators: crows. Parasites: mites, Laboulbeniales (Fungi), and hymenopterans. Dispersal power. Wing-polymorphic: brachypterous (incapable of flight) to macropterous (capable of flight). Occasional flier (to artificial lights at night). Moderate runner. Occasional climber (on trees and shrubs). Collecting techniques. Pitfall trapping; turning stones and logs; night collecting with a headlamp. References. King, 1919 (biology); Darlington, 1936 (dispersal power); Bell, 1960 (natural history); Lindroth, 1969a (biology, dispersal power); Erwin, 1981 (natural history); Anderson et al. 1995 (ecology); Will et al. 1995 (ecology, dispersal power).

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Chlaenius (Agostenus) alternatus G.H. Horn, 1871 Ecology. Lowlands and mountains. Edges of ponds, lakes, pools, marshes, slow rivers and drainage channels; swamps and cultivated fields (e.g., wheat, barley). Open ground; wet, clay soil covered with dense vegetation (e.g., Carex, Juncus). Nocturnal; sheltering during the day under dead leaves and stones. Biology. Seasonality: May-July, September. Oviposition: late May to late June (AB). Adult food, in captivity: mealworms and shrimps. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Treading the vegetation into the water. Reference. Lindroth, 1969a (natural history). Chlaenius (Anomoglossus) amoenus Dejean, 1831 Ecology. Lowlands. Deciduous forests. Moderately moist soil. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: March-August, October. Tenerals: early June (MS). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. Reference. Bell, 1960 (ecology). Chlaenius (Chlaenius) augustus Newman, 1838 Ecology. Lowlands. Overflow grasslands (AL). In inundation debris (NJ). Biology. Seasonality: February-August, December. Dispersal power. Brachypterous, incapable of flight. Reference. Löding, 1945 (ecology). Chlaenius (Chlaenius) azurescens Chaudoir, 1876 Natural History. Unknown. Chlaenius (Chlaeniellus) brevilabris brevilabris LeConte, 1848 Ecology. Lowlands. Flood-plain forests along rivers (mostly), brooks, lakes, ponds, and marshes. Open or slightly shaded ground; moist or wet, clayish, muddy, gravelly or sandy soil, bare or covered with sparse vegetation. Nocturnal; sheltering during the day under stones (mostly) and pieces of wood. Biology. Seasonality: March-November. Tenerals: mid-July (WV), August (MN). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; turning stones. References. Bell, 1960 (ecology); Lindroth, 1969a (dispersal power); Lee, 1978 (ecology). Chlaenius (Chlaeniellus) brevilabris texanellus Casey, 1914 Ecology. Lowlands and mountains. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight.

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Chlaenius (Agostenus) caeruleicollis Chaudoir, 1876 Ecology. Grasslands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Chlaenius (Brachylobus) caurinus (G.H. Horn, 1885) Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Chlaenius (Chlaenius) chaudoiri G.H. Horn, 1876 Ecology. A small pond with bare sandy-clayish bank (AZ). Biology. Seasonality: September. Dispersal power. Macropterous, probably capable of flight. Chlaenius (Chlaeniellus) circumcinctus Say, 1830 (Chlaenius perplexus Dejean, 1831) Ecology. Lowlands. Borders of lakes, ponds, and swamps. Shaded ground; wet, muddy or clayish soil. Nocturnal; sheltering during the day under dead grass, pieces of wood, and other debris. Biology. Seasonality: January-August. Tenerals: March-April (FL). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Treading the vegetation into the water. Reference. Leng, 1915 (ecology). Chlaenius (Chlaenius) cordicollis Kirby, 1837 Ecology. Lowlands and mountains. Shores of large lakes and rivers. Close to water and exposed to the wave action. Open ground; wet, gravelly-stony soil, clay- or silt-mixed, bare or covered with sparse vegetation (e.g., grass). Nocturnal; sheltering during the day under large stones and limestone slabs. Gregarious. Adults and larvae are active at night among the stones. Larval habitat: as the adult; under stones. Biology. Seasonality: AprilOctober. Gravids: May to late June (QU, VT). Tenerals: July-September. Adult food, in the field: dead or injured insects. Defense-mechanism: when alarmed, the adult may escape by diving into the water and staying submerged for nearly a minute, or emit a strong persistent odorous spray from the pygidial glands; discharge is four times as long as body length and used with accuracy and efficiency against ants. Dispersal power. Macropterous. Frequent flier (to artificial lights in captivity). Moderate runner. Good swimmer. Collecting technique. Turning large stones and limestone slabs. References. Schaupp, 1880b (biology; as leucoscelis Chevrolat); Bell, 1960 (natural history); Lindroth, 1969a (ecology); Erwin, 1981 (natural history); Will et al. 1995 (ecology, dispersal power). Chlaenius (Chlaenius) cumatilis LeConte, 1851 Ecology. Lowlands and mountains. Stream banks: along canyons. Shaded ground; moist, muddy soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality:

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January, March, May-July, September, December. Parasites: Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult may emit a fine spray from the pygidial glands. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones. Chlaenius (Chlaenius) cursor Chevrolat, 1835 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Chlaenius (Anomoglossus) emarginatus Say, 1823 Ecology. Lowlands and mountains. Deciduous forests: along paths, roadsides, small openings, clearings, and edges. Cultivated fields (rarely; e.g., corn). Shaded ground; moderately moist, gravelly or sandy soil. Nocturnal; sheltering during the day under well-embedded stones, logs, pieces of wood, and dead leaves. Gregarious (in winter). Biology. Seasonality: March-November. Copulating pairs: May to early July (QU). Gravids: May (QU). Tenerals: late July to October; mostly August. Adults found overwintering on higher and drier ground, in woods, at the edge of woods, in vacant lots, and in the upper zone of gravel pits; under large embedded stones (mostly), under pieces of wood or cardboard, and in moss. Adult food: immature stages of the Gypsy Moth and other small soft-bodied organisms (in the field); meat, a raisin, and pieces of banana, apple, and orange (in captivity). Predators: toads and starlings. Defense-mechanism: when alarmed, the adult may emit a strong musky smell from the pygidial glands. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strong burrower. Collecting techniques. Pitfall trapping; turning large embedded stones at the edge of woods; light trapping. References. Bell, 1960 (ecology); Lindroth, 1969a (dispersal power); Kirk, 1975c (dispersal power); Erwin, 1981 (natural history); Anderson et al. 1995 (ecology). Chlaenius (Chlaenius) erythropus Germar, 1824 Ecology. Lowlands and mountains. Mature flood-plain forests: along rivers. Edges of swamps, lakes, and ponds. Cultivated fields (rarely; e.g., corn). Shaded ground; wet or moist, clayish, sandy or muddy soil. Nocturnal; sheltering during the day under logs, boards, and other debris. Gregarious (in winter). Biology. Seasonality: January-December. Tenerals: November (TX). Adults found overwintering in colonies with Galerita and Brachinus in and under decayed pine logs. Adult food, in the field: lepidopterous caterpillars (mostly) and earthworms. Predators: alligators. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pitfall trapping; baiting traps with syrup; turning logs. References. Blatchley, 1910 (biology); Leng, 1915 (ecology); Bell, 1960 (natural history); Kirk, 1969 (dispersal power); Lindroth, 1969a (natural history).

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Chlaenius (Chlaeniellus) flaccidus G.H. Horn, 1876 Ecology. Lowlands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Chlaenius (Chlaeniellus) floridanus G.H. Horn, 1876 Ecology. Lowlands. Lake shores. Wet, sandy soil. Nocturnal; sheltering during the day. Biology. Seasonality: January-April, June-September, November-December. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Pitfall trapping. Chlaenius (Chlaenius) fuscicornis Dejean, 1831 Ecology. Lowlands. Swamp-forests. Shaded ground; moist soil. Nocturnal; sheltering during the day under logs and driftwood. Biology. Seasonality: January-July, September-December. Adults found overwintering in rotten logs. Adult food, in the field: small insects. Defense-mechanism: when alarmed, the adult may emit a repugnant smoky spray from the pygidial glands. Dispersal power. Macropterous. Occasional flier. Moderate runner. Collecting techniques. Pitfall trapping; turning logs. Chlaenius (Chlaeniellus) glaucus LeConte, 1856 Ecology. A small pond with bare, sandy-clayish bank (AZ). Biology. Seasonality: September. Dispersal power. Macropterous, probably capable of flight. Chlaenius (Agostenus) harpalinus Eschscholtz, 1833 Ecology. Lowlands and mountains. Edges of ponds and small lakes. Ground shaded by trees (e.g., Salix); moist soil. Nocturnal; sheltering during the day under dead dead leaves and grass. Biology. Seasonality: June, September. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. Reference. Lindroth, 1969a (ecology, dispersal power). Chlaenius (Chlaeniellus) impunctifrons Say, 1823 Ecology. Lowlands and mountains. Border of marshes (e.g., Typha) and swamps; swamp-forests, marshy lake shores and river banks; flood-plain forests; margins of slow streams, ponds, and pools. Open or shaded ground; soft, wet soil consisting of clay, mud or humus-rich clay, covered with some vegetation. Nocturnal; sheltering during the day under logs, fallen branches and pieces of bark, dead leaves, and stones. Biology. Seasonality: January-December. Copulating pairs: May and early June (NY, QU). Gravids: May to late July, mostly before mid-June (NY, QU). Eggs are laid in mud or clay cells stuck to the underside of leaves of Typha-plants, shrubs or trees (up to 2 m above ground). Tenerals: early July to mid-September. Adults found overwin-

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tering on higher and drier ground, about 7-8 m from the summer habitat, in the upper zone of sand pits and gravel pits, at the edge of woods, and in woods; in the soil (2.515 cm deep), under well-embedded stones, dead leaves, and the loose bark of fallen trees. Adult food: insects and other small arthropods (in the field); lepidopterous caterpillars, mealworms, dead carabids and slugs, meat and corn kernels (in captivity). Larval food, in captivity: mealworms. Predators: frogs, toads, and crows. Parasites: hymenopterans. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Regular climber (on plants, shrubs, and trees). Strong burrower. Collecting techniques. Pitfall trapping; turning debris; light trapping; sweeping or beating the vegetation. References. Riley, 1884 (biology); Claassen, 1919 (ecology, biology); King, 1919 (biology); Bell, 1960 (natural history); Erwin, 1981 (natural history). Chlaenius (Agostenus) interruptus G.H. Horn, 1876 Ecology. Lowlands. Border of a pond, on rather firm soil covered with some vegetation (Carex; AZ). Biology. Seasonality: April, June. Dispersal power. Macropterous, probably capable of flight. Reference. Lindroth, 1969a (natural history). Chlaenius (Chlaenius) laticollis Say, 1823 Ecology. Lowlands and mountains. Swamps, swamp-forests; borders of lakes, ponds, and rivers; cultivated fields (e.g., corn) and pastures. Shaded ground; moist or wet, muddy, clayish or sandy soil. Nocturnal; sheltering during the day under logs (mostly), driftwood, fallen pieces of bark, and stones. Gregarious (in winter). Biology. Seasonality: January-December. Copulating pairs: early June (NY). Eggs are laid on the ground. Tenerals: August (KY). Adults found overwintering in colonies with Galerita, Chlaenius, and Dicaelus in woods and on hills; under logs, stones, and the loose bark of trees. Larval food, in captivity: dipterous larvae. Predators: crows. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; turning logs; light trapping. References. Schaupp, 1880a (biology); Dimmock and Knab, 1904 (biology); Leng, 1915 (ecology); Bell, 1960 (natural history); Erwin, 1981 (natural history). Chlaenius (Chlaenius) leucoscelis monachus LeConte, 1851 Ecology. Mountains. Brook margins. Moist soil. Biology. Seasonality: September. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Chlaenius (Chlaenius) leucoscelis sanantonialis Casey, 1914 Ecology. Mountains. River banks. Open ground; moist, clayish or muddy, bare soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: July-August. Dispersal power. Macropterous, probably capable of flight. Moderate runner.

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Chlaenius (Chlaenius) leucoscelis sonomae Casey, 1920 Ecology. Unknown. Biology. Seasonality: April, October. Tenerals: September (CA). Dispersal power. Macropterous, probably capable of flight. Chlaenius (Brachylobus) lithophilus indigaceus (Casey, 1914) Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Chlaenius (Brachylobus) lithophilus lithophilus Say, 1823 Ecology. Lowlands and mountains. Borders of lakes, ponds, pools (e.g., in sand pits and gravel pits), marshes, rivers, and brooks; swamps. Often associated with quite small bodies of water. Open or shaded ground; rather firm (mostly) or soft, moist or wet, clayish, muddy, gravelly or sandy soil covered with dense vegetation (e.g., grass). Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering during the day under stones (mostly), logs, and dead leaves. Biology. Seasonality: March-December. Gravids: May to mid-June (IL, NY, QU). Tenerals: mid-July to mid-August. Adults found overwintering on higher and drier ground, about 7-8 m from the summer habitat, at the edge of woods, in woods, in the upper zone of sand pits and gravel pits; in the soil (2.515 cm deep), under stones, and under plant rosettes (Verbascum). Adult food, in captivity: earthworms, dead slugs, meat, and corn kernels. Predators: frogs, toads, and robins. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; to artifical lights in captivity). Moderate runner. Collecting techniques. Turning stones; light trapping. References. Blatchley, 1910 (ecology); Bell, 1960 (natural history); Lindroth, 1963a and 1969a (natural history). Chlaenius (Pseudanomoglossus) maxillosus G.H. Horn, 1876 Ecology. Coastal lowlands. Salt marshes, brackish marshes, and sea beaches. Open ground; wet, saline soil covered with some vegetation. Nocturnal; sheltering during the day under dead algae. Biology. Seasonality: January-October. Adults found overwintering in brackish marshes. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Pitfall trapping. Reference. Löding, 1945 (ecology). Chlaenius (Chlaeniellus) nebraskensis LeConte, 1856 Ecology. Uplands. Borders of small lakes, reservoirs, and cattle water holes. Open ground; rather firm, moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: AprilSeptember. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Pitfall trapping. References. Lindroth, 1969a (natural history); Bell, 1971 (ecology); Kirk, 1975c (dispersal power).

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Chlaenius (Chlaeniellus) nemoralis Say, 1823 Ecology. Lowlands and mountains. Marshy river banks; swamp and pond edges. Cultivated fields (e.g., corn, oat, alfalfa), clear-cut fields, hills, fencerows, city lots, gardens, and yards. Open or shaded places; wet or moist soil covered with some vegetation (e.g., grass). Often close to small bodies of water. Nocturnal; sheltering during the day under stones, pieces of wood, bark, and other debris. Gregarious (in winter). Biology. Seasonality: January-December. Adults found overwintering at the edge of woods and in woods; under logs and under the loose bark of fallen trees. Adult food, in the field: insects, earthworms, fungi, and plant fragments. Predators: frogs and crows. Defense-mechanism: when alarmed, the adult may emit a Brachinus-like smoke from the pygidial glands. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pitfall trapping; turning stones and pieces of wood; light trapping. References. Leng, 1915 (ecology); Bell, 1960 (ecology); Kirk, 1969 (dispersal power) and 1975c (ecology, dispersal power); Lindroth, 1969a (natural history); Lee, 1978 (ecology, dispersal power); Erwin, 1981 (natural history). Chlaenius (Agostenus) niger Randall, 1838 Ecology. Lowlands and mountains. Swamps; borders of eutrophic marshes; marshy borders of ponds, irrigation channels, and roadside ditches; edges of saline marshes; meadows. Open ground; soil, when flooded, covered with rich but discontinuous vegetation (e.g., Typha), or, when not flooded, soft, wet muddy, sometimes saline, rich in organic debris and covered with similar vegetation. Nocturnal; sheltering during the day among mats of dead, often floating plants (mostly), in clumps of broomsedges, under dead leaves or stones. Adults are often active under water (depths of 15-20 cm). Biology. Seasonality: January-September. Tenerals: early August (QU, MN). Adults found overwintering on higher and drier ground, in vacant lots; under dead leaves and in the soil. Adult and larval food, in captivity: mealworms. Predators: crows. Defense-mechanism: when alarmed, the adult may dive into the water and remain submerged for a few minutes. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Excellent swimmer and diver. Strongly favored by human activities (e.g., creation of drainage channels and roadside ditches). Collecting techniques. Treading mats of dead vegetation into the water; light trapping. References. Ulke, 1903 (dispersal power); Bell, 1960 (natural history); Lindroth, 1963a and 1969a (natural history). Chlaenius (Chlaeniellus) obsoletus LeConte, 1851 Ecology. Lowlands and mountains. Damp places (CA). Biology. Seasonality: September-October, December. Dispersal power. Macropterous, probably capable of flight. Moderate runner.

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Chlaenius (Chlaenius) orbus G.H. Horn, 1871 Ecology. Lowlands. Damp places (TX). Nocturnal; sheltering during the day under logs. Biology. Seasonality: September. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Chlaenius (Chlaeniellus) oxygonus Chaudoir, 1843 Ecology. Lowlands. Edges of deciduous forests (MS). A cypress-swamp (FL). Moist soil. Nocturnal; sheltering during the day under dead leaves and bark. Biology. Seasonality: April-May, July-August, December. Tenerals: May (FL). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. Chlaenius (Chlaenius) patruelis LeConte, 1844 Ecology. Lowlands. Edges of a small lake, under pieces of wood (FL). Biology. Seasonality: January-March, July, November. Dispersal power. Macropterous, probably capable of flight. Chlaenius (Chlaeniellus) pennsylvanicus blanditus Casey, 1920 Ecology. Lake shores. Open ground; wet, muddy, bare soil. Nocturnal; sheltering during the day under vegetal debris and pieces of wood. Biology. Seasonality: July. Tenerals: late July (UT). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Chlaenius (Chlaeniellus) pennsylvanicus pennsylvanicus Say, 1823 Ecology. Lowlands, mountains, subalpine and alpine zones. Borders of ponds, pools (e.g., in sand pits, gravel pits), lakes, eutrophic marshes, slow rivers and brooks; floodplain forests and swamps. Often associated with small bodies of standing water. Open or slightly shaded ground; wet, muddy or clayish soil, rich in organic debris, with bare spots surrounded by dense vegetation (e.g., Carex, grass, Typha). Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering during the day under dead fallen plants and leaves, pieces of wood, stones, and other debris. Gregarious. Biology. Seasonality: January-December. Copulating pairs: early June (QU); coitus lasting 2-49 minutes. Gravids: May to mid-June (QU, VT). Tenerals: mid-July to September; mostly August. Adults found overwintering on higher and drier ground, about 3 to 7 m from their summer habitat, at the edge of swamps and woods, in woods, pastures, on hills, in the upper zone of sand pits; in the soil, under stones, logs, fallen pieces of bark, and the loose bark of fallen trees. Adult food, in captivity: mealworms and meat. Larval food, in captivity: mealworms. Predators: frogs, toads, bluebirds, thrushes, and spiders. Parasites: podapolipid mites and Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult may emit a musky smoke-like smell from the pygidial glands, that adhers to fingers when the beetle is seized. Dispersal power. Macropterous. Frequent

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flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on trees). Collecting techniques. Treading the soil with the feet; treading the vegetation into the water; light trapping. References. Blatchley, 1910 (ecology); Lindroth, 1955a, 1963a, and 1969a (natural history); Bell, 1960 (biology); Kirk, 1975c (natural history); Erwin, 1981 (natural history). Chlaenius (Chlaeniellus) pertinax Casey, 1920 Ecology. Lowlands. Biology. Tenerals: March. Dispersal power. Macropterous, probably capable of flight. Chlaenius (Eurydactylus) pimalicus Casey, 1914 Ecology. Mountains. Dry open situations. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Chlaenius (Chlaenius) platyderus Chaudoir, 1856 Ecology. Lowlands and lower mountains. Cultivated fields (e.g., alfalfa, soybean, corn, flax, clover, rye), wastelands, pastures, fencerows, hills, sand pits, gravel pits, and forests (open; flood plain). Open ground; moderately moist soil covered with some vegetation. Nocturnal; sheltering during the day under stones, logs, and dry cow dung. Biology. Seasonality: March-October. Tenerals: mid-July (KS) and September (SD). Adult food, in the field: lepidopterous caterpillars, coleopterous larvae, fungi, and plant material. Predators: raccoons. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones and logs. References. Bell, 1960 (natural history); Kirk, 1975c (ecology, dispersal power). Chlaenius (Chlaenius) prasinus Dejean, 1826 Ecology. Lowlands and mountains. Borders of rivers (preferably large), brooks, lakes, and ponds. Open ground; moist, sandy or sandy-gravelly, often silt-mixed, bare soil. Nocturnal; sheltering during the day under stones and pieces of wood. Larval habitat: under stones. Biology. Seasonality: January-December. Tenerals: September (IL, FL). Adult food, in the field: insects. Larval food, in captivity: mealworms. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; turning stones and pieces of wood. References. Blatchley, 1910 (ecology); Bell, 1960 (natural history); Thompson and Allen, 1974 (ecology, biology); Erwin, 1981 (natural history). Chlaenius (Chlaenius) purpureus Chaudoir, 1876 Ecology. Mountains. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight.

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Chlaenius purpuricollis frostii Carr, 1920 Ecology. Uplands. Biology. Seasonality: February, May-June. Dispersal power. Macropterous, probably capable of flight. Chlaenius purpuricollis purpuricollis Randall, 1838 Ecology. Lowlands, mountains, and subalpine zone. Alvar pavements, alvar savanna grasslands, alvar grasslands, tall-grass prairies, meadows, sandhills, sand dunes, rocky slopes, and city yards. Not associated with water. Open ground; dry limestone or sandy soil covered with relatively dense grass, sparse herbaceous vegetation or shrubs. Biology. Seasonality: April-September. Tenerals: July-September. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pitfall trapping. References. Bouchard et al. 1998 (ecology, biology); Tinerella and Rider, 2001 (ecology). Chlaenius (Anomoglossus) pusillus Say, 1823 Ecology. Lowlands. Deciduous forests, coniferous forests (e.g., Pinus), orchards, and hedges; pastures and cultivated fields (e.g., corn, tobacco, soybean, cotton, sugarcane); vicinity of rivers and brooks. Shaded or open ground; moderately moist soil covered with dead leaves or some vegetation. Nocturnal; sheltering during the day in leaf litter, under stones, and debris. Biology. Seasonality: January-October. Tenerals: August-September. Adults found overwintering under pieces of wood. Adult food, in the field: lepidopterous eggs and staphylinids. Predators: ducks. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; light trapping; raking the leaf litter. References. Blatchley, 1910 (ecology); Leng, 1915 (ecology); Lee, 1978 (ecology, dispersal power); Kirk, 1969 (dispersal power); Lindroth, 1969a (dispersal power); Hsin et al. 1979 (biology). Chlaenius (Callistometus) ruficauda Chaudoir, 1856 Ecology. Mountains. River banks and other damp situations. Nocturnal; sheltering during the day under logs. Biology. Seasonality: September. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning logs. Chlaenius (Chlaenius) sericeus regularis LeConte, 1851 Ecology. Lowlands and mountains. Biology. Seasonality: January-July. Dispersal power. Macropterous, probably capable of flight. Moderate runner.

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Chlaenius (Chlaenius) sericeus sericeus (Forster, 1771) Ecology. Lowlands and mountains. Borders of lakes, ponds, reservoirs, pools (e.g., in gravel pits, sand pits), cattle water holes, rivers, brooks, and marshes; swamps, wet pastures, and beaver houses. Cultivated fields (e.g., corn, oat, wheat, tobacco, cabbage, potato, carrot), gardens, orchards (e.g., apple), and open forests. Open ground; rather firm, moderately moist soil consisting of clay, mud or gravel, moderately rich in organic debris, covered with some vegetation (e.g., grass, Carex). Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering during the day under stones (mostly), pieces of wood, and other debris. Gregarious (in winter). Larval habitat: under stones and pieces of wood. Biology. Seasonality: March-December. Copulating pairs: May-June (CT, QU); coitus lasting 1-10 minutes. Gravids: May to early July (QU, VT). Eggs are laid in mud or clay cells stuck on stones or on the lower part of dead plant stems lying on the surface of moist ground. Tenerals: mid-July to November; mostly August. Adults found overwintering on higher and drier ground, in the upper zone of sand pits and gravel pits, on hills, in pastures, at the edge of woods, and in woods; in the soil (2.5-20 cm deep), under well-embedded stones, and pieces of wood. Adult food: carabids, scarabaeids, wireworms, other insects, and earthworms (in the field); mealworms, carabids, dead or dying insects, earthworms, dead slugs, meat, and grapefruit (in captivity). Larval food, in captivity: mealworms. Predators: frogs, toads, crows, and raccoons. Parasites: nematodes. Defense-mechanism: when alarmed, the adult may emit a strong pungent musky odor that clings to fingers when the beetle is seized. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Regular in sea- and lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on trees). Strong burrower. Good swimmer. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; light trapping; turning drift material. References. Wickham, 1895 (biology); Blatchley, 1910 (ecology); King, 1919 (biology); Lindroth, 1955a, 1963a, and 1969a (natural history); Bell, 1960 (natural history); Kirk, 1969 (dispersal power) and 1975c (natural history); Erwin, 1981 (natural history). Chlaenius (Chlaenius) sericeus sierricola Casey, 1914 Ecology. Mountains. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Chlaenius (Chlaenius) sericeus viridifrons Eschscholtz, 1833 Ecology. Unknown. Biology. Seasonality: December. Dispersal power. Macropterous, probably capable of flight. Chlaenius (Chlaeniellus) simillimus Chaudoir, 1856 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight.

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Chlaenius (Chlaenius) solitarius Say, 1823 Ecology. Lowlands and uplands. River banks. Open ground; moist, muddy or sandy, bare soil. Nocturnal; sheltering during the day under logs, dead branches, and in the soil. Gregarious (in spite of its name). Biology. Seasonality: April, June-October, December. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; turning logs. Reference. Bell, 1960 (dispersal power). Chlaenius (Chlaeniellus) texanus G.H. Horn, 1876 Ecology. Lowlands. Borders of lakes and rivers. Open ground; wet, muddy soil, bare or covered with sparse vegetation. Nocturnal; sheltering during the day under stones. Biology. Seasonality: June-August. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Turning stones. Chlaenius (Eurydactylus) tomentosus tomentosus (Say, 1823) Ecology. Lowlands, mountains, subalpine and alpine zones. Meadows, pastures, rangelands, relict prairies, old fields, vacant lots, gardens, cultivated fields (e.g., corn, wheat, hay, oat, clover, alfalfa, soybean, tobacco, flax, strawberry), fencerows, roadsides, sand pits, gravel pits, sand dunes, sand hills, orchards (e.g., peach, plum), and open forests. In the countryside and in cities. Open ground; dry, sandy soil covered with some vegetation (e.g., grass, weeds). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under embedded stones (mostly), dead leaves and plants, reindeer moss (Cladonia), plant rosettes (Verbascum), and logs. Biology. Seasonality: February-December. Copulating pairs: May-June (QU); coitus lasting about 9 minutes. Gravids: early June (QU). Tenerals: August-September, November. Adults found overwintering on higher and drier ground, in vacant lots and on hills; in the soil under well-embedded stones (mostly), dead leaves, and plant rosettes (Verbascum). Adult food: lepidopterous caterpillars, scarabaeids, other insects, earthworms, fungi, and vegetal tissue (in the field); earthworms, dead carabids, and meat (in captivity). Predators: toads, bluebirds, crows, starlings, and raccoons. Parasites: nematodes and tapeworms. Defense-mechanism: when alarmed, the adult may emit a musky or smoke-like smell from the pygidial glands. Dispersal power. Macropterous. Frequent flier (sometimes swarming to artificial lights at night). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on trees). Strong burrower. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning embedded stones; light trapping; turning drift material. References. Blatchley, 1910 (dispersal power); Bell, 1960 (natural history); Kirk, 1969 (dispersal power) and 1975c (natural history); Lindroth, 1969a (natural history) and 1970 (biology); Larochelle, 1974a (ecology); Erwin, 1981 (natural history).

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Chlaenius (Eurydactylus) tomentosus zunianus Casey, 1914 Ecology. Mountains. A cut grassy field (AZ). Biology. Seasonality: September. Adult food, in the field: lepidopterous caterpillars. Dispersal power. Macropterous, probably capable of flight. Chlaenius (Chlaeniellus) tricolor tricolor Dejean, 1826 Ecology. Lowlands and mountains. Border of large rivers, lakes, ponds, marshes, and pools; flood-plain forests. Deciduous forests: along edges, paths, and openings. Orchards (e.g., apple), pastures, and cultivated fields (e.g., tobacco, barley, asparagus, alfalfa, soybean, potato, corn, carrot, cabbage). Often at a considerable distance from water. Open or shaded ground; firm, wet or moist, gravelly, sandy or muddy soil. Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering under well-embedded stones (mostly), logs, and dead leaves. Gregarious (in winter). Biology. January-December. Copulating pairs: mid-May (QU) to mid-June (CT); coitus lasting about 22 minutes. Gravids: early June (QU). Eggs are laid in mud cells stuck to the stems of sedges (e.g., Eleocharis) and grasses. Tenerals: late May (rarely; VT), August-November (QU, VT). Adults found overwintering on higher and drier ground, in colonies with Brachinus, in the upper zone of sand pits and gravel pits, in vacant lots and pastures, on hills, in clearings, at the edge of woods, and in woods; under well-embedded stones and in the soil (mostly; 17.5-50 cm deep), and under pieces of wood. Adult food: lepidopterous caterpillars and slugs (in the field); carabids, mealworms, earthworms, dead slugs, and meat (in captivity). Larval food, in captivity: mealworms. Predators: frogs, toads, crows, and starlings. Parasites: hymenopterans (egg parasites) and Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult may emit a strong musky smell from the pygidial glands, that clings to fingers when the beetle is seized. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants and trees). Strong burrower. Strongly favored by human activities. Collecting techniques. Pitfall trapping; lifting embedded stones; light trapping; sweeping plants. References. King, 1919 (biology); Bell, 1960 (ecology); Lindroth, 1963a and 1969a (natural history); Kirk, 1975c (ecology, dispersal power); Erwin, 1981 (natural history); Will et al. 1995 (biology, dispersal power). Chlaenius (Chlaeniellus) tricolor vigilans Say, 1830 Ecology. Lowlands and uplands. Greenhouses (occasionally). Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Chlaenius (Chlaeniellus) vafer LeConte, 1852 Ecology. Lowlands. Edges of deciduous forests. Nocturnal; sheltering during the day. Biology. Seasonality: May, July-August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Reference. Kirk, 1975c (dispersal power).

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Chlaenius (Chlaeniellus) variabilipes Eschscholtz, 1833 Ecology. Edge of a water hole in the coastal prairie (TX). Grasslands (Mexico). Biology. Seasonality: April, December. Dispersal power. Macropterous, probably capable of flight. Reference. Bell, 1960 (ecology). Chlaenius (Chlaenius) viduus G.H. Horn, 1871 Ecology. Uplands. Biology. Seasonality: January, March-April. Adults found overwintering under pieces of wood and in rotten wood. Dispersal power. Brachypterous, incapable of flight. Clivina americana Dejean, 1831 Ecology. Lowlands. Margins of eutrophic marshes, ponds, lakes, reservoirs, slow rivers and brooks; swamps, flood-plain forests, bogs, wet meadows, and beaver houses. Open or half-shaded ground; moist or wet soil consisting of mud, clay or sand-mixed clay, rich in organic debris (e.g., leaf litter), covered with carpets of depressed grass. Nocturnal; sheltering during the day in burrows dug in the soil, under mats of dead plants, well-embedded fallen branches, logs, stones, dead leaves, and in decaying floating debris. Biology. Seasonality: January-December. Tenerals: April (rarely; FL), mid-June to October. Adults found overwintering on higher and drier ground, at the edge of woods and in the upper zone of lake shores; in burrows dug in the soil (as deep as 10 cm), under well-embedded stones, and under pieces of wood. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Slow runner. Strong burrower. Clearly effective as a colonist. Collecting techniques. Treading the vegetation into the water; treading the soil with the feet; raking the leaf litter; light trapping. References. Leng, 1915 (ecology, dispersal power; Lindroth, 1961 (natural history); Kirk, 1969 (biology, dispersal power) and 1975c (dispersal power); Erwin, 1981 (natural history); Nichols, 1988 (ecology); Will et al. 1995 (ecology). Clivina analis Putzeys, 1846 Natural History. Unknown. Clivina (Paraclivina) bipustulata (Fabricius, 1801) Ecology. Lowlands. Margins of rivers, brooks, ponds, and lakes. Cultivated fields (e.g., corn, tobacco, cotton, wheat, alfalfa, soybean), gardens, vacant lots, roadsides, and orchards (e.g., peach). Open ground; soil wet or moist, muddy or clayish, rich in organic debris, covered with some vegetation. Nocturnal; sheltering during the day in burrows dug in the soil in layers of leaf litter mixed with silt (mostly), under stones, and logs. Associated with staphylinids (Bledius) and ants (Formica). Biology. Seasonality: January-November. Tenerals: August-September (MD). Adults found overwintering in wood debris. Predators: ducks and trashers. Dispersal power. Macropterous. Frequent flier (to artificial lights at night).

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Occasional in lakeshore drift material, which indicates flight ability. Slow runner. Strong burrower. Strongly favored by human activities. Collecting techniques. Treading the soil with the feet; pouring water over the ground; pitfall trapping; baiting traps with molasses and fermented sugar; light trapping. References. Leng, 1915 (ecology, dispersal power; Lindroth, 1961 (natural history); Kirk, 1969 (biology, dispersal power) and 1975c (dispersal power); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Clivina californica Van Dyke, 1925 Ecology. Lake shores. Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: May, November. Dispersal power. Wing-condition unknown. Slow runner. Reference. Van Dyke, 1925 (ecology). Clivina (Clivina) collaris (Herbst, 1784) Ecology. Lowlands. Cultivated fields, vacant lots, and gravel pits. Gardens and greenhouses (Europe). Open ground; black, moderately moist soil, rich in humus, bare or covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil (mostly; 10 cm deep) or under pieces of wood. Biology. Seasonality: March-September. Predators: toads. Defense-mechanism: when alarmed, the adult may slowly emit a discharge from the pygidial glands, at a range shorter than its body length. Dispersal power. Macropterous. Flight records (Europe). Slow runner. Strong burrower. Clearly effective as a colonist. Collecting technique. Pitfall trapping. References. Larsson, 1939 (biology); Lindroth, 1945, 1961, and 1985 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology). Clivina (Paraclivina) convexa LeConte, 1844 Ecology. Unknown. Biology. Seasonality: June-July. Dispersal power. Unknown. Clivina (Semiclivina) dentipes Dejean, 1825 Ecology. Lowlands and mountains. Margins of rivers, brooks, irrigation canals, lakes, pools, and sloughs; flood-plain forests. Open ground; well-drained soil composed of sand-mixed clay, clay or mud, bare or with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil (mostly), under stones, logs, pieces of wood, and in leaf litter. Biology. Seasonality: January-October, December. Tenerals: mid-July and August. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Slow runner. Strong burrower. Collecting techniques. Treading the soil with the feet; pouring water over the ground; light trapping; using interception traps. References. Löding, 1945 (ecology); Kirk, 1969 and 1975c (dispersal power); Erwin, 1981 (natural history). Clivina (Paraclivina) fasciata Putzeys, 1846 Ecology and Biology. Unknown. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner.

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Clivina (Paraclivina) ferrea LeConte, 1857 Ecology. Lowlands and uplands. Banks of rivers and brooks. Cultivated fields (e.g., corn, cotton); orchards (e.g., peach). Open ground; moist, sandy or silty soil covered with some vegetation. Nocturnal; sheltering during the day in burrows dug in the soil, under stones and other debris. Biology. Seasonality: February-October, December. Tenerals: July-August (MS). Adults found overwintering in woods; under logs. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Occasional climber (on plants). Strong burrower. Collecting techniques. Pitfall trapping; sweeping. References. Kirk, 1975c (ecology, dispersal power); Erwin, 1981 (natural history). Clivina (Paraclivina) floridae Csiki, 1927 Ecology. A bay beach, on damp, sandy soil. Nocturnal; sheltering during the day in burrows dug in the soil and under pieces of wood. Biology. Seasonality: December. Dispersal power. Wing-condition unknown. Slow runner. Clivina (Clivina) fossor (Linné, 1758) Ecology. Lowlands and mountains. Cultivated fields (e.g., corn, carrot, clover, barley, bean, raspberry, potato), pastures, meadows, vacant lots, lawns, parks, orchards, gardens, roadsides, gravel pits, and sand pits. Edges of pools, ponds, marshes, lakes, and slow rivers. Forest edges, open forests, and flood-plain forests. Open ground (mostly) or lightly shaded ground; moderately moist to wet, usually clay soil covered with some vegetation. Mostly nocturnal; usually sheltering during the day in burrows dug in the soil (as deep as 20cm), under stones, pieces of wood, dead leaves; sometimes active on plants. Active over the entire 24 hour cycle, but reaching its maximum activity in the evening (Europe). Reproductive adults are mostly grounddwelling while larvae are subterranean. Biology. April-October. Gravids: early July (QU). Tenerals: April-May, July-October. Adults found overwintering on higher and drier ground, in the upper zone of gravel pits, in vacant lots, at the edge of woods, and in woods; under stones (mostly) pieces of wood, and the loose bark of fallen trees. Adults hibernate at their breeding site in Europe. Adult food: wireworms (in the field); chrysomelid eggs and larvae, curculionid immature stages, seed carrot, plant material (Europe), insects (Europe), and worms (Europe) (in captivity). Economic importance: sometimes a pest of corn, sugar beet, and strawberry (Europe). Predators: frogs and toads. Defense-mechanism: when alarmed, the adult may feign death for a few seconds or slowly emit a discharge from the pygidial glands (at a range shorter than its body length). Dispersal power. Wing-polymorphic: macropterous (capable of flight) to brachypterous (incapable of flight). Frequent flier (sometimes swarming; at sunset; to artificial lights at night). Frequent in drift material, which indicates flight ability. Slow runner. Frequent climber (on plants, e.g., Gramineae). Strong burrower. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Pitfall trapping; pouring water over the ground;

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treading the soil with the feet; sweeping plants; light trapping; turning drift material. References. Lindroth, 1945, 1955a, 1961, and 1963a (natural history); Finlayson and Campbell, 1976 (biology); Tyler and Ellis, 1979 (biology); Sharova, 1981 (ecology, biology); Desender, 1983 (ecology, biology); Desender and Pollet, 1985 (biology, dispersal power); Lindroth, 1985 (natural history); Levesque and Levesque, 1987 (biology); Nelson and Reynolds, 1987 (ecology, dispersal power); Koch, 1989 (ecology); Luff, 1993 (biology); Levesque and Levesque, 1994 (biology). Clivina (Clivina) impressefrons LeConte, 1844 Ecology. Lowlands and mountains. Cultivated fields (e.g., corn, soybean, barley, asparagus). Banks of rivers and brooks; vicinity of cattle water holes; flood-plain forests. Open ground; low, moist or wet soil consisting of peat, mud, clay or sandy clay, covered with some vegetation. Crepuscular and nocturnal; sheltering during the day in burrows dug in the soil, under stones, and pieces of wood. Gregarious (in winter). Larval habitat: burrows dug in the soil (as deep as 60 cm). Biology. Seasonality: March-November. Tenerals: July (OH), September (TX). Adult food, in the field: lepidopterous caterpillars and corn seeds. Adult and larval food, in captivity: chrysomelid eggs and larvae, rarely corn seed. Larval food, in the field: animal matter and corn seed. Adults and larvae are primarily carnivorous, but feed on plant matter when their primary food is missing or in short supply. Larvae are cannibalistic in captivity. Economic importance: sometimes a severe pest of corn. Predators: swallows. Parasites: podapolipid mites and Laboulbeniales (Fungi). Dispersal power. Macropterous. Frequent flier (sometimes swarming; at sunset; to artificial lights at night). Frequent in lakeshore drift material, which indicates flight ability. Slow runner. Strong burrower. Strongly favored by human activities. Collecting techniques. Pitfall trapping; pouring water over the ground; treading the soil with the feet; light trapping; turning drift material. References. Webster, 1906 (biology); Phillips, 1909 (biology); Blatchley, 1910 (ecology, biology); Lindroth, 1961 (natural history); Bell, 1971 (ecology); Sechriest et al. 1971 (biology); Kirk, 1975c (ecology, dispersal power); Pausch and Pausch, 1980 (biology); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Clivina (Paraclivina) marginipennis Putzeys, 1846 Ecology and Biology. Unknown. Dispersal power. Macropterous. Occasional flier (to artificial lights at light). Clivina morula LeConte, 1857 Ecology. Lowlands. River banks. Moist or wet soil. Nocturnal; sheltering during the day in burrows dug in the soil and under stones. Biology. Seasonality: January-March, July-October. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet.

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Clivina (Clivina) myops Bousquet, 1997 Ecology. Unknown. Biology. Seasonality: April. Dispersal power. Macropterous, probably capable of flight. Reference. Bousquet, 1997b (taxonomy, dispersal power). Clivina (Clivina) oregona Fall, 1922 Ecology. River banks. Open ground; moist, sandy, clayish or muddy soil. Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: April-July. Dispersal power. Wing-dimorphic: macropterous (probably capable of flight) or brachypterous (incapable of flight). Slow runner. Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Reference. Lindroth, 1961 (natural history). Clivina (Clivina) pallida Say, 1823 Ecology. Lowlands. Forests and saw mills. Moist soil. Nocturnal; sheltering during the day under tree bark (Pinus), in tree hollows, fermenting pieces of wood and saw dust, and burrows dug in the soil under stones. Biology. Seasonality: January-July, September-October, December. Adults found overwintering in tree hollows. Dispersal power. Wing-condition unknown. Slow runner. Occasional climber (on trees). Strong burrower. Collecting technique. Sifting (e.g., with Berlese funnels) fermenting pieces of wood or sawdust. References. Löding, 1945 (ecology); Kirk, 1969 (ecology, biology); Nelson, 1991 and 1995 (ecology). Clivina (Clivina) planicollis LeConte, 1857 Ecology. Unknown. Biology. Seasonality: April-September. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Reference. Kirk, 1969 (dispersal power). Clivina (Paraclivina) postica LeConte, 1848 Ecology. On a dock (LA). Biology. Seasonality: May-August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Reference. Kirk, 1975c (dispersal power). Clivina (Clivina) punctigera LeConte, 1857 Ecology. Lowlands. Swamps. Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: May-August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. References. Ulke, 1903 (dispersal power); Kirk, 1969 (dispersal power).

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Clivina (Clivina) punctulata LeConte, 1852 Ecology. Moist situations. Biology. Seasonality: April-May, July, November. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. Clivina (Antroforceps) rubicunda LeConte, 1857 Ecology. Lowlands. A beaver pond. A woodland pool. A hammock forest. Close to water or at some distance from it. Nocturnal; sheltering during the day in burrows dug in the soil, under logs and pieces of wood, and in leaf litter. Biology. Seasonality: JanuaryJune, August. Adults found overwintering (IN). Dispersal power. Wing-dimorphic: macropterous or brachypterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. Collecting technique. Pitfall trapping. References. Blatchley, 1910 (ecology, biology); Ball, 2001 (taxonomy, natural history). Clivina rufa LeConte, 1857 Ecology. Lowlands. Banks of river and brooks. Moist, muddy soil. Nocturnal; sheltering during the day in burrows dug in the soil or under stones. Biology. Seasonality: January-December, except November. Adults found overwintering. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Clivina (Antroforceps) sasajii Ball, 2001 Ecology. Uplands. In a forest, in the burrow of a Pocket Gopher. Biology. Seasonality: April, December. Dispersal power. Brachypterous, incapable of flight. Reference. Ball, 2001 (taxonomy, natural history). Clivina (Paraclivina) stigmula Putzeys, 1846 Ecology. Lowlands. Biology. Seasonality: August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Clivina (Paraclivina) striatopunctata Dejean, 1831 Ecology. Lowlands. Swamps, lake shores, brook banks, salt meadows, cultivated fields (e.g., cotton), and orchards. Nocturnal; sheltering during the day in burrows dug in the soil and under debris. Biology. Seasonality: February-December, except April. Predators: catbirds, woodpeckers, and robins. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Slow runner. Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping. Reference. Leng, 1915 (ecology).

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Clivina (Paraclivina) sulcipennis Putzeys, 1866 Ecology. Lowlands. Biology. Seasonality: March, June-August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Clivina (Semiclivina) vespertina Putzeys, 1866 Ecology. Lowlands. Biology. Seasonality: April-June. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Colliuris (Cosnania) lengi (Schaeffer, 1910) Ecology. Open forests (Quercus). Dry soil. Sheltering during the day on oak trees. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Reference. Ball and Bousquet, 2001 (ecology). Colliuris (Cosnania) lioptera (Bates, 1891) Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Colliuris (Calocolliuris) ludoviciana (Sallé, 1849) Ecology. Lowlands. Edges of swamps, marshes, and eutrophic ponds; meadows; floodplain forests. Open ground; wet, muddy or sandy soil covered with some vegetation (e.g., grass, Carex). Mostly diurnal; active in the sunshine on plants; sheltering on cloudy or cool days under pieces of wood. Gregarious. Biology. Seasonality: January-December. Adults found overwintering near swamps; under boards and other pieces of wood. Adult food, in captivity: small insects. When the beetle seizes its prey, it proceeds to drag it off, imitating certain ant species. Defense-mechanism: at the least noise, the adult escapes by running rapidly while vibrating its antennae. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Frequent climber (on plants). Collecting techniques. Treading the vegetation (grass, Carex) into the water; sweeping plants. References. Summers, 1873 (ecology, biology); Ulke, 1903 (ecology); Davidson, 1995 (ecology). Colliuris (Cosnania) pensylvanica (Linné, 1767) Ecology. Lowlands, mountains, subalpine and alpine zones. Grasslands, meadows, pastures, lawns, abandoned fields, old fields, cultivated fields (e.g., corn, alfalfa, barley, soybean, cotton, oat, tobacco, peanut), stubble fields, fencerows, ski fields, sand pits, moist areas near marshes, pond edges, peat bogs, orchards, forest edges, and open forests. Open ground; usually dry soil covered with some vegetation (e.g., grass or weeds). Diurnal and nocturnal; active in the sunshine on plants (Solidago, Verbascum, Medicago) or sheltering on cloudy or cool days at the roots of plants, in clumps of

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broomsedges, under stones, logs, pieces of wood, in leaf litter, and under heaps of hay. Biology. Seasonality: January-December. Adults found overwintering on higher and drier ground, in open places, fencerows, and in woods; under stones, dead leaves, wood debris, and in ant nests. Adult food, in the field: lepidopterous caterpillars and eggs (mostly), lygaeids, and nabid nymphs. Predators: toads, salamanders, quails, crows, sparrows, starlings, and swallows. Also eaten by insectivorous plants (Sarracenia). Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Frequent in lake- and seashore drift material, which indicates flight ability. Fast runner. Regular climber (on plants). Strongly favored by human activities. Collecting techniques. Sweeping the vegetation; pitfall trapping; sweeping plants; light trapping; turning drift material. References. Schwarz, 1890 (biology); Blatchley, 1910 (ecology); Leng, 1915 (ecology); Kirk, 1969 (biology, dispersal power), 1970 (ecology) and 1975c (ecology, dispersal power); Lindroth, 1969a (natural history); Erwin, 1981 (natural history). Colliuris (Cosnania) picta picta (Chaudoir, 1843) Ecology. Unknown. Biology. Seasonality: January, May. Dispersal power. Macropterous, probably capable of flight.

Colpodes buchannani Hope, 1831, see Metacolpodes buchannani (Hope, 1831). References. Liebherr, 1998 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Coptodera aerata Dejean, 1825 Ecology. Lowlands and mountains. Deciduous forests (e.g., Quercus, Liriodendron). Mostly nocturnal; active at night on logs or fallen tree trunks; sheltering during the day under the loose bark of trees, on tree trunks, tree-stumps and logs, in dead wood crevices, in the vicinity of piled wood, and under dead leaves; sometimes active on cloudy days on the vegetation. Biology. Seasonality: January, April-October, December. Tenerals: midJuly (IN). Adults found overwintering in woods; under the loose bark of trees. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Fast runner. Frequent climber (on trees and plants). Collecting techniques. Lifting the loose bark of trees; light trapping; beating or sweeping the vegetation; baiting traps with carrion. References. Ulke, 1903 (ecology); Löding, 1945 (ecology); Kirk, 1969 (ecology, dispersal power); Erwin, 1981 (natural history); Ball and Bousquet, 2001 (ecology). Coptodera brunnea Shpeley and Ball, 1993 Ecology. Mountains. A pasture tree, on bromeliads (Mexico). Biology. Seasonality: February, June-August. Dispersal power. Macropterous. Occasional flier (to artificial lights

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at night). Collecting techniques. Light trapping; bait trapping. Reference. Shpeley and Ball, 1993 (taxonomy, natural history). Coptodera festiva Dejean, 1825 Ecology. Lowlands. A pine forest (FL). Tropical forests (Mexico, Trinidad, Peru). Nocturnal; sheltering during the day, e.g., in a wood pile. Biology. Seasonality: July-September, December. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Collecting techniques. Light trapping; fogging tree canopy with insecticides; baiting traps with rotting lemons; using glass fruitfly traps. Reference. Shpeley and Ball, 1993 (taxonomy, natural history). Coptodera nitidula (Buquet, 1834) Ecology. Mountains. Forests (e.g., Quercus). Nocturnal; sheltering during the day under the loose bark of trees and in bromeliads. Biology. Seasonality: July-September. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Climber (on trees). Collecting techniques. Lifting the loose bark of trees; light trapping. Reference. Shpeley and Ball, 1993 (taxonomy, natural history). Coptodera picea Dejean, 1826 Ecology. Lowlands. Forests (e.g., palmettos). Nocturnal; sheltering during the day on trees (e.g., bromeliads, leaves) and logs. Biology. Seasonality: April. Dispersal power. Macropterous, probably capable of flight. Climber (on trees). Reference. Shpeley and Ball, 1993 (taxonomy, natural history). Cratacanthus dubius (Palisot de Beauvois, 1811) Ecology. Lowlands and mountains. Cultivated fields (e.g., corn, wheat, oat, alfalfa, clover, flax, tobacco, soybean), relict prairies, rangelands, pastures, wastelands, old fields, fencerows, lawns, hills, blowouts; vicinity of rivers, brooks, lakes, and ponds; open forests. Open ground; dry clay soil, sometimes sandy, covered with some vegetation (e.g., grass). Mostly nocturnal; sometimes active in the sunshine in spring; usually sheltering in burrows dug in the soil (mostly), under heaps of grass, dry cow dung, stones, logs, sticks, and boards. Biology. Seasonality: January-November. Tenerals: June and August (VA), July (SD). Adults found overwintering under stones. Adult food, in the field: small insects. Predators: toads, bluebirds, catbirds, crows, flycatchers, robins, sparrows, starlings, trashers, and thrushes. Parasites: gregarines and tapeworms. Dispersal power. Wing-dimorphic: macropterous (capable of flight) or brachypterous (incapable of flight). Frequent flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Strong burrower. Strongly favored by human activities. Collecting techniques. Pitfall trapping; light trapping; turning debris; sweeping. References. Blatchley, 1910 (ecology); Lindroth, 1968 (natural history); Kirk, 1975c

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(natural history); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power); Ball and Bousquet, 2001 (ecology). Cychrus hemphillii hemphillii G.H. Horn, 1878 Ecology. Uplands and mountains. Fragments of forests and brushes (e.g. Alnus, Philadelphicus): along rivers and canyons. A coniferous forest (Populus-Abies) (ID). Shaded ground; rather dry soil covered with dense vegetation. Nocturnal; sheltering during the day. Biology. Seasonality: May-June, September. Tenerals: June (ID). Adult food: snails (in the field); snails and slugs (in captivity). Defense-mechanism: when alarmed, the adult may emit a pungent defensive fluid from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Slow runner (with tenebrionid-like locomotion). Collecting technique. Pitfall trapping. Reference. Greene, 1975 (ecology, biology). Cychrus hemphillii rickseckeri LeConte, 1884 Ecology. Mountains. Mixed and coniferous forests: often along streams. Shaded ground; rather dry soil covered with some vegetation (e.g., weeds or grass). Nocturnal; sheltering during the day under stones and big pieces of wood. Biology. Seasonality: MayAugust. Tenerals: June. Adult food, in the field: snails. Dispersal power. Brachypterous, incapable of flight. Slow runner (with tenebrionid-like locomotion). Collecting techniques. Pitfall trapping; turning stones and big pieces of wood. References. Stace Smith, 1948 (ecology, biology); Lindroth, 1961 (natural history); LaBonte, 1988 (ecology). Cychrus tuberculatus T.W. Harris, 1839 Ecology. Lowlands, mountains, subalpine and alpine zones. Coniferous forests, forest clearings, and moraines. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under logs and stones. Biology. Seasonality: April-October. Tenerals: May-July. Adult food, in the field: mollusks. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning logs and stones. References. Lindroth, 1961 (natural history); Kavanaugh, 1992 (natural history). Cyclotrachelus (Evarthrus) alabamae (Van Dyke, 1926) Ecology. Lowlands. Mixed forests (Pinus, Quercus). Shaded ground; moderately moist soil. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: JanuaryMarch, September-October. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. Reference. Freitag, 1969 (ecology). Cyclotrachelus (Cyclotrachelus) alabamensis (Casey, 1920) Ecology. Lowlands. Coniferous forests (Pinus). Open ground; moderately moist, sandy soil covered with sparse vegetation. Nocturnal; sheltering during the day under dead

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leaves and pieces of wood. Biology. Seasonality: February, July, November. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning debris. Reference. Löding, 1945 (ecology). Cyclotrachelus (Evarthrus) alternans (Casey, 1920) Ecology. Uplands. Cultivated fields (e.g., corn, oat, flax, alfalfa, sorghum, wheat, soybean), relict prairies, gardens, and lawns. Open ground; dry, sandy soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day under stones and logs. Biology. Seasonality: April-October. Copulating pairs: June-July (SD). Adult food, in captivity: lepidopterous caterpillars, dead and live invertebrates, and seeds. Parasites: rickettsias. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones and logs; light trapping. Reference. Kirk, 1975c (natural history). Cyclotrachelus (Cyclotrachelus) approximatus (LeConte, 1848) Ecology. Lowlands and mountains. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under pieces of wood. Biology. Seasonality: JuneJuly, September. Tenerals: July (MD). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning pieces of wood. Reference. Erwin, 1981 (natural history). Cyclotrachelus (Evarthrus) blatchleyi (Casey, 1918) Ecology. Lowlands. Open dry disturbed situations. Nocturnal; sheltering during the day. Biology. Seasonality: March-April, June-July, September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; baiting traps with carrion. Reference. Freitag, 1969 (ecology). Cyclotrachelus (Cyclotrachelus) brevoorti (LeConte, 1848) Ecology. Lowlands. Forests (e.g., Quercus, Pinus). Shaded ground; moderately moist soil. Nocturnal; sheltering during the day in leaf litter and rotten logs. Biology. Seasonality: March-May, August-November. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. Reference. Freitag, 1969 (ecology). Cyclotrachelus (Evarthrus) constrictus (Say, 1823) Ecology. Uplands. Grasslands, prairies, cultivated fields (e.g., corn), and pastures. Open ground; dry soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day under stones and logs. Biology. Seasonality: April-October. Tenerals: June (SD). Parasites: tapeworms. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones and

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logs. References. Freitag, 1969 (ecology); Bell, 1971 (ecology); Kirk, 1975c (ecology, biology); Lavigne, 1978 (ecology). Cyclotrachelus (Evarthrus) convivus (LeConte, 1852) Ecology. Lowlands and mountains. Deciduous and mixed forests: often along rivers. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day in leaf litter and under logs. Biology. Seasonality: January-December. Adult food, in the field: small insects. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter; turning logs. References. Freitag, 1969 (ecology); Allen and Thompson, 1977 (ecology, biology). Cyclotrachelus (Cyclotrachelus) dejeanellus (Csiki, 1930) (Cyclotrachelus morio (Dejean, 1828)) Ecology. Lowlands. Deciduous (e.g., Quercus) or coniferous (e.g., Pinus) forests. Shaded ground; moderately moist, sandy soil. Nocturnal; sheltering during the day in thick leaf litter, under pieces of wood, and bunches of fallen Spanish moss. Biology. Seasonality: January-December. Tenerals: spring (mostly) and autumn (FL). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. Reference. Leng, 1915 (ecology; as morio (Dejean)); Freitag, 1969 (ecology; as morio (Dejean)); Harris and Whitcomb, 1971 and 1974 (ecology, biology; as morio (Dejean)). Cyclotrachelus (Evarthrus) engelmani (LeConte, 1852) Ecology. Lowlands. Forests. Shaded ground; moderately moist soil. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Freitag, 1969 (ecology). Cyclotrachelus (Cyclotrachelus) faber (Germar, 1824) Ecology. Lowlands. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: January-March, July-August. Adult food, in the field: fungal spores. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. Reference. Freitag, 1969 (ecology, biology). Cyclotrachelus (Evarthrus) floridensis (Freitag, 1969) Ecology. Lowlands. Biology. Seasonality: February. Dispersal power. Brachypterous, incapable of flight.

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Cyclotrachelus (Cyclotrachelus) freitagi Bousquet, 1993 Ecology. Lowlands and mountains. Deciduous forests; coniferous forests (e.g., Pinus). Shaded ground; moderately moist soil. Nocturnal; sheltering during the day in leaf litter, under logs, sticks, and stones. Biology. Seasonality: January, March-November. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter; night collecting with a headlamp. References. Leng, 1915 (ecology; as obsoleta (Say)); Freitag, 1969 (ecology; as obsoletus (Say)); Will et al. 1995 (dispersal power). Cyclotrachelus (Cyclotrachelus) fucatus (Freitag, 1969) Ecology. Lowlands and mountains. Forests (e.g., deciduous). Shaded ground; moderately moist soil. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: June-July, September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. Reference. Freitag, 1969 (ecology). Cyclotrachelus (Evarthrus) furtivus (LeConte, 1852) Ecology. Lowlands and mountains. Deciduous forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under debris. Biology. Seasonality: April-July, October. Parasites: mites. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Freitag, 1969 (ecology). Cyclotrachelus (Evarthrus) gigas (Casey, 1918) Ecology. Lowlands. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Cyclotrachelus (Evarthrus) gravesi (Freitag, 1969) Ecology. Lowlands. Biology. Seasonality: March. Dispersal power. Brachypterous, incapable of flight. Cyclotrachelus (Evarthrus) gravidus (Haldeman, 1853) Ecology. Lowlands. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Cyclotrachelus (Cyclotrachelus) hernandensis (Van Dyke, 1943) Ecology. Lowlands. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under debris. Biology. Seasonality: January. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping.

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Cyclotrachelus (Evarthrus) heros (Say, 1823) Ecology. Lowlands. Cultivated fields (e.g., cotton). Open ground; dry soil. Biology. Seasonality: March. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Freitag, 1969 (ecology). Cyclotrachelus (Evarthrus) hypherpiformis (Freitag, 1969) Ecology. Lowlands. Biology. Seasonality: April, June. Dispersal power. Brachypterous, incapable of flight. Cyclotrachelus (Evarthrus) incisus (LeConte, 1848) Ecology. Lowlands and uplands. Deciduous forests (e.g., Quercus, Carya), cultivated fields (e.g., corn), pastures, and wastelands. Shaded or open ground; moderately moist soil. Nocturnal; sheltering during the day in leaf litter and under logs. Biology. Seasonality: March-June, August-November. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Freitag, 1969 (ecology); Kirk, 1975c (ecology); Allen and Thompson, 1977 (ecology); Davidson, 1995 (ecology). Cyclotrachelus (Evarthrus) iowensis (Freitag, 1969) Ecology. Uplands. Prairies, cultivated fields (e.g., corn), and open forests. Open ground; moderately moist soil. Nocturnal; sheltering during the day under stones and logs. Biology. Seasonality: May-October. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones and logs. References. Freitag, 1969 (ecology); Kirk, 1975c (ecology, dispersal power). Cyclotrachelus (Cyclotrachelus) iuvenis (Freitag, 1969) Ecology. Mountains. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: April-October. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. Reference. Freitag, 1969 (ecology). Cyclotrachelus (Cyclotrachelus) laevipennis (LeConte, 1848) Ecology. Lowlands and mountains. Deciduous forests, coniferous forests (e.g., Pinus), and sheltered pastures. Shaded ground; moderately moist soil covered with thick leaf litter. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: JanuaryDecember. Tenerals: January, October (FL). Adults found overwintering (SC). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Freitag, 1969 (ecology); Kirk, 1970 (biology); Harris and Whitcomb, 1971 and 1974 (ecology, biology).

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Cyclotrachelus (Cyclotrachelus) levifaber (Freitag, 1969) Ecology. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under logs. Biology. Seasonality: May, October-December. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning logs. Cyclotrachelus (Cyclotrachelus) macrovulum (Freitag, 1969) Ecology. Lowlands. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under debris. Biology. Seasonality: November-December. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Cyclotrachelus (Evarthrus) nonnitens (LeConte, 1873) Ecology. Lowlands. Deciduous forests; coniferous forests (e.g., Pinus). Shaded ground; moderately moist soil. Biology. Seasonality: February, May. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Allen and Thompson, 1977 (ecology). Cyclotrachelus (Cyclotrachelus) ovulum (Chaudoir, 1868) Ecology. Lowlands. Coniferous forests (e.g., Pinus). Shaded ground; moderately moist soil covered with thick leaf litter. Nocturnal; sheltering during the day in leaf litter and under bark. Biology. Seasonality: January-May, October-December. Tenerals: OctoberMarch (FL). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Freitag, 1969 (ecology); Harris and Whitcomb, 1971 and 1974 (ecology, biology). Cyclotrachelus (Cyclotrachelus) parafaber (Freitag, 1969) Ecology. Lowlands. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under debris. Biology. Seasonality: November. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Cyclotrachelus (Evarthrus) parasodalis (Freitag, 1969) Ecology. Lowlands. Deciduous forests (e.g., Quercus, Carya); coniferous forests (e.g., Pinus). Shaded ground; moderately moist soil. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: March-November. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Freitag, 1969 (ecology); Allen and Thompson, 1977 (ecology, biology).

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Cyclotrachelus (Evarthrus) sallei (LeConte, 1873) Ecology. Unknown. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Cyclotrachelus (Evarthrus) seximpressus (LeConte, 1848) Ecology. Lowlands and uplands. Cultivated fields (e.g., cotton, corn, clover), meadows, pastures, wastelands, roadsides, and flower beds. Deciduous (e.g., Quercus, Carya) and coniferous forests. Open (mostly) or shaded ground; moderately moist soil covered with some vegetation. Nocturnal; sheltering during the day under logs, stones, and in the soil. Biology. Seasonality: February-March. Adult and larval food, in captivity: mealworms and meat. Larval food: earthworms (in the field) and mealworms (in captivity). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning logs and stones. References. Blatchley, 1910 (ecology); Freitag, 1969 (ecology); Allen and Thompson, 1977 (ecology, biology). Cyclotrachelus (Evarthrus) sigillatus (Say, 1823) Ecology. Lowlands and uplands. Deciduous forests (mostly; e.g., Ulmus, Acer, Castanea, Fagus, Quercus, Magnolia, Carya), mixed forests (e.g., Quercus, Carya, Pinus), coniferous forests (e.g., Pinus), pastures, grasslands, low prairies, and cultivated fields (e.g., tobacco, corn). Shaded (mostly) or open ground; moderately moist soil covered with thick leaf litter or grass. Nocturnal; sheltering during the day in leaf litter (mostly), under stones, logs, pieces of wood, and other debris. Biology. Seasonality: January-December. Tenerals: spring (FL). Predators: woodpeckers and starlings. Parasites: mites. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter; light trapping. References. Freitag, 1969 (ecology); Harris and Whitcomb, 1971 and 1974 (ecology, biology); Erwin, 1981 (natural history); Anderson et al. 1995 (ecology, biology). Cyclotrachelus (Evarthrus) sinus (Freitag, 1969) Ecology. Lowlands. Mixed forests (Pinus, Quercus). Shaded ground; moderately moist soil. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Freitag, 1969 (ecology). Cyclotrachelus (Evarthrus) sodalis colossus (LeConte, 1848) Ecology. Lowlands. Cultivated fields (e.g., corn, alfalfa, sweet clover), pastures, meadows, fencerows, roadsides, and farm yards. Open ground; dry soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day under pieces of wood. Biology. Seasonality: June-August. Adult food, in the field: lepidopterous caterpillars

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(mostly), scarabaeid larvae, ants, fungi, and algae. Predators: trashers, thrushes, and foxes. Parasites: parantennulid mites. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; lifting pieces of wood. References. Freitag, 1969 (ecology, biology). Cyclotrachelus (Evarthrus) sodalis lodingi (Van Dyke, 1926) Ecology. Lowlands. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: May-July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. Cyclotrachelus (Evarthrus) sodalis sodalis (LeConte, 1848) Ecology. Lowlands and uplands. Cultivated fields (e.g., cotton, cabbage, corn, alfalfa, asparagus, bean, tomato), pastures, old fields, roadsides, lawns, borders between fields and forests, open forests, and railway embankments. Open ground; dry, sandy or clayish soil covered with some vegetation. Nocturnal; sheltering during the day under pieces of wood. Biology. Seasonality: April-October. Adult food: lepidopterous caterpillars, curculionids, lygaeids, flies, and ants (in the field); lepidopterous caterpillars, grasshopper nymphs, pentatomids, curculionids, hymenopteran larvae, weed and corn seeds, and cheese (in captivity). Adults are cannibalistic in captivity. Economic importance: may play an important role in the control of alfalfa insect pests. Predators: crows and robins. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on plants). Strongly favored by human activities. Collecting techniques. Pitfall trapping; lifting pieces of wood; night collecting with a headlamp; sweeping plants. References. Freitag, 1969 (ecology); Barney and Armbrust, 1980 (biology); Barney and Pass, 1986a and 1986b (biology); Perfecto et al. 1986 (ecology, biology); Epstein and Kulman, 1990 (ecology, biology); Will et al. 1995 (dispersal power). Cyclotrachelus (Cyclotrachelus) spoliatus (Newman, 1838) Ecology. Lowlands. Deciduous forests; coniferous forests (e.g., Pinus). Shaded ground; moderately moist soil. Nocturnal; sheltering during the day in leaf litter, under logs and pieces of wood. Biology. Seasonality: March-November. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter; light trapping. References. Freitag, 1969 (ecology); Kirk, 1969 (ecology). Cyclotrachelus (Evarthrus) substriatus (LeConte, 1848) Ecology. Uplands. Cultivated fields (e.g., corn), pastures, meadows, grasslands (shortgrass), semi-deserts, and open forests (e.g., Quercus, Pinus). Open ground; dry soil covered with vegetal debris and grass. Nocturnal; sheltering during the day under stones and dry cattle dung. Biology. Seasonality: May-October. Adult food, in captivity: grasshopper nymphs, boiled egg yolks, and wheat. Defense-mechanism: when alarmed, the

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adult may escape by burrowing under vegetal debris. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones. References. Freitag, 1969 (ecology); Kirk, 1975c (ecology); Hsin et al. 1979 (biology); Chen and Willson, 1996 (biology); McIntyre, 1998 (ecology, biology). Cyclotrachelus (Cyclotrachelus) texensis (Freitag, 1969) Ecology. Lowlands. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under debris. Biology. Seasonality: December. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Cyclotrachelus (Evarthrus) torvus deceptus (Casey, 1918) Ecology. Lowlands. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Cyclotrachelus (Evarthrus) torvus torvus (LeConte, 1863) Ecology. Lowlands and uplands. Cultivated fields (e.g., corn, alfalfa), pastures, wastelands, roadsides, farmyards; upper zone of streams; open forests. Open ground; dry soil covered with sparse vegetation (e.g., grass). Nocturnal; sheltering during the day under logs, boards, and stones. Biology. Seasonality: February-December. Parasites: tapeworms. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Favored by human activities. Collecting techniques. Pitfall trapping; turning debris. References. Freitag, 1969 (ecology); Kirk, 1975c (ecology). Cyclotrachelus (Cyclotrachelus) unicolor (Say, 1823) Ecology. Lowlands. Open forests (e.g., Quercus, Pinus), grasslands, and pastures. Shaded or open ground; moderately moist soil. Nocturnal; sheltering during the day under debris. Biology. Seasonality: January-April, July-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Cyclotrachelus (Cyclotrachelus) vinctus (LeConte, 1852) Ecology. Mountains. Forests (mostly), pastures, and grasslands. Shaded ground; moderately moist soil covered with thick leaf litter. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: May-June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter; baiting traps with carrion. Reference. Freitag, 1969 (ecology). Cyclotrachelus (Evarthrus) whitcombi (Freitag, 1969) Ecology. Lowlands and mountains. Biology. Seasonality: October. Dispersal power. Brachypterous, incapable of flight. Moderate runner.

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Cylindronotum aeneum Putzeys, 1845 Ecology. On vine foliage. Biology. Seasonality: July. Dispersal power. Wing-condition unknown. Climber. Cymindis (Pinacodera) abbreviatus (Casey, 1920) Natural History. Unknown. Cymindis (Cymindis) americanus Dejean, 1826 Ecology. Lowlands, mountains, and subalpine zone. Grasslands, old fields, pastures, meadows, cultivated fields (e.g., corn), fencerows, hedges, sand pits, gravel pits, hills, orchards, thickets, forest edges and clearings, and open deciduous (mostly) or coniferous forests. Open or shaded ground; dry, sandy or clayish soil covered with sparse vegetation. Nocturnal; sheltering during the day under well-embedded stones (mostly), dead leaves, and pieces of wood. Gregarious (in winter). Biology. Seasonality: February-December. Tenerals: late July (QU). Adults found overwintering at the edge of woods (mostly), in pastures, on hills and hillocks, in clearings and in woods; under stones (mostly), in the soil, under logs, in rotten pieces of wood, and in ant nests. Adult food, in captivity: meat. Predators: toads. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strong burrower. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning well-embedded stones. References. Blatchley, 1910 (biology); Darlington, 1936 (dispersal power); Lindroth, 1969a (natural history); Larochelle, 1974a (ecology); Kirk, 1975c (dispersal power); Larochelle, 1977e (dispersal power); Epstein and Kulman, 1990 (ecology, biology); Anderson et al. 1995 (ecology). Cymindis (Pinacodera) ampliatus (Casey, 1920) Ecology. Uplands. Biology and Dispersal power. Unknown. Cymindis (Cymindis) arizonensis Schaeffer, 1910 Ecology. Mountains.Vacant lots and roadsides. Open ground; dry, gravelly soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day under stones. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones. Cymindis (Pinacodera) atripennis (Casey, 1920) Ecology. Lowlands. Biology. Seasonality: February-March, June, December. Adults found overwintering in moss. Dispersal power. Unknown.

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Cymindis (Pinacodera) blandus Casey, 1913 Natural History. Unknown. Cymindis (Cymindis) borealis LeConte, 1863 Ecology. Lowlands, mountains, subalpine and alpine zones. Abandoned fields, meadows, pastures, grasslands, cultivated fields (e.g., wheat), hills, roadsides, sand pits, gravel pits, and dry arctic tundra. Open ground; dry sandy soil covered with short sparse vegetation (e.g., grass). Nocturnal; sheltering during the day under stones, dead leaves, and pieces of cardboard. Biology. Seasonality: April-November. Gravids: late July (QU). Tenerals: late June, July. Adults found overwintering in pastures and on hills; under plant rosettes (Verbascum). Adult food, in captivity: mealworms and meat. Predators: toads. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pitfall trapping; turning stones; raking the leaf litter. References. Darlington, 1936 (dispersal power); Lindroth, 1963a and 1969a (natural history); Larochelle, 1974a (ecology) and 1977e (dispersal power). Cymindis (Cymindis) californicus G.H. Horn, 1895 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Cymindis (Pinacodera) complanatus Dejean, 1826 Ecology. Lowlands. Biology. Seasonality: February-May. Dispersal power. Macropterous, probably capable of flight. Cymindis (Cymindis) cribricollis Dejean, 1831 Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, meadows, cultivated fields and their borders (e.g., flax, wheat, barley), vacant lots, roadsides, sand pits, gravel pits, hedges, coastal tundra, thickets (Alnus); forest edges, paths, roadsides, and clearings; light forests. Open (mostly) or slightly shaded ground; dry sandy soil covered with sparse vegetation (e.g., grass) or dead leaves. Nocturnal; sheltering during the day under stones and dead leaves (mostly), logs, boards, pieces of paper, heaps of hay, and fallen pieces of bark. Gregarious (in winter). Biology. Seasonality: March-November. Gravids: early July to early August (AB, QU). Tenerals: late June to August. Adults found overwintering on higher and drier ground, in fields neighboring woods, in clearings, at the edge of woods, and in woods; under stones, pieces of wood, clumps of grass, in the soil (as deep as 25 cm), in rotten logs, under logs, and under the loose bark of fallen trees. Adult food: lepidopterous caterpillars (in the field); mealworms, meat, and banana skin (in captivity). Larval food, in captivity: mealworms. Predators: toads (regularly) and birds of prey. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting

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techniques. Pitfall trapping; turning stones; raking dead leaves. References. Darlington, 1936 (dispersal power); Lindroth, 1955a, 1963a, and 1969a (natural history); Schmoller, 1970 (biology); Frank, 1971b (biology); Larochelle, 1972e (dispersal power); Kirk, 1975c (ecology); Levesque et al. 1979 (biology); Reeves et al. 1983 (biology); Levesque and Levesque, 1986 (biology); Parmenter and MacMahon, 1988 (biology). Cymindis (Cymindis) elegans elegans LeConte, 1848 Ecology. Lowlands. Vacant lots and sand hills. Open ground; dry, sandy, almost bare soil. Nocturnal; sheltering during the day under stones and vegetal debris. Biology. Seasonality: March-June, August-December. Tenerals: April-June, December (NC). Adults found overwintering in vacant lots; under vegetal debris. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; turning debris. Reference. Darlington, 1936 (dispersal power). Cymindis (Cymindis) elegans mobilensis Casey, 1920 Ecology. Lowlands. Macrohabitat unknown. Nocturnal; sheltering during ther day under logs. Biology. Seasonality: February-November. Dispersal power. Wing-condition unknown. Cymindis (Cymindis) evanescens Casey, 1913 Ecology. Mountains. Shrub-steppes and pine forests. Open ground; dry soil, sometimes rocky, with some vegetation (e.g., Artemisia, Chrysothamnus, Bromus). Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Cymindis (Cymindis) interior Lindroth, 1969 Ecology. Uplands. Biology. Seasonality: May-September, November. Tenerals: August (KS). Dispersal power. Brachypterous, incapable of flight. Reference. Lindroth, 1969a (biology, dispersal power). Cymindis (Cymindis) laticollis Say, 1830 Ecology. Lowlands and mountains. A flood-plain forest (IL). Edge of a pond (SD). Sheltering under logs. Biology. Seasonality: June-September. Dispersal power. Wing-dimorphic: brachypterous (incapable of flight) or macropterous (probably capable of flight). Cymindis (Pinacodera) limbatus Dejean, 1831 Ecology. Lowlands and mountains. Deciduous forests (e.g., Quercus, Carya, Castanea), mixed forests (Pinus, Quercus, Carya), coniferous forests (e.g., Pinus), and orchards (e.g., apple,

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peach). Shaded ground; moderately moist soil covered with thick leaf litter. Mostly crepuscular and nocturnal; sheltering during the day under the loose bark of standing and fallen trees, on trees, in leaf litter, under stones, and in squirrel nests. The larva is grounddwelling. Biology. Seasonality: February-December. Tenerals: July (MD, WV). Adults found overwintering in forests; in and under wood debris. Adult food, in the field: lepidopterous caterpillars. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Regular in lake- and seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (on trees; up to 7 m above ground). Collecting techniques. Lifting the loose bark of trees; beating trees; using sticky boards; pyrethrum spraying; using malaise traps; light trapping; sugaring trees; turning drift material; baiting traps with molasses and fermented sugar; baiting traps with carrion. References. Blatchley, 1910 (ecology); Kirk, 1969 (biology); Lindroth, 1969a (dispersal power); Liebherr and Mahar, 1979 (natural history); Erwin, 1981 (natural history); Krinsky and Godwin, 1990 (ecology). Cymindis (Cymindis) neglectus Haldeman, 1843 Ecology. Lowlands and mountains. Light forests (mostly deciduous, e.g., Acer, Quercus; also mixed), their clearings and edges, thickets (e.g., Alnus, Juniper), adjacent moraines, hills, vacant lots, and roadsides. Ground shaded by trees, shrubs, and bushes; moderately or very dry, sandy soil covered with thick leaf litter. Nocturnal; sheltering during the day mostly in leaf litter and under reindeer moss (Cladonia), also under stones. Gregarious (in spring). Biology. Seasonality: February-November. Gravids: early July (QU). Tenerals: early July to mid-August (QU, ME). Adults found overwintering in woods, thickets, pastures, on hills, and in vacant lots; in leaf litter and under stones. Adult food, in captivity: meat and apple. Predators: toads. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Raking the leaf litter; turning reindeer moss; pitfall trapping. References. Darlington, 1936 (dispersal power); Larochelle, 1971c (ecology) and 1972e (dispersal power). Cymindis (Pinacodera) obscurus (Casey, 1920) Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Cymindis (Cymindis) pilosus Say, 1823 Ecology. Lowlands and mountains. Grasslands, meadows, prairies, pastures, rangelands, vacant lots, cultivated fields (e.g., hay, barley, corn, alfalfa), hills, roadsides, sand pits, gravel pits, orchards (e.g., peach), and light forests (deciduous, coniferous). Open ground; well-drained, dry, sandy, rarely gravelly soil covered with sparse vegetation. Nocturnal; sheltering during the day under stones and dead leaves (mostly), heaps of grass, dry cow dung, plant rosettes (Verbascum), and boards. Sometimes gregarious. Biology. Seasonality: January-December. Gravids: early June (once) and early August (QU). Tenerals: late May to late July. Adults found overwintering on hills, in pastures, vacant lots,

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and gravel pits; under stones and plant rosettes (Verbascum). Predators: toads, frogs, birds of prey, crows, and starlings. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; raking the leaf litter. References. Blatchley, 1910 (ecology); Darlington, 1936 (dispersal power); Lindroth, 1969a (natural history); Larochelle, 1974a (ecology); Kirk, 1975c (ecology, dispersal power). Cymindis (Cymindis) planipennis LeConte, 1863 Ecology. Lowlands and mountains. Grasslands, pastures, prairies, rangelands, cultivated fields and their borders (e.g., wheat, barley), hills, and gravel pits. Open ground; dry, sandy soil covered with sparse vegetation (e.g., grass). Nocturnal; sheltering during the day under stones, dry cow dung, about roots of plants, and under other debris. Gregarious (up to 9 individuals under a single cow dung cake). Biology. Seasonality: MarchOctober. Tenerals: June (NT), early July (MB), and late August (QU). Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; turning stones. References. Lindroth, 1969a and 1971a (natural history); Bell, 1971 (ecology); Kirk, 1975c (ecology, dispersal power); Lavigne, 1978 (ecology); Krinsky and Godwin, 1990 (ecology). Cymindis (Pinacodera) platicollis (Say, 1823) Ecology. Lowlands and mountains. Decidous forests (mostly; e.g., Quercus, Acer), mixed forests (Pinus, Quercus, Carya), coniferous forests (e.g., Pinus), and orchards (e.g., peach). Shaded ground; moderately moist soil covered with thick leaf litter. Mostly crepuscular and nocturnal; sheltering during the day under the loose bark of tree-stumps and standing trees, on trees and bushes, in bunches of Spanish Moss, in leaf litter, in moss, and under stones. The larva is ground-dwelling. Biology: Seasonality: January-December. Adult found overwintering in woods; under loose bark and in leaf litter. Adult food, in the field and in captivity: lepidopterous caterpillars. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (on tree-stumps, trees, and bushes). Collecting techniques. Lifting the loose bark of trees; beating trees (including Spanish Moss); using sticky boards; pyrethrum spraying; using tree bands; light trapping; using malaise traps; baiting traps with molasses and fermented sugar. References. Blatchley, 1910 (ecology); Leng, 1915 (ecology); Löding, 1945 (biology); Lindroth, 1969a (natural history); Liebherr and Mahar, 1979 (natural history). Cymindis (Pinacodera) punctifer (LeConte, 1884) Ecology. A mixed forest (Quercus-Pinus; Mexico). Biology and Dispersal power. Unknown.

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Cymindis (Pinacodera) punctiger LeConte, 1851 Ecology. Lowlands and mountains. Mexico: semi-deserts and open forests (Quercus, Pinus). Nocturnal; sheltering during the day in leaf litter, under stones and logs, or active on yucca plants. Biology. Seasonality: January-March, July, November-December. Dispersal power. Wing-condition unknown. Moderate runner. Occasional climber (on plants). Collecting techniques. Pitfall trapping; turning stones and logs. Cymindis (Cymindis) seriatus Hatch, 1953 Ecology. Mountains. Biology. Seasonality: September. Dispersal power. Brachypterous, incapable of flight. Cymindis (Pinacodera) subcarinatus (Casey, 1920) Natural History. Unknown. Cymindis (Cymindis) unicolor Kirby, 1837 Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, meadows, and vicinity of snowfields. Open ground; dry, gravelly soil covered with sparse vegetation. Nocturnal; sheltering during the day under stones. Biology. Seasonality: June-September. Gravids: early July (QU). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones. References. Lindroth, 1955a, 1963a, and 1969a (natural history). Cymindis (Cymindis) uniseriatus Bates, 1884 Ecology. Mountains. Mexico: open forests (Quercus, Pinus). Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Cymindis (Cymindis) vaporariorum (Linné, 1758) Ecology. Lowlands and mountains. Tundra, in Salix leaf litter. Europe: moraines, heaths, and light forests (e.g., Pinus); open (mostly) or shaded ground; dry, sandy or gravelly soil, rarely peaty, with sparse vegetation. Biology. Unknown. Dispersal power. Wingdimorphic: brachypterous (incapable of flight) or macropterous. Occasional flier (Europe). Moderate runner. Collecting techniques. Pitfall trapping; baiting traps with dung; ° 1986a sifting leaf litter. References. Lindroth, 1945 and 1986 (natural history); H urka, (biology); Luff, 1993 (biology); Shpeley and Ball, 1999 (taxonomy, natural history). Darlingtonea kentuckensis kentuckensis Valentine, 1952 Ecology. Caves: along stream banks, flowstones, stalagmites, benches and mud flats shelves raised above the ground. Moist, clayish or muddy soil. Biology. Seasonality: January-

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March, June, August-November. Adult food, in the field: worms, cave-cricket eggs, and dipterous larvae. Adults are cannibalistic in captivity. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. Good climber. Darlingtonea kentuckensis lexingtoni Valentine, 1952 Ecology. Caves. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Fast runner. Dercylinus impressus (LeConte, 1853) Ecology. Lowlands. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Reference. Bousquet, 1996a (taxonomy). Diacheila arctica amoena (Faldermann, 1835) Ecology. Borders of marshes with icy standing water. Open ground; wet soil covered with moss carpets. Nocturnal; sheltering during the day in moss. Biology. Seasonality: June, August-September. Tenerals: September (AK). Defense-mechanism: when alarmed, the adult may stridulate. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Treading moss carpets into the water. References. Lindroth, 1954 and 1961 (natural history). Diacheila polita (Faldermann, 1835) Ecology. Open tundra with peaty soil (mostly). Pool margins with soft ground covered with Carex. Drier habitats covered with Betula nana, Vaccinium, Rubus, Carex, and moss. Rather dry soil. Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: June, August-September. Tenerals: August (NT). Adult and larval food, in captivity: mealworms. Defense-mechanism: when alarmed, the adult may stridulate. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. References. Lindroth, 1945, 1954, 1961, and 1985 (natural history); Goulet, 1983 (ecology, dispersal power). Dicaelus (Dicaelus) alternans alternans Dejean, 1826 Ecology. Lowlands. A roadside, under a piece of wood (FL). Biology. Seasonality: January-April, June-August, November. Dispersal power. Brachypterous, incapable of flight. Reference. Leng, 1915 (ecology). Dicaelus (Dicaelus) alternans subtropicus Casey, 1913 Ecology. Lowlands. Macrohabitat unknown. Sheltering under a piece of wood. Biology. Seasonality: February, May, September, December. Dispersal power. Brachypterous, incapable of flight. Moderate runner.

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Dicaelus (Paradicaelus) ambiguus LaFerté-Sénectère, 1841 Ecology. Forests (mostly; deciduous, mixed); cultivated fields (e.g., cotton). Moderately moist soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: March-December. Tenerals: September (MD). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Erwin, 1981 (natural history). Dicaelus (Liodicaelus) chermocki Ball, 1959 Ecology. Mountains. A coniferous forest (Abies, Pinus). A stream bank (in a canyon). Nocturnal; sheltering during the day under stones. Biology. Seasonality: March, JuneJuly, September. Dispersal power. Brachypterous, incapable of flight. Dicaelus (Dicaelus) costatus LeConte, 1853 Ecology. Lowlands. A scrub-oak area Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Dicaelus (Dicaelus) crenatus LeConte, 1853 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Dicaelus (Paradicaelus) dilatatus dilatatus Say, 1823 Ecology. Lowlands and mountains. Deciduous forests (mostly; e.g., Acer, Quercus), mixed forests (Pinus, Quercus), forest edges, open fields, grasslands, pastures, and cultivated fields (e.g., corn). Shaded (mostly) or open ground; moderately moist soil covered with thick leaf litter. Nocturnal; sheltering during the day under logs and stones. Larval habitat: woods and edges of woods, under pieces of wood and stones. Biology. Seasonality: March-November. Tenerals: August-September (NY, MD). Adults found overwintering in woods and at the edge of woods; under stones and in the soil (as deep as 10 cm). Adult food: snails, Gypsy Moth immature stages, and scarabaeid larvae (in the field); flies and lepidopterous pupae (in captivity). Defense-mechanism: when alarmed, the adult may emit a Brachinus-like smoke from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; baiting traps with malt; turning logs and stones; light trapping. References. Schaupp, 1878a, 1879a, and 1879b (biology); Liebeck, 1887 (biology); Dimmock and Knab, 1904 (biology); McCullough, 1967 (biology); Erwin, 1981 (natural history); Anderson et al. 1995 (ecology). Dicaelus (Paradicaelus) dilatatus sinuatus Ball, 1959 Ecology. Lowlands and mountains. Deciduous forests. Shaded ground; moderately moist soil. Biology. Seasonality: April-October. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping.

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Dicaelus (Paradicaelus) elongatus Bonelli, 1813 Ecology. Lowlands and mountains. Deciduous forests (mostly), hillsides, swamps, and old fields. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under stones (mostly), logs, pieces of wood, and among dead leaves. Gregarious (in fall and winter; up to 24 individuals under a single stone). Larval habitat: woods, under stones. Biology. Seasonality: January-November. Tenerals: early June to November. Adults found overwintering in colonies with Galerita and Chlaenius in woods; under stones and logs. Adult food, in the field: lepidopterous caterpillars and undetermined animal matter. Defense-mechanism: when alarmed, the adult may emit a powerful discharge of gas from the pygidial glands, which turns to a dense smoke, at a range of 60 cm. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on plants). Collecting techniques. Pitfall trapping; turning stones; sweeping plants. References. Schaupp, 1878b, 1879a, and 1879b (biology); Dimmock and Knab, 1904 (biology); Sherman, 1928 (biology); Fattig, 1929 and 1933 (biology); Lindroth, 1969a (natural history); Kirk, 1970 (biology) and 1975c (ecology); Erwin, 1981 (natural history); Anderson et al. 1995 (ecology). Dicaelus (Paradicaelus) furvus carinatus Dejean, 1831 Ecology. Lowlands. Flood-plain forests: along river banks. Shaded ground; moist soil. Nocturnal; sheltering during the day under debris. Biology. Seasonality: April, JuneOctober. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Leng, 1915 (ecology). Dicaelus (Paradicaelus) furvus furvus Dejean, 1826 Ecology. Lowlands and mountains. Deciduous forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under stones and logs. Gregarious (in winter). Biology. Seasonality: March-December. Adults found overwintering in colonies with Galerita and Chlaenius in woods; under stones. Predators: crows. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones and logs; light trapping. Reference. Erwin, 1981 (natural history). Dicaelus (Liodicaelus) laevipennis laevipennis LeConte, 1848 Ecology. Mountains. Fields, deserts, and rocky slopes. Open ground; rather moist soil. Nocturnal; sheltering during the day under debris. Biology. Seasonality: April-September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Ball, 1959 (ecology). Dicaelus (Paradicaelus) politus Dejean, 1826 Ecology. Lowlands and mountains. Deciduous forests (mostly; e.g., Acer, Quercus) and mixed forests (e.g., Pinus, Tsuga, Fagus, Quercus, Carya). Shaded ground; rather

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moist soil covered with thick leaf litter. Nocturnal; sheltering during the day under stones, in wet rotten logs and tree-stumps (rich in large land slugs), under the loose bark of fallen trees, under logs, pieces of wood, and in thick leaf litter (at the base of tree-stumps). Gregarious. Biology. Seasonality: March-October. Gravids: mid-May (IL). Tenerals: August-September. Adults found overwintering in woods and at the edge of woods; under stones and in leaf litter. Adult food: lepidopterous immature stages (in the field); mealworms and meat (in captivity). Larval food, in captivity: mealworms. Predators: toads. Defense-mechanism: when alarmed, the adult may emit a Brachinus-like smoke from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Frequent climber (on logs and tree-stumps). Collecting techniques. Pitfall trapping; turning stones and logs. References. Schaupp, 1878b, 1879a, and 1879b (biology); Gilbert, 1957 (biology); Ball, 1959 (ecology); Lindroth, 1969a (natural history); Erwin, 1981 (natural history); Anderson et al. 1995 (ecology). Dicaelus (Dicaelus) purpuratus darlingtoni Fall, 1932 Ecology. Lowlands. Open forests. Shaded ground; rather dry soil. Nocturnal; sheltering during the day under dead fallen palm fronds. Biology. Seasonality: March-April, June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Dicaelus (Dicaelus) purpuratus purpuratus Bonelli, 1813 Ecology. Lowlands and mountains. Deciduous forests (mostly; e.g., Acer, Quercus, Ulmus), cultivated fields (e.g., cotton, corn), old fields, and pastures. Shaded (mostly) or open ground; moderately moist soil. Nocturnal; sheltering during the day under the loose bark of fallen trees, under logs and stones. Larval habitat: woods, in rotten logs. Biology. Seasonality: February-November. Tenerals: September (MI, NY). Adult food: snails and scarabaeid larvae (in the field); snails, earthworms, and pieces of slugs (in captivity). Adults do not attack live slugs. Predators: crows. Defensemechanism: when alarmed, the adult may emit a Brachinus-like smoke or a darkcolored defensive fluid from the pygidial glands. When alarmed, the larva may eject a dark-colored fluid from the anal tube. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; lifting the loose bark of trees. References. Schwarz, 1876 (ecology, biology); Dimmock and Knab, 1904 (ecology, biology); McCullough, 1969b (biology); Erwin, 1981 (natural history); Anderson et al. 1995 (ecology). Dicaelus (Dicaelus) purpuratus quadratus LeConte, 1848 Ecology. Lowlands. Border of a temporary pond, under an old boat (FL). Biology. Seasonality: January-July, September. Dispersal power. Brachypterous, incapable of flight. Moderate runner.

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Dicaelus (Dicaelus) purpuratus splendidus Say, 1823 Ecology. Lowlands and uplands. A cultivated field (wheat; SD). A damp site, under logs (OK). Biology. Seasonality: April-August, October. Larval food, in captivity: snails. Predators: raccoons. Defense-mechanism: when alarmed, the adult may emit a defensive fluid from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Kirk, 1975c (ecology). Dicaelus (Paradicaelus) sculptilis intricatus LeConte, 1873 Ecology. Lowlands. Low deciduous forests. Shaded ground; moist soil. Often found among myrtle (Vinca minor Linné). Nocturnal; sheltering during the day under logs and stones. Biology. Seasonality: March-April, July-October. Tenerals: late July (PA). Adult food, in the field: worms. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on tree trunks; up to 1 m above ground). Collecting techniques. Pitfall trapping; turning logs and stones; night collecting with a headlamp. Reference. Will et al. 1995 (ecology, dispersal power). Dicaelus (Paradicaelus) sculptilis sculptilis Say, 1823 Ecology. Lowlands and uplands. Relict prairies. Open ground; soil covered with some vegetation. Nocturnal; sheltering during the day under debris. Biology. Seasonality: June-August. Predators: foxes. Defense-mechanism: when alarmed, the adult may emit a repugnant Brachinus-like smoky gas from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. References. Blatchley, 1910 (ecology); Fattig, 1933 (biology); Erwin, 1981 (natural history). Dicaelus (Paradicaelus) sculptilis upioides Ball, 1959 Ecology. Lowlands and mountains. Deciduous forests, coniferous forests, wooded canyons, meadows, old fields, cultivated fields (e.g., corn, wheat, oat), gardens, and upper zone of stream banks. Shaded or open ground; moderately dry, clayish or sandy soil. Nocturnal; sheltering during the day under logs, stones, and dead leaves. Biology. Seasonality: April-September. Tenerals: September (MB). Adult food, in the field: snails. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning logs and stones. References. Lindroth, 1969a (biology); Kirk, 1975c (ecology; as sculptilis Say); Richardson and Holliday, 1982 (ecology); Epstein and Kulman, 1990 (ecology, biology); Holliday, 1991 (ecology, biology). Dicaelus (Liodicaelus) suffusus (Casey, 1913) Ecology. Mountains. Mexico: mixed forests (Quercus, Pinus) and meadows. Biology. Seasonality: July, October. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping.

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Dicaelus (Paradicaelus) teter Bonelli, 1813 Ecology. Lowlands and mountains. Deciduous (mostly; e.g., Acer, Quercus), mixed and coniferous (e.g., Abies, Picea) forests: often along gorges. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under stones (mostly), the loose bark of fallen trees, pieces of wood, and in rotten logs or loose piles of decaying bark at the base of dead trees; active at night on standing and fallen trees. Biology. Seasonality: March-October. Tenerals: late July (WV), September (MD). Adult food, in the field: snails and lepidopterous immature stages. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on trees and logs). Collecting techniques. Pitfall trapping; turning stones. References. Ball, 1959 (ecology); Barr, 1969 (ecology); Erwin, 1981 (natural history). Dicheirotrichus (Trichocellus) cognatus (Gyllenhal, 1827). Ecology. Lowlands, mountains, subalpine and alpine zones. Open or light forests and thickets; neighboring sand pits and gravel pits. Moraines, grasslands, meadows, cultivated fields (e.g., wheat, barley, cabbage), pastures, vacant lots, roadsides, and alkali flats. Slightly shaded or open ground; dry, usually sandy soil covered with sparse vegetation (e.g., tiny Polytrichum, grass). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under small stones, dead leaves, heaps of hay, in rotten grass, under pieces of wood or cardboard, dry cow dung, in the soil at the base of plants, and in mouse nests. Gregarious. Biology. Seasonality: January, March-November. Copulating pairs: October (AB). Gravids: late May (NY). Tenerals: July to late August; mostly after mid-July. Adults found overwintering on higher and drier ground, in the upper zone of sand pits and gravel pits, on hills and hillocks, in hawthorn hedges, and at the edge of woods; mostly in the soil under small well-embedded stones (as deep as 7.5 cm), also in leaf litter. Active at low temperatures under snow in October-November. Adult food: Calluna-seeds (in the field, Europe); lepidopterous eggs (in captivity). Dispersal power. Macropterous. Frequent flier (in the sunshine; to artificial lights at night; to artifical lights in captivity). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning small stones; sweeping plants; using window traps. References. Larsson, 1939 (biology). Lindroth, 1945, 1955a, 1963b, 1968, and 1986 (natural history); Frank, 1971b (biology); Boer, 1980 (biology); Kirk, 1975c (ecology, dispersal power). Koch, 1989 (ecology); Kavanaugh, 1992 (natural history); Kryzhanovskij et al. 1995 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Dicheirotrichus (Oreoxenus) mannerheimii (R.F. Sahlberg, 1844). Ecology. Lowlands and mountains. Moraines. Open ground; dry, sandy or gravelly soil covered with sparse low vegetation. Mostly nocturnal; usually sheltering during the day under stones and vegetal debris; sometimes active on cloudy days on plants. Biology. Seasonality: May to mid-August. Tenerals: mid-August (AK). Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Moderate runner. Occasional climber (on plants). Collecting techniques. Turning stones; sweep-

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ing plants. References. Lindroth, 1968 (natural history) and 1986 (ecology); Koch, 1989 (ecology); Kryzhanovskij et al. 1995 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Dicheirus brunneus (Dejean, 1829) Ecology. Lowlands and mountains. Grassy areas. Open ground. Nocturnal; sheltering during the day under debris. Biology. Seasonality: April. Dispersal power. Unknown. Reference. Noonan, 1968 (ecology). Dicheirus dilatatus angulatus Casey, 1914 Ecology. Lowlands. Biology and Dispersal power. Unknown. Dicheirus dilatatus dilatatus (Dejean, 1829) Ecology. Lowlands and mountains. Grasslands and hillside ravines. Open ground; dry soil. Nocturnal; sheltering during the day under stones and other debris. Biology. Seasonality: April. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones and other debris. Dicheirus obtusus LeConte, 1852 Ecology. Lowlands and mountains. An open grassy area (OR). Biology and Dispersal power. Unknown. Reference. Noonan, 1968 (ecology). Dicheirus piceus (Ménétriés, 1843) Ecology. Lowlands and mountains. Grasslands, vacant lots, rocky hillsides, and shorelines. Open ground; dry soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day under stones, logs, and pieces of wood. Larval habitat: burrows (about 10 cm deep). Biology. April-May, September. Tenerals: August-September (WA). Larval food, in the field: seeds. Predators: spiders. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning debris. References. Lindroth, 1968 (natural history); Noonan, 1968 (ecology); Alcock, 1976 (ecology, biology). Dicheirus strenuus (G.H. Horn, 1868) Ecology. Lowlands. Grassy areas. Open ground. Nocturnal; sheltering during the day under debris. Biology. Seasonality: September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Reference. Noonan, 1968 (ecology). Diplochaetus desertus Van Dyke, 1953, see Diplochaetus emaciatus (Bates, 1891) Reference. Bousquet and Laplante, 1997 (taxonomy).

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Diplochaetus emaciatus (Bates, 1891) (Diplochaetus desertus Van Dyke, 1953) Ecology. Coastal lowlands. Seashore. Open ground; moist, saline soil. Biology. Seasonality: March, June. Dispersal power. Macropterous, probably capable of flight. Reference. Bousquet and Laplante, 1997 (taxonomy). Diplochaetus lecontei (G.H. Horn, 1876), see Diplochaetus rutilus (Chevrolat, 1863). Reference. Bousquet and Laplante, 1997 (taxonomy). Diplochaetus megacephalus Bousquet, 1997 Ecology. Gypsum flats. Nocturnal; sheltering during the day under debris. Biology. Seasonality: August. Dispersal power. Macropterous, probably capable of flight. Reference. Bousquet and Laplante, 1997 (taxonomy, ecology). Diplochaetus planatus (G.H. Horn, 1876) Ecology. Lowlands. Sea beaches; shores of saline lakes and ponds; alkali flats. Open ground; moist, clayish or muddy, saline soil. Nocturnal; sheltering during the day in clay cracks, under sticks, and other debris. Biology. Seasonality: May, September-October. Adult food, in the field: flies. Dispersal power. Macropterous, probably capable of flight. Collecting techniques. Treading the soil with the feet; pouring water over the ground. Reference. Bousquet and Laplante, 1997 (taxonomy, ecology). Diplochaetus rutilus (Chevrolat, 1863) (Diplochaetus lecontei (G.H. Horn, 1876)) Ecology. Lowlands. Borders of small salt marshes, ponds, and lagoons. Open ground: dry or wet, muddy or clayish, saline soil. Nocturnal; sheltering during the day mostly under layers of dried thickened mud and in cracks in the soil, also under driftwood. Biology. Seasonality: January-August, October-November. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Collecting techniques. Treading the soil with the feet; pouring water over the ground. References. Smith, 1910 (ecology; as lecontei (G.H. Horn)); Bousquet and Laplante, 1997 (taxonomy, ecology). Diplocheila (Isorembus) assimilis (LeConte, 1844) Ecology. Lowlands. Margins of eutrophic marshes and irrigation canals; swamps, low forests, and flood-plain forests. Open or shaded ground; wet, muddy or clayish soil, rich in organic debris, covered with dense vegetation (e.g., Typha, Carex). Nocturnal; sheltering during the day in beds of floating plant debris (Typha, grass), under dead leaves, and pieces of wood. Biology. Seasonality: May-September. Gravids:

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early June to late July (ON, QU). Tenerals: August (NH). Defense-mechanism: when alarmed, the adult may escape by swimming. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Good swimmer. Collecting technique. Treading the dead floating vegetation into the water. Reference. Lindroth, 1968 (natural history). Diplocheila (Isorembus) crossi Will, 1998 Ecology. Lowlands. A hardwood swamp (MS). Nocturnal; sheltering during the day under logs and boards. Biology. Seasonality: April-May, July. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Reference. Will, 1998b (taxonomy, natural history). Diplocheila (Isorembus) impressicollis (Dejean, 1831) Ecology. Lowlands. Margins of eutrophic marshes and irrigation canals; swamps, marshy banks of rivers and brooks; margins of pools and ponds in forests; flood-plain forests. Open or shaded ground; muddy or clayish soil, rich in organic debris, covered with dense vegetation (e.g., Typha). Nocturnal; sheltering during the day in beds of floating plant debris (e.g., Typha), under logs, pieces of wood, and stones. Biology. Seasonality: February-October. Tenerals: early May (rarely), July-October; mostly after mid-July. Adults found overwintering on higher and drier ground, in the upper zone of sand pits; under logs. Adult food, in captivity: lepidopterous caterpillars and pupae. Defense-mechanism: when alarmed, the adult may escape by swimming. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Occasional climber (on trees). Good swimmer. Collecting techniques. Treading the dead floating vegetation into the water; light trapping. References. Lindroth, 1969a (natural history); Larochelle, 1974a (ecology); Kirk, 1975c (dispersal power); Will et al. 1995 (dispersal power). Diplocheila (Isorembus) major major (LeConte, 1848) Ecology. Lowlands. Margins of large eutrophic marshes and ponds (Typha); cypressswamps and low deciduous forests. Open or shaded ground; wet, muddy or clay soil, rich in organic debris, covered with dense vegetation (e.g., Typha). Nocturnal; sheltering during the day in beds of floating plant debris or under logs. Biology. Seasonality: March-October. Defense-mechanism: when alarmed, the adult may escape by swimming. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Good swimmer. Collecting technique. Treading the dead floating vegetation into the water. References. Leng, 1915 (dispersal power); Lindroth, 1969a (natural history); Kirk, 1975c (dispersal power); Will et al. 1995 (dispersal power).

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Diplocheila (Isorembus) major melissisa Ball, 1859 Ecology. Lowlands. Margins of eutrophic ponds; flood-plain forests and low deciduous forests. Open or shaded ground; wet, muddy or clayish soil covered with dense vegetation. Nocturnal; sheltering during the day under mats of floating plant debris. Biology. Seasonality: January-June, August, October. Adults found overwintering in woods; in rotten logs. Defense-mechanism: when alarmed, the adult may escape by diving into the water and swimming like a hydrophilid. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Good swimmer and diver. Collecting techniques. Treading mats of foating plant debris into the water; light trapping. Diplocheila (Isorembus) nupera Casey, 1897 Ecology. Unknown. Biology. Seasonality: January-July. Dispersal power. Macropterous. Occasional in seashore drift material, which indicates flight ability. Diplocheila (Isorembus) obtusa (LeConte, 1848) Ecology. Lowlands and mountains. Grasslands, pastures, cultivated fields (e.g., oat, asparagus, alfalfa), lawns, vacant lots, hills, roadsides, vicinity of sidewalks (even of cities), gravel pits, sand pits, and upper zone of lake shores and river banks. Open ground; firm, often sloping, well-drained, warm, dry, gravelly, sandy, stony, or chalky soil covered with sparse grass (e.g., Phleum). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under embedded stones (mostly), dead leaves, and pieces of wood. Semi-gregarious. Biology. Seasonality: March-November. Tenerals: July (rarely)-September. Adults found overwintering on higher and drier ground, at the edge of woods, in woods, in the upper zone of sand pits, gravel pits, and river banks, in pastures, on hills and hillocks; in the soil (as deep as 10 cm) and under well-embedded stones. Adult food, in captivity: earthworms, meat, corn kernels, and a raisin. Predators: frogs and toads. Dispersal power Macropterous. Occasional flier (to artificial lights at night). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Strong burrower. Occasional climber (on plants). Strongly favored by human activities and spring floods along large rivers. Collecting techniques. Turning embedded stones; pitfall trapping; sweeping plants. References. Lindroth, 1969a (natural history); Kirk, 1975c (ecology, dispersal power). Diplocheila (Isorembus) oregona (Hatch, 1951) Ecology. Lowlands and mountains. Borders of ponds and lakes. Open ground; moist, often alkaline soil. Nocturnal; sheltering during the day under dry cow dung. Biology. Seasonality: May-June, August. Adult food, in captivity: snails. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Lindroth, 1968 (natural history).

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Diplocheila (Isorembus) striatopunctata (LeConte, 1844) Ecology. Lowlands. Margins of eutrophic marshes, ponds, lakes, pools, slow irrigation canals, rivers, and brooks; swamps and flood-plain forests. Open or shaded ground; wet, muddy or clayish soil, sometimes moderately saline, rich in organic debris, covered with tall dense vegetation (e.g., Typha). Nocturnal; sheltering during the day under mats of dead floating plant debris (mostly), in leaf litter, under logs, and under pieces of wood. Biology. Seasonality: April-September. Gravids: late May to late August; mostly before early July (QU, VT). Tenerals: late June to September; mostly after mid-July. Adults found overwintering on higher and drier ground, at the edge of woods, in woods, and in the upper zone of sand dunes; under well-embedded stones, fallen pieces of bark, and in rotten logs. Predators: toads. Phoretic mites found on adult body. Defense-mechanism: when alarmed, the adult may escape by swimming. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Good burrower. Good swimmer. Collecting techniques. Treading the dead floating vegetation into the water; pitfall trapping; light trapping. References. Lindroth, 1968 (natural history); Kirk, 1975c (ecology, dispersal power); Will et al. 1995 (dispersal power). Diplocheila (Isorembus) undulata Carr, 1920 Ecology. Lowlands and uplands. Borders of a marsh and a pond. Tall-grass prairies. Open ground; moist, muddy or calcareous soil covered with dense vegetation (e.g., sedges, grasses, Sphagnum). Nocturnal. Biology. Seasonality: April-July. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Ball, 1992 (ecology). Diplous aterrimus (Dejean, 1828) Ecology. Lowlands, mountains, subalpine and alpine zones. Margins of cold, swift brooks and rivers. Close to water. Open (mostly) or half-shaded ground; well-drained, moist, bare soil consisting of coarse gravel and scattered stones. Adults respond to chemical cues associated with green algae. Nocturnal; sheltering during the day under stones (mostly), among gravel, and under logs. Larval exuvia and pupae found in a mud cell under a boulder lying on a flat rock in a stream bed (WY). Gregarious. Biology. Seasonality: April-August. Tenerals: late June to late August. Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Turning stones; raking the loose gravel. References. Emden, 1942 (biology); Lindroth, 1961 (natural history); Evans, 1983 (ecology); Kavanaugh, 1992 (natural history). Diplous californicus (Motschulsky, 1859) Ecology. Lowlands and mountains. Margins of cold, swift brooks and rivers. Close to water. Open ground; well-drained, moist, gravelly or sandy soil, bare (mostly) or cov-

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ered with sparse vegetation. Nocturnal; sheltering during the day under stones and among gravel. Gregarious (up to15 individuals under a single stone). Biology. Seasonality: April, June-August. Tenerals: July (ID, OR, WA). Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Turning stones; raking the loose gravel. Reference. Lindroth, 1961 (natural history). Diplous filicornis (Casey, 1918) Ecology. Lowlands, mountains, subalpine and alpine zones. Margins of cold, swift brooks and rivers. Close to water. Open or half-shaded ground; well-drained, moist, bare soil consisting of coarse gravel. Nocturnal; sheltering during the day among gravel and under stones. Biology. Seasonality: July. Tenerals: mid-July. Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Raking the loose gravel; turning stones. Reference. Lindroth, 1961 (natural history). Diplous rugicollis (Randall, 1838) Ecology. Lowlands, mountains, subalpine and alpine zones. Margins of cold, swift brooks and rivers. Close to water. Open (mostly) or half-shaded ground; well-drained, wet, gravelly, sandy or stony, bare soil. Nocturnal; sheltering during the day under stones and among gravel. Gregarious. Biology. Seasonality: April-September. Gravids: August (NH). Tenerals: late June to August; mostly after mid-July. Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Turning stones; raking the loose gravel. Reference. Lindroth, 1961 (natural history). Discoderus (Discoderus) aequalis Casey, 1914 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Discoderus (Discoderus) amoenus LeConte, 1863 Ecology. Uplands. Rangelands and deserts. Open ground; dry soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day under stones. Biology. Seasonality: July. Tenerals: early June (MS). Predators: asilids (Diptera). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones. References. Lavigne, 1977 (ecology); Ball and Bousquet, 2001 (taxonomy). Discoderus (Discoderus) congruens Casey, 1914 Ecology. Uplands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Reference. Ball and Bousquet, 2001 (taxonomy).

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Discoderus (Selenalius) cordicollis G.H. Horn, 1891 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Discoderus (Discoderus) crassicollis G.H. Horn, 1891 Ecology and Biology. Dispersal power. Macropterous, probably capable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Discoderus (Discoderus) dallasensis Casey, 1924 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Discoderus (Discoderus) impotens (LeConte, 1858) Ecology. Uplands. Biology. Seasonality: May-June. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Reference. Ball and Bousquet, 2001 (taxonomy). Discoderus (Discoderus) longicollis Casey, 1914 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Discoderus (Discoderus) obsidianus Casey, 1914 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Ball and Bousquet, 2001 (taxonomy). Discoderus (Discoderus) papagonis Casey, 1924 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Discoderus (Discoderus) parallelus (Haldeman, 1843) Ecology. Lowlands and mounains. Grasslands, relict prairies, rangelands, pastures, cultivated fields (e.g., tobacco, cotton, soybean, oat), roadsides, fencerows, and the upper zone of stream banks. Open ground; dry, sandy soil covered with some vegetation (e.g., Artemisia, grass). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in the soil, under stones, logs, and pieces of wood. Biology. Seasonality: January, March-November. Predators: toads. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in lake- and seashore drift material,

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which indicates flight ability. Moderate runner. Occasional climber (on plants, e.g., Lespedeza). Strongly favored by human activities. Collecting techniques. Pitfall trapping; light trapping; sweeping plants. References. Lindroth, 1968 (natural history); Kirk, 1969 (dispersal power), 1970 (ecology), and 1975c (ecology, dispersal power); Ball and Bousquet, 2001 (taxonomy). Discoderus (Selenalius) parilis (Casey, 1914) Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Discoderus (Discoderus) peregrinus Casey, 1924 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Ball and Bousquet, 2001 (taxonomy). Discoderus (Discoderus) pinguis Casey, 1884 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Discoderus (Discoderus) robustus piceus Casey, 1914 Natural History. Unknown. Reference. Ball and Bousquet, 2001 (taxonomy). Discoderus (Discoderus) robustus robustus G.H. Horn, 1883 Ecology. Unknown. Biology. Seasonality: April. Dispersal power. Macropterous, probably capable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Discoderus (Discoderus) subviolaceus Casey, 1914 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Discoderus (Discoderus) symbolicus Casey, 1914 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Discoderus (Discoderus) tenebrosus (LeConte, 1848) Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Reference. Ball and Bousquet, 2001 (taxonomy).

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Discoderus (Discoderus) texanus Casey, 1924 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Reference. Ball and Bousquet, 2001 (taxonomy). Dromius (Dromius) fenestratus (Fabricius, 1794) Ecology. Lowlands. Europe: forests (mostly Pinus); sheltering during the day under the loose bark of trees. Biology. Seasonality: April, October. Dispersal power. Macropterous. Flight records (Europe). Moderate runner. Frequent climber (on trees; Europe). References. Larsson, 1939 (biology); Lindroth, 1945 and 1986 (natural history); Koch, 1989 (ecology); Downie and Arnett, 1996 (first North American record); Larson, 1998 (biology). Dromius (Dromius) piceus Dejean, 1831 Ecology. Lowlands and mountains. Deciduous forests (e.g., Alnus, Acer, Populus, Quercus, Liriodendron), mixed forests, coniferous forests (e.g., Picea, Abies, Pinus, Larix), and orchards (e.g., apple). Shaded ground. Nocturnal; sheltering during the day under the loose bark and bark scales of standing trees; rarely active during the day on shrubs or flowering plants; active on tree trunks shortly after sunset and in the canopy during the night. Biology. Seasonality: January-December. Eggs are laid on tree trunks. Tenerals: July-September (ME, QU, VA). Adults found overwintering in woods, at the edge of woods; at the base of trees, under pieces of fallen tree bark and wood, in leaf litter, in rotten logs, and in moss. Adult food, in captivity: ant eggs and meat. Predators: toads. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (adult and larva; on trees, shrubs, and plants). Collecting techniques. Beating tree trunks with a mallet to make beetles fall on a ground sheet; using sticky boards; pyrethrum spraying; light trapping; night collecting with a headlamp; sugaring trees. References. Löding, 1945 (biology); Lindroth, 1969a (natural history); Kirk, 1970 (ecology, dispersal power) and 1975c (dispersal power); Erwin, 1981 (natural history); Mahar et al. 1983 (natural history); Krinsky and Godwin, 1990 (ecology); Dearborne and Donahue, 1993 (ecology); Will, 1998a (biology). Dyschiriodes (Dyschiriodes) abbreviatus (Putzeys, 1846) Ecology. Lowlands. Banks of water bodies. Moist soil. Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: February-May, August-October, December. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Slow runner. Strong burrower. Collecting techniques. Pouring water over the ground; using fruit fly traps. Reference. Fedorenko, 1996 (taxonomy).

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Dyschiriodes (Dyschiriodes) aeneolus (LeConte, 1850) (Dyschirius frigidus Mannerheim, 1853) Ecology. Lowlands. An open mossy marsh. A sand-mixed clay area at some distance from a river. Moist soil. Nocturnal; sheltering during the day in burrows dug in the soil. Associated with staphylinids (Bledius). Biology. Seasonality: April-October. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. References. Lindroth, 1961 (natural history; as frigidus); Fedorenko, 1996 (taxonomy). Dyschiriodes (Paradyschirius) affinis (Fall, 1921) Ecology. Lowlands and mountains. River banks; margins of ponds (in sand pits and gravel pits); shores of lakes and reservoirs. Open ground; steep, moist or wet, sandy bare soil, sometimes mixed with silt. Nocturnal; sheltering during the day in burrows dug in the soil. Gregarious. Associated with staphylinids (Bledius). Biology. Seasonality: April-September. Tenerals: August-September (QU, VT). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Lindroth, 1961 (natural history); Choate, 1977 (ecology); Erwin, 1981 (natural history); Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) alticola (Lindroth, 1961) Ecology. Mountains and subalpine zone. Margins of a pond. A thin mixed forest. A subalpine meadow. Ground open or shaded by bushes (e.g., Alnus); wet or moist soil consisting of mud or fine sand, covered with some vegetation. Nocturnal; sheltering during the day in burrows dug in the soil and under stones. Biology. Seasonality: MayAugust. Tenerals: early July (WY). Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. References. Lindroth, 1961 (natural history); Papp, 1978 (ecology, biology); Fedorenko, 1996 (taxonomy). Dyschiriodes (Paradyschirius) analis (LeConte, 1852) Ecology. River banks, pool margins, and mud flats. Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: April-August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) aratus (LeConte, 1852) Ecology. Lowlands and mountains. River banks, lake shores, and reservoir borders. Open ground; wet, sandy-muddy soil, bare or covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil. Associated with staphylinids (Bledius). Biology. Seasonality: June-July. Dispersal power. Macropterous, probably capa-

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ble of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Eudyschirius) brevispinus (LeConte, 1878) Ecology. Lowlands. Gravel pits. Open ground; moderately dry soil composed of sand or fine gravel, covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil or under stones. Biology. Seasonality: May-August. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) campicola (Lindroth, 1961) Ecology. Banks of rivers (mostly) and brooks, lake shores, salt flats; borders of salt marshes and roadside ditches. Open ground; moist, sandy soil, sometimes silt-mixed, alkaline or saline, bare or covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil. Seasonality: April, June-September. Tenerals: August (ND). Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Davidson and Lee, 1990 (ecology); Fedorenko, 1996 (taxonomy). Dyschiriodes carri (Bousquet, 1996) Ecology. A sandy roadside ditch. Biology and Dispersal power. Unknown. Reference. Bousquet, 1996b (taxonomy). Comments. Subgeneric placement unknown. Dyschiriodes (Dyschiriodes) cerberus (Larson, 1968) Ecology. Unknown. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Eudyschirius) comatus (Bousquet, 1988) Ecology. A sea beach, on sandy soil. Nocturnal; sheltering during the day in burrows dug in the soil or under debris. Biology. Seasonality: April-July. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) compactus (Lindroth, 1961) Ecology. Uplands. Banks of rivers and brooks; lake shores. Open or shaded ground; moist, sandy soil, bare or covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil. Associated with staphylinids (Bledius). Biology. Seasonality: May-June, August. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy).

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Dyschiriodes (Dyschiriodes) consobrinus (LeConte, 1852) Ecology. Lowlands. Banks of small pools (in dunes) and streams. Open ground; moist, sandy soil, bare or covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil. Associated with staphylinids (Bledius). Biology. Seasonality: July. Tenerals: late July (OR). Dispersal power. Macropterous, capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Eudyschirius) criddlei (Fall, 1925) Ecology. Uplands. Saline or alkaline places. Open ground; moist soil. Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) curvispinus (Putzeys, 1846) Ecology. Lowlands. Banks of rivers and brooks; shores of lakes and ponds; borders of bogs and salt marshes. Open (mostly) or shaded ground; slightly moist, sandy, rarely saline soil covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil. Associated with staphylinids (Bledius). Biology. Seasonality: June, September. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Whitehead, 1970 (ecology); Fedorenko, 1996 (taxonomy). Dyschiriodes dejeanii (Putzeys, 1846) Natural History. Unknown. Comments. The subgeneric status was not dealt with by Fedorenko (1996). Dyschiriodes (Eudyschirius) edentulus (Putzeys, 1846) Ecology. Lowlands. Lake shores and river banks. Open ground; moist, sandy, alkaline or saline, bare soil. Nocturnal; sheltering during the day in burrows dug in the soil. Associated with staphylinids (Bledius). Biology. Seasonality: June-July. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) erythrocerus (LeConte, 1857) Ecology. Lowlands and mountains. Sea beaches and river banks. Open ground; wet, clayish or muddy soil. Nocturnal; sheltering during the day in burrows dug in the soil or under vegetal debris. Biology. Seasonality: April-September, November. Dispersal

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power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. Collecting techniques. Pouring water over the ground; light trapping. References. Lindroth, 1961 (natural history); Kirk, 1975c (dispersal power); Will et al. 1995 (dispersal power); Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) exochus (Whitehead, 1970) Ecology. Lowlands. Sea beaches. Open ground; moist, sandy, saline, bare soil. Nocturnal; sheltering during the day in burrows dug in the soil. Associated with staphylinids (Bledius). Biology. Seasonality: August. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Whitehead, 1970 (dispersal power); Fedorenko, 1996 (taxonomy). Dyschiriodes (Eudyschirius) ferrugineus (Bousquet, 1988) Ecology. Coastal lowlands. Estuarine flats. Open ground; wet, muddy soil covered with some vegetation (e.g., Salicornia). Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: November. Dispersal power. Wing-condition unknown. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) filiformis (LeConte, 1857) Ecology. Lowlands. Sea beaches, sand flats, and lake shores; borders of salt marshes; swamps; banks of pools and ditches. Open ground; moist, sandy, saline or alkaline soil, bare or covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil (mostly) and under pieces of wood. Gregarious. Associated with staphylinids (Bledius). Biology. Seasonality: January-October. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Whitehead, 1970 (ecology); Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) gibbipennis (LeConte, 1857) Ecology. Banks of brooks and rivers; lake shores. Open or shaded ground; moist, sandy soil, bare or covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil. Associated with staphylinids (Bledius). Biology. Seasonality: MarchApril, June. Predators: flycatchers. Dispersal power. Macropterous. Occasional in seashore drift material, which indicates flight ability. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Eudyschirius) globosus (Herbst, 1784) Ecology. Lowlands. Macrohabitat unknown. Under grass clippings (BC). Biology. Seasonality: July. Dispersal power. Unknown. References. Bousquet, 1996b and 2001 (taxonomy).

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Dyschiriodes (Dyschiriodes) globulosus (Say, 1823) Ecology. Lowlands and mountains. Wastelands, cultivated fields (e.g., corn, soybean, alfalfa, oat, flax, clover, carrot), grasslands, meadows, pastures, lawns, hills, sand pits, gravel pits, roadsides, ski fields, fencerows, forest clearings, and the upper zone of river banks. Open ground; moderately moist soil consisting of sand or sand-mixed clay, bare or covered with sparse tiny vegetation (e.g., moss). Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering during the day under small stones and dead leaves (mostly), plant debris, pieces of wood, in moss, under reindeer moss (Cladonia), heaps of grass, dry cow dung, and in burrows dug in the soil. Biology. Seasonality: January-December. Tenerals: January-February, April, June-August, October; mostly after mid-July. Adults found overwintering on higher and drier ground, in vacant lots, pastures, meadows, on hills and hillocks, in the upper zone of roadsides and sand pits, in hawthorn hedges, at the edge of woods, and in woods; in burrows (as deep as 7.5 cm), under small stones, dead leaves, and reindeer moss (Cladonia). Adult food: a small maggot-like larva. Predators: toads, bluebirds, and thrushes. Dispersal power. Wing-dimorphic: macropterous or brachypterous (incapable of flight). Occasional flier (to artificial lights at night; to artifical lights in captivity). Frequent in drift material, which indicates flight ability. Slow runner. Strong burrower. Strongly favored by human activities and spring floods along large rivers. Collecting techniques. Pitfall trapping; turning small stones; raking the leaf litter; turning drift material; baiting traps with molasses and fermented sugar. References. Lindroth, 1961 and 1963a (natural history); Kirk, 1969 (biology) and 1975c (ecology, dispersal power); Erwin, 1981 (natural history); Fedorenko, 1996 (taxonomy). Dyschiriodes (Eudyschirius) glomerosus (Bousquet, 1996), see Dyschiriodes (Eudyschirius) globosus (Herbst, 1784). Reference. Bousquet, 2001 (taxonomy). Dyschiriodes (Paradyschirius) haemorrhoidalis (Dejean, 1831) Ecology. Lowlands and mountains. Margins of rivers, brooks, pools, and ponds. Cultivated fields (e.g., cotton, tobacco). Open ground; wet or moist, bare clay soil, sometimes mixed with sand. Nocturnal; sheltering during the day in soil cracks or burrows dug in the ground. Gregarious. Biology. Seasonality: February-September. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Slow runner. Strong burrower. Collecting techniques. Pouring water over the ground; light trapping; turning drift material; pitfall trapping. References. Ulke, 1903 (ecology); Kirk, 1975c (dispersal power); Erwin, 1981 (natural history); Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) hiemalis (Bousquet, 1987), see Dyschiriodes (Dyschiriodes) melancholicus hiemalis (Bousquet, 1987). Reference. Fedorenko, 1996 (taxonomy).

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Dyschiriodes (Dyschiriodes) integer (LeConte, 1852) Ecology. Lowlands and mountains. Borders of eutrophic marshes; swamps; margins of ponds (freshwater, alkaline), lakes, pools, slow rivers, brooks, and irrigation ditches; vicinity of cattle water holes; mud flats; wet pastures, cultivated fields (e.g., carrot), and flood-plain forests. Open ground; moist wet clay soil (sometimes sand-mixed), often rich in organis debris, sometimes alkaline or saline, with bare spots surrounded by depressed sparse or moderate vegetation. Nocturnal; sheltering during the day in burrows and cracks in the soil, under mats of dead grass, stones, and at the base of plants. Gregarious. Associated with staphylinids (Bledius). Biology. Seasonality: March-November. Tenerals: late June-August. Adults found overwintering on higher and drier ground, in the upper zone of marsh borders and lake shores, and at the edge of woods; in burrows (as deep as 7.5 cm), moss, and leaf litter. Predators: frogs, toads, bluebirds, and robins. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in drift material, which indicates flight ability. Slow runner. Occasional climber (on plants). Strong burrower. Collecting techniques. Treading the vegetation into the water; light trapping; sweeping plants. References. Lindroth, 1955a (as nigripes), 1961, and 1963a (natural history); Kirk, 1975c (ecology, dispersal power); Will et al. 1995 (dispersal power); Fedorenko, 1996 (taxonomy). Dyschiriodes (Eudyschirius) interior (Fall, 1922) Ecology. Uplands. Margins of lakes, ponds, rivers, and brooks. Open ground; sandy, saline (mostly) or alkaline, bare soil. Nocturnal; sheltering during the day in burrows dug in the soil. Associated with staphylinids (Bledius). Biology. Seasonality: April, JuneJuly. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Lindroth, 1961 (natural history); Fedorenko, 1996 (taxonomy). Dyschiriodes (Antidyschiriodes) laevifasciatus (G.H. Horn, 1878) Ecology. Mountains. River banks. Open ground; moist bare soil composed of sand-mixed clay. Nocturnal; sheltering during the day in burrows dug in the soil. Associated with staphylinids (Bledius). Biology. Seasonality: April, June-July. Tenerals: mid-July (WA). Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Lindroth, 1961 (natural history); Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) larochellei (Bousquet, 1988) Ecology. Coastal lowlands. Sea beaches; borders of salt marshes and lagoons. Open ground; wet, loamy, muddy or clayish soil covered with sparse vegetation (e.g., Juncus, Salicornia). Nocturnal; sheltering during the day in burrows and cracks in the soil, and under mats of dead algae. Gregarious. Biology. Seasonality: January-September, November-December. Dispersal power. Macropterous, probably capable of flight. Slow

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runner. Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) longulus (LeConte, 1850) Ecology. Lowlands and mountains. Wastelands, grasslands, and margins of snowfields. Open ground; firm, moderately moist, sandy, gravelly or clayish soil, with some organic debris, covered with sparse vegetation (e.g. Agrostis, Potentilla). Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: April-August. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous (probably capable of flight). Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Lindroth, 1955a, 1961, and 1963a (natural history); Kirk, 1975c (dispersal power); Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) melancholicus hiemalis (Bousquet, 1987) Ecology. Lowlands, mountains, and subalpine zone. Peaty situations. Open ground; moderately moist soil. Nocturnal; sheltering during the day among moss, dead leaves, and in burrows dug in the soil. Biology. Seasonality: June-August. Tenerals: August (AB). Dispersal power. Brachypterous, incapable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) melancholicus melancholicus (Putzeys, 1866) Ecology. Lowlands and mountains. Lake beaches, pond edges, and mossy situations. Open ground; moist, sandy soil, bare or covered with sparse vegetation or moss carpets. Nocturnal; sheltering during the day in burrows dug in the soil, moss or leaf litter. Biology. Seasonality: June-July. Dispersal power. Brachypterous, incapable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) montanus (LeConte, 1879) Ecology. Lowlands and mountains. River banks (mostly) and pond margins. Moist or wet soil consisting of fine sand, sometimes silt-mixed, bare or covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil. Gregarious. Associated with staphylinids (Bledius). Biology. Seasonality: April-September. Tenerals: July (QU, WY). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Lindroth, 1961 (natural history); Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) pacificus (Lindroth, 1961) Ecology. Coastal lowlands. Sea beaches: sand flats (along dunes and freshwater flowing into or toward the ocean). Open ground; moist, sandy, saline soil, bare or covered

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with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil or under vegetal debris. Gregarious. Associated with staphylinids (Bledius). Biology. Seasonality: May-July. Dispersal power. Brachypterous, incapable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Lindroth, 1961 (natural history); Kavanaugh, 1992 (natural history); Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) pallipennis (Say, 1823) Ecology. Lowlands and mountains. River banks (mostly); shores of lakes, ponds, and pools; sea beaches. At a some distance from water. Open ground; steep, firm, moist, often saline soil consisting of fine sand, bare or covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil. Associated with staphylinids (Bledius). Biology. Seasonality: May, August, November. Defense-mechanism: body color-pattern matching the soil. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet. References. Lindroth, 1961 (natural history); Kirk, 1975c (ecology); Fedorenko, 1996 (taxonomy). Dyschiriodes (Eudyschirius) patruelis (LeConte, 1852) Ecology. Sandy places; mud flats. Open ground; moist soil. Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: April-May. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) perversus (Fall, 1922) Ecology. Mountains. River banks. Open ground; wet, sandy-muddy, bare soil. Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: JuneJuly. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) pilosus (LeConte, 1857) Ecology. Lowlands. Banks of rivers (mostly) and brooks; shores of lakes and ponds; pool margins. Open or half-shaded ground; moist or wet, clay soil, bare or covered with very sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil (mostly), under stones, and pieces of wood. Gregarious. Associated with staphylinids (Bledius). Biology. Seasonality: February-September. Tenerals: late July and August. Predators: thrushes. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Ulke, 1903 (ecology); Blatchley, 1910 (ecology; as hispidus (LeConte)); Lindroth, 1961 (natural history); Erwin, 1981 (natural history); Fedorenko, 1996 (taxonomy).

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Dyschiriodes (Dyschiriodes) planatus (Lindroth, 1961) Ecology. Grasslands. Open ground; rather dry soil covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: March-July, October. Dispersal power. Macropterous. Occasional in drift material, which indicates flight ability. Slow runner. Strong burrower. Collecting technique. Pitfall trapping. References. Lindroth, 1961 (natural history); Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) politus politus (Dejean, 1825) Ecology. Lowlands and mountains. Banks of rivers, brooks, and irrigation canals; shores of lakes, ponds, and marshes; borders of pools ( e.g., in sand pits and gravel pits). Open ground; moderately moist soil consisting of more or less clay-mixed sand, covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil. Gregarious. Associated with staphylinids (Bledius). Biology. Seasonality: April-September, November. Tenerals: July (WY), September (QU). Adults found overwintering on higher and drier ground, in the upper zone of gravel pits, in vacant lots, and in woods; under dead leaves, stones, and in rotten logs. Adult food, in the field: staphylinids (Europe). Predators: toads. Dispersal power. Macropterous. Frequent flier (at sunset; to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Slow runner. Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping. References. Lindroth, 1945, 1961, 1963a, and 1985 (natural history); Erwin, 1981 (natural history); Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) pumilus (Dejean, 1825) Ecology. Lowlands and uplands. Salt marshes, lake shores, pool margins, and river banks. Open ground; wet, sandy soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day in burrows dug in the soil and under debris. Biology. Seasonality: January, March-December. Tenerals: late July (AL). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Slow runner. Occasional climber (on plants). Strong burrower. Collecting techniques. Pouring water over the ground; light trapping; sweeping plants. References. Ulke, 1903 (ecology); Leng, 1915 (ecology, dispersal power); Fedorenko, 1996 (taxonomy). Dyschiriodes (Paradyschirius) quadrimaculatus (Lindroth, 1961) Ecology. Mountains. Upper zone of river banks (mostly) and lake shores. Open ground; steep, wet soil composed of sand-mixed clay, bare or covered with sparse vegetation (e.g., Equisetum). Nocturnal; sheltering during the day in burrows dug in the soil. Gregarious. Associated with staphylinids (Bledius). Biology. Seasonality: May-July, September. Tenerals: early July (WY). Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Lindroth, 1961 (natural history); Fedorenko, 1996 (taxonomy).

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Dyschiriodes (Dyschiriodes) salivagans (LeConte, 1875) Ecology. Lake shores, mud flats, and marsh borders. Open ground; moist soil, consisting of a loam-like mud, bare, close to clumps of vegetation. Nocturnal; sheltering during the day in burrows dug in the soil. Associated with staphylinids (Bledius). Biology. Seasonality: May-July. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) sculptus (Bousquet, 1988) Ecology. Coastal lowlands. Sea beaches. Open ground; moist, sandy bare soil. Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: February. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) sellatus (LeConte, 1857) Ecology. Lowlands and uplands. Sea beaches; saline banks of rivers, brooks, lakes, pools, and lagoons. Close to water. Open ground; wet, sandy, saline or alkaline soil, bare or almost so. Mostly nocturnal; sheltering during the day in burrows dug in the soil. Larvae travel under the soil surface, making dendritic networks of ridges. Gregarious. Associated with staphylinids (Bledius). Biology. Seasonality: May-September. Tenerals: mid-July (NY), August (QU). Defense-mechanism: when alarmed, the adult may escape by flying; body color-pattern matching the soil. Dispersal power. Macropterous. Occasional flier (during the day). Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Lindroth, 1955a, 1961, and 1963a (natural history); Kirk, 1975c (dispersal power); Wright, 1989 (ecology, biology); Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) setosus (LeConte, 1857) Ecology. Lowlands and mountains. River banks (mostly); margins of ponds and lakes; edges of pools (e.g., in sand pits and gravel pits); roadsides and meadows. Open ground; moderately moist, sandy or clayish soil, bare or almost so. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in burrows dug in the soil. Gregarious. Associated with staphylinids (Bledius). Biology. Seasonality: April-October. Tenerals: May-June (rarely), August, October. Dispersal power. Macropterous. Occasional in seashore drift material, which indicates flight ability. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Lindroth, 1961 (natural history); Fedorenko, 1996 (taxonomy).

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Dyschiriodes (Dyschiriodes) sextoni (Bousquet, 1987) Ecology. A tall-grass prairie (WI). Biology. Seasonality: June-July, September-October. Dispersal power. Macropterous, probably capable of flight. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) sphaericollis (Say, 1823) Ecology. Lowlands and mountains. Banks of rivers, brooks, and irrigation canals; shores of lakes, ponds, and pools (e.g., in sand pits and gravel pits); sea beaches; borders of lagoons; saline or alkaline flats. Close to water. Open ground; moist or wet bare soil consisting of sand (somewhat clay-mixed), often saline or alkaline. Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering during the day in burrows dug in the ground (mostly), in cracks in the soil, under vegetal debris or stones. Gregarious. Associated with staphylinids (Bledius). Biology. Seasonality: April-November. Copulating pairs: May (QU). Tenerals: late May-October. Adult food, in the field: staphylinid (Bledius) and heterocerid (Heterocera) larvae. Predators: toads. Phoretic mites found on adult body. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Slow runner. Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping. References. Leng, 1915 (dispersal power); Lindroth, 1961 and 1963a (natural history); Kirk, 1975c (dispersal power)); Erwin, 1981 (natural history); Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) subarcticus (Lindroth, 1961) Ecology. Banks of rivers and brooks. At some distance from water. Open or shaded ground; moist soil consisting of very fine sand, bare or covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil. Associated with staphylinids (Bledius). Biology. Seasonality: June-August. Tenerals: August (AK). Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Lindroth, 1961 (natural history); Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) sublaevis (Putzeys, 1846) Ecology. Lowlands. Sea beaches and salt marshes; banks of freshwater ponds and lakes. Open ground; moist or wet, sandy or sandy-muddy, often alkaline soil, bare or covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil. Gregarious. Associated with staphylinids (Bledius). Biology. Seasonality: June-August. Tenerals: late July (TX). Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy).

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Dyschiriodes (Eudyschirius) tenuispinus (Lindroth, 1961) Ecology. Uplands. Prairies. Open ground; soil covered with some vegetation (e.g., short grass). Nocturnal; sheltering during the day in tunnels dug in dry cow dung. Biology. Seasonality: April-June. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pitfall trapping. References. Bell, 1971 (ecology); Fedorenko, 1996 (taxonomy). Dyschiriodes (Paradyschirius) terminatus (LeConte, 1848) Ecology. Lowlands and uplands. Banks of large rivers. Open ground; moderately moist, sandy, bare soil. Nocturnal; sheltering during the day in burrows dug in the soil (mostly) and under debris. Associated with staphylinids (Bledius). Biology. Seasonality: MarchSeptember. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Slow runner. Strong burrower. Collecting techniques. Pouring water over the ground; light trapping. References. Blatchley, 1910 (biology); Kirk, 1969 (ecology); Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) timidus (Lindroth, 1961) Ecology. Uplands. Biology. Seasonality: May-July. Dispersal power. Brachypterous, incapable of flight. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Eudyschirius) tridentatus (LeConte, 1852) Ecology. Lowlands and mountains. Banks of rivers and brooks. Open (mostly) or shaded ground; moist or wet, bare sandy soil. Nocturnal; sheltering during the day in burrows dug in the soil. Associated with staphylinids (Bledius). Biology. Seasonality: February, April-August. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy). Dyschiriodes (Dyschiriodes) truncatus (LeConte, 1857) Ecology. Lowlands and mountains. Margins of lakes, slow rivers, and brooks. Open ground; moist bare soil consisting of clay-mixed sand. Nocturnal; sheltering during the day in burrows dug in the soil. Associated with staphylinids (Bledius). Biology. Seasonality: April-August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Lindroth, 1961 (natural history); Fedorenko, 1996 (taxonomy). Dyschiriodes (Eudyschirius) unipunctatus (Fall, 1901) Ecology. Unknown. Biology. Seasonality: May-July. Dispersal power. Macropterous, probably capable of flight. Reference. Fedorenko, 1996 (taxonomy).

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Dyschiriodes (Eudyschirius) varidens (Fall, 1910) Ecology. Mountains. Banks of rivers and brooks; shores of springs and lakes. Open or shaded ground; moist, sandy soil, bare or covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil. Gregarious. Associated with staphylinids (Bledius). Biology. Seasonality: March-August. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Fedorenko, 1996 (taxonomy). Dyschirius Bonelli, 1810, see Akephorus LeConte, 1851 (for marinus LeConte and obesus LeConte) and Dyschiriodes Jeannel, 1941 (for all other species). Reference. Fedorenko, 1996 (taxonomy). Elaphropus anceps (LeConte, 1848) Ecology. Lowlands, mountains, and subalpine zone. Upper banks of rivers, brooks, roadside ditches, lakes, pools (e.g., in sand pits and gravel pits), and cattle water holes; vacant lots and cultivated fields (e.g., corn). Open ground; moderately dry soil consisting of sand (often clay-mixed) or mud, bare or covered with sparse vegetation. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under stones and dead leaves (mostly), pieces of wood, plant rosettes (Verbascum), and in cracks in the soil. Biology. Seasonality: March-September. Tenerals: late June (rarely), August. Adults found overwintering on higher and drier ground, in the upper zone of brook banks and in vacant lots; under stones and dead leaves. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting techniques. Pouring water over the ground; using an aspirator. References. Lindroth, 1966 (natural history); Bell, 1971 (ecology); Will et al. 1995 (ecology, dispersal power). Elaphropus anthrax (LeConte, 1852) Ecology. Mountains. Drier zone of river and brook banks. Open ground; moderately moist, sandy, bare soil. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under stones. Biology. Seasonality: April, July. Dispersal power. Wing-condition unknown. Fast runner. Collecting technique. Pouring water over the ground. Elaphropus brevis (Casey, 1918) Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Elaphropus brunnicollis (Motschulsky, 1862) Ecology. Coastal lowlands. Sea beaches, lagoons, and pond margins. Open ground; moist, sandy, saline, bare soil. Diurnal; active in the sunshine. Biology. Seasonality: February,

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April, July. Dispersal power. Wing-condition unknown. Fast runner. Collecting technique. Pouring water over the ground. Elaphropus capax (LeConte, 1863) Ecology. Lowlands. Borders of marshes, swamps, bogs, pools, and streams. Open ground; wet, clayish or muddy soil, with some vegetation. Diurnal and nocturnal; active in the sunshine or sheltering on cloudy or cool days among sedges and cattails, under bark and moss. Biology. Seasonality: January-July, October-November. Dispersal power. Macropterous, probably capable of flight. Occasional flier (to artificial lights at night). Fast runner. Collecting techniques. Pouring water over the ground; lifting bark and moss from logs; treading sedges. References. Blatchley, 1910 (ecology); Bousquet, 1987a (ecology). Elaphropus cockerelli (Fall, 1907) Natural History. Unknown. Elaphropus congener (Casey, 1918) Ecology. Borders of potholes. Open ground. Biology. Seasonality: May-September. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Elaphropus conjugens (Notman, 1919) Natural History. Unknown. Elaphropus dolosus (LeConte, 1848) Ecology. Lowlands and mountains. River banks, lake shores, and reservoir borders. Open ground; moist, sandy, sometimes alkaline, bare soil. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days in cracks in the soil (mostly) and under debris. Gregarious. Biology. Seasonality: May-September. Tenerals: mid-July and August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Blatchley, 1910 (ecology); Lindroth, 1966 (natural history); Kirk, 1975c (dispersal power). Elaphropus fatuus (Casey, 1918) Natural History. Unknown. Elaphropus ferrugineus (Dejean, 1831) Ecology. Damp places; caves. Diurnal; active in the sunshine; frequently occurring in ant nests (Lasius aphidicola Walsh (= Lasius alienus Förster); L. umbratus Nylander), in rotting

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tree-stumps, under pieces of wood or under stones. Biology. Seasonality: April-September. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting techniques. Turning pieces of wood and stones; inspecting ant nests; dismembering rotting tree-stumps. References. Hayward, 1900 (ecology); Wickham, 1900 (ecology); Blatchley, 1910 (ecology); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Elaphropus fuscicornis (Chaudoir, 1868) Ecology. Vicinity of pools. Biology. Seasonality: January-April, September, November. Dispersal power. Wing-condition unknown. Fast runner. Elaphropus granarius (Dejean, 1831) Ecology. Lowlands and mountains. Vacant lots, pastures, meadows, roadsides, hills, sand pits, gravel pits; upper zone of river banks and pond shores; puddle margins and forest clearings. Open ground; dry, sandy, rarely gravelly soil, bare or covered with sparse vegetation (e.g., grass). Nocturnal; sheltering during the day under well-embedded stones and dead leaves (mostly), lichen, heaps of hay, and in the soil at the base of plants (e.g., Rumex). Gregarious. Biology. Seasonality: January-October, December. Tenerals: February (AL), August (VT). Adults found overwintering on higher and drier ground, in pastures, on hills and hillocks, and at the edge of woods; under well-embedded stones, in the soil (as deep as 25 cm), at the base of plants (e.g., grass), under dead leaves, and in trees. Dispersal power. Wing-dimorphic: macropterous or brachypterous (incapable of flight). Occasional flier (to artificial lights at night). Occasional in drift material, which indicates flight ability. Fast runner. Strong burrower. Collecting techniques. Pouring water over the ground; raking dead leaves. References. Blatchley, 1910 (ecology); Leng, 1915 (ecology); Lindroth, 1966 (natural history); Erwin, 1981 (natural history). Elaphropus incurvus (Say, 1830) Ecology. Lowlands and mountains. Upper zone of river banks, brooks, lakes, ponds, reservoirs, pools (even temporary), and roadside ditches. Wastelands, cultivated fields (e.g., corn, cabbage, raspberry), meadows, roadsides, sand pits, blowouts, dunes, ski fields, gravel pits, and open forests. Often near water. Open ground; rather dry, sandy soil (often clay-mixed), rarely alkaline, bare or covered with sparse vegetation. Mostly diurnal; active in the sunshine; frequently occurring in ant nests (Formica exsectoides Forel; Formica ulkei Emery), under stones (mostly), dead leaves, pieces of wood, dead fallen plants, silt slabs, dry bison dung, in cracks in the soil, and at the base of plants. Sometimes gregarious. Biology. Seasonality: January-November. Tenerals: May (rarely), mid-June to September; mostly August. Adults found overwintering on higher and drier ground, on the top of sandy mound and dunes; in ant nests (Formica exsectoides), under stones and in rotten logs. Adult food, in the field: ants (observed in ant food stores, attacking dead or disabled individuals). Adult food, in captivity: dipterous eggs. Predators: frogs and ants. Dispersal power. Macropterous. Frequent flier (in the sunshine in the afternoon; at sunset; to artificial lights at night; to artifical lights in captivi-

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ty). Occasional in seashore drift material, which indicates flight activity. Fast runner. Strong burrower. Favored by human activities (strongly) and spring floods along large rivers. Collecting techniques. Inspecting ant nests; turning stones; pouring water over the ground; pitfall trapping. References. Schwarz, 1890 (ecology); Ulke, 1893 (ecology); Leng, 1908 and 1915 (ecology); Blatchley, 1910 (ecology); Smith, 1910 (ecology); Holmquist, 1928 (ecology); Park, 1929 (ecology, biology); Lindroth, 1966 (natural history); Erwin, 1981 (natural history). Elaphropus levipes (Casey, 1918) Ecology. Lowlands. River banks. Open ground; soil composed of coarse sand mixed with silt. Nocturnal; sheltering during the day in the soil or under bark. Biology. Seasonality: April, August-September. Dispersal power. Macropterous. Occasional flier. Fast runner. Moderate borrower. Collecting technique. Pouring water over the ground. Reference. Erwin, 1981 (natural history). Elaphropus liebecki (Hayward, 1900) Natural History. Unknown. Elaphropus monticola (Casey, 1918) Natural History. Unknown. Elaphropus nebulosus (Chaudoir, 1868) Ecology. Lowlands and mountains. River and brook margins, lake shores, and reservoir borders. Open ground; moist, clayish, muddy or sandy, bare soil. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under stones and debris. Biology. Seasonality: May-August. Tenerals: late July (TX). Dispersal power.Wing-condition unknown. Fast runner. Collecting technique. Pouring water over the ground. Elaphropus obesulus (LeConte, 1852) Ecology. In decaying tree trunks (CA). Biology and Dispersal power. Unknown. Elaphropus occultus (LeConte, 1848) Natural History. Unknown. Elaphropus parvulus (Dejean, 1831) Ecology. Lowlands and mountains. Banks of rivers, brooks, and lakes; saline places close to the sea. Open ground; moist, gravelly or sandy, sometimes saline soil. Diurnal; active in the sunshine. Biology. Seasonality: April-July. Dispersal power. Macropterous. Flight

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records (ID, Europe). Fast runner. Clearly effective as a colonist. Collecting technique. Pouring water over the ground. References. Lindroth, 1945 and 1985 (ecology, dispersal power); Koch, 1989 (ecology); Luff, 1993 (biology); LaBonte and Nelson, 1998 (ecology). Elaphropus renoicus (Casey, 1918) Natural History. Unknown. Elaphropus rubricauda (Casey, 1918) Natural History. Unknown. Elaphropus saturatus (Casey, 1918) Ecology. Lowlands and mountains. Banks of rivers and brooks, drier zone of lake shores and pond margins, borders of marshes, wet meadows, cultivated fields (e.g., corn), and vacant lots. Open ground; moderately moist, sandy, clayish or gravelly soil, bare or covered with sparse vegetation. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under stones, among gravel, and under vegetal debris. Biology. Seasonality: March-September. Tenerals: mid-June to August; mostly from mid-July on. Adults found overwintering on higher and drier ground, in the upper zone of river and brook banks, in hawthorn hedges and vacant lots; in leaf litter. Dispersal power. Macropterous. Occasional in seashore drift material, which indicates flight ability. Fast runner. Collecting techniques. Pouring water over the ground; raking the soil. References. Lindroth, 1966 (natural history); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Elaphropus sectator (Casey, 1918) Natural History. Unknown. Elaphropus sedulus (Casey, 1918) Natural History. Unknown. Elaphropus tahoensis (Casey, 1918) Ecology. Mountains. Biology and Dispersal power. Unknown. Elaphropus tripunctatus (Say, 1830) Ecology. Lowlands and mountains. River banks. Open ground; moist, sandy-gravelly, bare soil. Crepuscular and nocturnal; sheltering during the day under stones and among gravel, also in crevices of steep banks. Gregarious. Biology. Seasonality: April-November. Tenerals: late June to August. Adults found overwintering on higher and drier ground, in the upper zone of river banks; in leaf litter. Predators: frogs and swallows.

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Dispersal power. Macropterous. Occasional flier (at sunset; to artificial lights at night). Fast runner. Favored by spring floods along large rivers. Collecting techniques. Pouring water over the ground; raking the soil. References. Lindroth, 1966 (natural history); Erwin, 1981 (natural history). Elaphropus vernicatus (Casey, 1918) (Tachys unionis Csiki, 1928) Ecology. Lowlands and mountains. Banks and shores of rivers and brooks (mostly), lakes, ponds, and marshes; roadsides, sand pits, vacant lots, and cultivated fields. Open ground; moist or wet soil consisting of clay, mud or sand, bare or covered with sparse vegetation. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days in ant nests (in large numbers), under stones, dead leaves, fallen dead plants, pieces of wood, and in cracks in the soil. Biology. Seasonality: February, April-September. Tenerals: early July (NB), September (MA). Adults found overwintering. Adult food, in the field: cruciferous plants. Dispersal power. Macropterous. Occasional in seashore drift material, which suggests flight ability. Fast runner. Strong burrower. Collecting techniques. Inspecting ant nests; pouring water over the ground; treading the soil with the feet. References. Casey, 1918 (ecology; as Tachyura laetifica n. sp.); Lindroth, 1966 (natural history; as unionis Csiki); Erwin, 1981 (natural history). Elaphropus vivax (LeConte, 1848) Ecology. Lowlands. Banks and shores: along rivers (mostly), brooks, lakes, and ponds. Open ground; moist or wet, sandy soil, bare or covered with sparse vegetation. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under stones, among gravel, in cracks in the soil, under dead leaves, and other debris. Gregarious. Biology. Seasonality: January, March-November. Tenerals: July-August (MD). Adults found overwintering on higher and drier ground, in forested areas; in wood debris and leaf litter. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Fast runner. Collecting techniques. Pouring water over the ground; raking the soil; using an aspirator. References. Lindroth, 1966 (natural history); Kirk, 1969 (biology); Erwin, 1981 (natural history); Hoffman, 1981 (ecology); Will et al. 1995 (dispersal power). Elaphropus xanthopus (Dejean, 1831) Ecology. Lowlands. River banks, margins of pools (in forests), lake shores, and mud flats in marshes. Cultivated fields (e.g., cotton) and abandoned fields. Open ground; moist, clayish, muddy or sandy-clayish soil, bare or covered with sparse vegetation. Diurnal; active in the sunshine. Biology. Seasonality: January-November. Adults found overwintering (SC). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight activity. Fast runner. Occasional climber (on trees). Collecting techniques. Pouring water over the ground; using tanglefoot screens. References. Blatchley, 1910 (ecology); Lindroth, 1966 (natural history).

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Elaphrus (Elaphrus) americanus americanus Dejean, 1831 Ecology. Lowlands and mountains. Beaches of ponds, marshes, lakes, pools, slow brooks and rivers; mud flats in sand pits, gravel pits, and fields; sea beaches. Open ground; firm, well-drained, almost horizontal, moist or wet soil consisting of coarse sand or clay, bare or with bare spots surrounded by sparse vegetation. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days at the base of plants (e.g., Carex, Juncus), in cracks in the soil, under vegetal debris, and stones. Gregarious. Biology. April-August. Copulating pairs: June (QU). Tenerals: mid-June to July. Adults found overwintering on higher and drier ground, in the upper zone of pond banks; under dead leaves. Adult food, in the field: flies and other insects. Adult and larval food, in captivity: mealworms. Predators: frogs and toads. Defense-mechanism: when alarmed, the adult may stridulate or escape by flying; body color matching the soil. Dispersal power. Macropterous. Frequent flier (in the sunshine; to artificial lights in captivity). Fast runner. Collecting technique. Hunting with a butterfly net. References. Lindroth, 1961 and 1963a (natural history); Goulet, 1983 (ecology). Elaphrus (Elaphrus) americanus sylvanus Goulet, 1982 Ecology. Lowlands, mountains, and subalpine zone. Beaches of streams, ponds, marshes, bogs, and seepages; vicinity of snowfields. Open ground; wet, clay soil, bare or covered with patchy vegetation (riparian habitats); moist soil, rich in organic debris, bare or covered with matted vegetation (snowfields). Diurnal; active in the sunshine. Biology. Seasonality: July-August. Adult and larval food, in captivity: mealworms. Defense-mechanism: when alarmed, the adult may stridulate; body color matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Hunting with a butterfly net. References. Goulet, 1983 (ecology); Kavanaugh, 1992 (natural history). Elaphrus (Elaphroterus) angusticollis angusticollis R.F. Sahlberg, 1844 Ecology. Mountains, subalpine and alpine zones. Beaches of large rivers. Often close to water. Ground shaded by bushes; wet soil consisting of sand-mixed clay, bare or covered with sparse vegetation (e.g., Equisetum) or some leaf litter (in the Salix-zone). Diurnal; active in the sunshine; often among leaves. Biology. Seasonality: June-August. Adult and larval food, in captivity: mealworms. Defense-mechanism: when alarmed, the adult may stridulate; body color matching the soil. Dispersal power. Wing-dimorphic: macropterous (mostly; probably capable of flight) or brachypterous (incapable of flight). Fast runner. Collecting technique. Hunting with a butterfly net. References. Lindroth, 1945, 1961, and 1985 (natural history); Goulet, 1983 (ecology). Elaphrus (Elaphrus) californicus Mannerheim, 1843 Ecology. Lowlands and mountains. Beaches of rivers and brooks (mostly), ponds, lakes, reservoirs, marshes, dugouts, and ditches; wet pastures. Open ground; well-drained, almost horizontal, moist (mostly) or wet clayish, sometimes saline or alkaline soil, bare

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or with bare spots surrounded by sparse vegetation. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under dead leaves and dead fallen plants, pieces of wood, dry cow dung, stones, and in cracks in the soil. Gregarious. Biology. Seasonality: April-December. Copulating pairs: May-June (QU). Tenerals: late May to August; mostly from early July on. Adults found overwintering on higher and drier ground, in the upper zone of sand pits and marshes; in the sandy soil (as deep as 5 cm). Adult food: flies, hymenopterans, other insects, spiders, and carrion (in the field); mealworms, earthworms, and meat (in captivity). Larval food, in captivity: mealworms. Predators: frogs. Defense-mechanism: when alarmed, the adult may stridulate or escape by running (mostly) or flying; body color matching the soil. Dispersal power. Macropterous. Frequent flier (in the sunshine; to artificial lights in captivity). Fast runner. Collecting technique. Hunting with a butterfly net. References. Lindroth, 1961 (natural history); Goulet, 1964, 1965a, and 1983 (natural history); Erwin, 1981 (natural history). Elaphrus (Neoelaphrus) cicatricosus LeConte, 1848 Ecology. Lowlands and mountains. Beaches of slow brooks running through swamps and swamp-forests. Ground shaded by bushes (e.g., Alnus) or trees; rather firm, wet, muddy soil, rich in organic debris. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under dead leaves. Biology. Seasonality: April-September. Tenerals: August (QU). Adult and larval food, in captivity: mealworms. Defense-mechanism: when alarmed, the adult may stridulate; body color matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting techniques. Collecting with with a butterfly net; raking the leaf litter. References. Frost, 1910b (ecology); Lindroth, 1961 (natural history); Goulet, 1983 (ecology). Elaphrus (Neoelaphrus) clairvillei Kirby, 1837 Ecology. Lowlands and mountains. Borders of eutrophic marshes, ponds, lakes, bogs, pools, slow rivers and brooks; borders of seepages and swamps; wet meadows and pastures; borders of salt marshes; swamp-forests and flood-plain forests. Frequently at some distance from water. Ground shaded by tall plants, shrubs or trees; soft, wet, muddy, rarely saline soil, rich in organic debris, sometimes with bare spots surrounded by vegetation. Diurnal; active in the sunshine. Larval habitat: mostly in the soil. Biology. Seasonality: March-December. Copulating pairs: May (QU). Oviposition: mid-May to late July. Tenerals: June-October. Adults found overwintering on higher and drier ground, on the upper banks of marshes, ponds, and sand pits; in hawthorn hedges, at the edge of woods, and in woods; in leaf litter, in the soil (as deep as 15 cm), under stones, the loose bark of fallen trees and logs. Life span: 1-2 summers. Adult food, in captivity: mealworms and meat. Larval food, in captivity: mealworms. Predators: frogs. Parasites: ichneumonids (attacking larvae). Phoretic mites found on adult body. Defensemechanism: when alarmed, the adult may stridulate; body color matching the soil. Dispersal power. Macropterous. Frequent flier (in the sunshine; at sunset; to artificial lights in captivity; into rotary traps). Frequent in seashore drift material, which indicates flight activity. Fast runner. Occasional climber (on plants). Collecting techniques. Hunting

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with a butterfly net; treading the soil with the feet; turning drift material; using rotary traps; sweeping plants. References. Lindroth, 1955a, 1961, and 1963a (natural history); Goulet, 1983 (natural history); Kavanaugh, 1992 (natural history). Elaphrus (Elaphrus) finitimus Casey, 1920 Ecology. Uplands. Banks of brooks, lakes, ponds, and ditches. Open ground; wet, clay, almost flat, bare soil. Diurnal; active in the sunshine. Biology. Seasonality: April-July. Adult and larval food, in captivity: mealworms. Defense-mechanism: when alarmed, the adult may stridulate; body color matching the soil. Dispersal power. Macropterous. Frequent flier (in the sunshine). Fast runner. Collecting technique. Hunting with a butterfly net. Reference. Goulet, 1983 (ecology). Elaphrus (Neoelaphrus) fuliginosus Say, 1830 Ecology. Lowlands and mountains. Margins of ponds and pool (in sand pits). Open ground; wet, sandy soil covered with sparse vegetation. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under dead leaves or stones. Solitary. Biology. Seasonality: April-September, November. Copulating pairs: early June. Tenerals: late July to August (QU). Adult and larval food, in captivity: mealworms. Predators: toads and starlings. Defense-mechanism: when alarmed, the adult may stridulate; body color matching the soil. Dispersal power. Macropterous. Occasional flier (to artificial lights in captivity). Fast runner. Collecting techniques. Hunting with a butterfly net; treading the soil with the feet; pouring water over the ground; raking the leaf litter; turning stones. References. Lindroth, 1961 (natural history); Larochelle, 1976b (ecology); Goulet, 1983 (ecology). Elaphrus (Neoelaphrus) laevigatus LeConte, 1852 Ecology. Lowlands and mountains. Marshy places. Ground shaded by trees; soft, black, muddy soil, rich in organic debris, covered with dense vegetation (e.g., Juncus). Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under plant debris. Biology. Seasonality: April-September. Oviposition: April (CA). Tenerals: late September (CA). Adult and larval food, in captivity: mealworms. Parasites: Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult may stridulate; body color matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting techniques. Hunting with a butterfly net; turning vegetal debris. Reference. Goulet, 1983 (natural history). Elaphrus (Arctelaphrus) lapponicus lapponicus Gyllenhal, 1810 Ecology. Lowlands, mountains, subalpine and alpine zones. Borders of wells, springs, brooks, small ponds, and trickles. Associated with cold water. Open ground; wet soil covered with thick moss carpets and sometimes scattered conifers. Diurnal; active in the sunshine on moss carpets. Biology. Seasonality: June-August. Tenerals: July (LB), August (QU). Adult and larval food, in captivity: mealworms. Defense-mechanism: when

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alarmed, the adult may stridulate; body color matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting techniques. Hunting with a butterfly net; treading thick moss carpets. References. Lindroth, 1945, 1961, and 1985 (natural history); Goulet, 1983 (natural history). Elaphrus (Arctelaphrus) lapponicus obliteratus Mannerheim, 1853 Ecology. Sphagnum-bogs. Open ground; wet soil. Diurnal; active in the sunshine on moss. Biology. Seasonality: unknown. Defense-mechanism: when alarmed, the adult may stridulate; body color matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Hunting with a butterfly net. Reference. Goulet, 1983 (ecology). Elaphrus (Elaphrus) lecontei Crotch, 1876 Ecology. Uplands. Beaches of lakes, ponds, reservoirs, marshes, sloughs, rivers, and brooks. Open ground; moist or wet, muddy, alkaline, bare soil. Close to water. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under stones or in soil crevices. Biology. Seasonality: April-August. Tenerals: early July (MB), August (WY). Adult and larval food, in captivity: mealworms. Defense-mechanism: when alarmed, the adult may stridulate; body color matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Hunting with a butterfly net. References. Lindroth, 1961 (natural history); Goulet, 1983 (natural history). Elaphrus (Neoelaphrus) lindrothi Goulet, 1983 Ecology. Lowlands. Cypress-swamps: on clay flats. Ground shaded by cypresses (Taxodium); wet, bare clay soil covered with scattered dead leaves or logs. Diurnal; active in the sunshine. Solitary. Biology. Seasonality: April-May, July. Tenerals: late July (VA). Defense-mechanism: when alarmed, the adult may stridulate; body color matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Hunting with a butterfly net. References. Blatchley, 1910 (ecology; as cicatricosus LeConte); Goulet, 1983 (ecology). Elaphrus (Elaphrus) marginicollis Goulet, 1983 Ecology. Mountains. Margin of a small spring, with moderately firm, wet soil, rich in organic debris. Diurnal; active in the sunshine. Biology. Seasonality: July. Defense-mechanism: when alarmed, the adult may stridulate; body color matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Reference. Goulet, 1983 (ecology). Elaphrus (Elaphrus) mimus Goulet, 1983 Ecology. Beaches of a small lake, on open ground with clay soil. Diurnal; active in the sunshine. Biology. Seasonality: May. Defense-mechanism: when alarmed, the adult may

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stridulate. Dispersal power. Macropterous, probably capable of flight. Fast runner. Reference. Goulet, 1983 (ecology). Elaphrus (Neoelaphrus) olivaceus LeConte, 1863 Ecology. Lowlands and mountains. Borders of eutrophic ponds, lakes, marshes, bogs, pools, slow rivers and brooks; swamps and margins of pasture puddles. Open ground; very soft, wet soil consisting of black mud, rich in organic debris, with bare spots surrounded by dense vegetation (e.g., Carex, Typha). Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under dead leaves and stones. Biology. Seasonality: April-September. Tenerals: July-September. Adults found overwintering on higher and drier ground; in moss and under the loose bark of fallen trees. Adult and larval food, in captivity: mealworms. Predators: toads. Defense-mechanism: when alarmed, the adult may stridulate or escape by running; body color matching the soil. Dispersal power. Macropterous. Occasional flier (in the sunshine; to artificial lights in captivity). Occasional in seashore drift material, which indicates flight activity. Fast runner. Collecting techniques. Hunting with a butterfly net; treading the soil with the feet. References. Lindroth, 1955a, 1961, and 1963a (natural history); Goulet, 1983 (natural history). Elaphrus (Elaphrus) parviceps Van Dyke, 1925, see Elaphrus (Elaphrus) trossulus Semenov, 1904. Reference. Goulet and Smetana, 1997 (taxonomy). Elaphrus (Elaphroterus) purpurans Hausen, 1891 (Elaphrus pallipes G.H. Horn, 1878) Ecology. Lowlands and mountains. Upper zone of river and brook banks. At some distance from water. Ground shaded by trees and bushes; rather dry, usually silty soil, bare or covered with dead leaves. Diurnal; active in the sunshine. Biology. Seasonality: AprilJuly. Tenerals: mid-July (WA). Adult and larval food, in captivity: mealworms. Defensemechanism: when alarmed, the adult may stridulate; body color matching the soil. Dispersal power. Macropterous. Occasional flier (into rotary traps). Fast runner. Collecting techniques. Hunting with a butterfly net; using rotary traps. References. Lindroth, 1961 (natural history; as pallipes G.H. Horn); Goulet, 1983 (ecology). Elaphrus (Elaphrus) ruscarius Say, 1830 Ecology. Lowlands and mountains. Beaches of slow brooks, small rivers, ditches, springs, lakes, and ponds (in swamps). Close to water. Open ground; wet, bare soil consisting of clay, sand, and silt, rich in organic debris. Diurnal; active in the sunshine. Biology. Seasonality: March-September. Tenerals: spring (AR). Adult food, in the field: lepidopterous caterpillars and grasshoppers. Adult and larval food, in captivity: mealworms. Predators: frogs, crows, redwings, robins, swallows, thrushes, and foxes. Defense-mechanism: when alarmed, the adult may run or stridulate; may also mimick the evasive

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behavior of wolf spiders and young toads (alternating between short, rapid runs and periods of immobility); body color matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Hunting with a butterfly net. References. Lindroth, 1961 (natural history); Brown and Thrall, 1974 (biology); Erwin, 1981 (natural history); Goulet, 1983 (ecology). Elaphrus (Elaphrus) trossulus Semenov, 1904 (Elaphrus (Elaphrus) parviceps Van Dyke, 1925) Ecology. Lowlands. Shores of small lakes and ponds. Open ground. Diurnal; active in the sunshine. Biology. Seasonality: June. Adults found overwintering under tree bark (over 1 km from its breeding habitat). Defense-mechanism: when alarmed, the adult may stridulate; body color matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Hunting with a butterfly net. References. Goulet, 1983 (ecology; as parviceps Van Dyke); Nelson, 1988 (ecology; as parviceps Van Dyke); Goulet and Smetana, 1997 (taxonomy). Elaphrus (Elaphrus) tuberculatus Mäklin, 1878 Ecology. Banks of large rivers. Open ground; moist, silty soil covered with dense vegetation (e.g., Equisetum). Diurnal; active in the sunshine. Biology. Seasonality: June-August. Adult and larval food, in captivity: mealworms. Defense-mechanism: when alarmed, the adult may stridulate; body color matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Hunting with a butterfly net. Reference. Goulet, 1983 (ecology). Elaphrus (Elaphrus) viridis G.H. Horn, 1878 Ecology. Margins of vernal pools. Moist, clayish-muddy soil, between Juncus-plants. Diurnal; active in the sunshine. Biology. Seasonality: May. Defense-mechanism: when alarmed, the adult may stridulate; body color matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Hunting with a butterfly net. Reference. Goulet, 1983 (natural history). Comments. Regarded as an endangered species. Elliptoleus acutesculptus Bates, 1882 Ecology. Mountains. Open pine-grassland savannahs. Ground shaded by trees. Biology. Seasonality: July (USA); April-May, July-August (Mexico). Dispersal power. Wingpolymorphic: brachypterous (mostly; incapable of flight) to macropterous. Moderate runner. Reference. Liebherr, 1991a (ecology, dispersal power). Eucaerus (Eucaerus) varicornis LeConte, 1853 Ecology. Lowlands. Swamp-forests and borders of swamps. Close to water. Shaded ground; wet soil covered with some vegetation (e.g., Cladium). Nocturnal; sheltering

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during the day in leaf litter about sedge roots, in half-submerged logs, or under boards. Biology. Seasonality: February, May, July-August. Adults found overwintering under vegetal debris. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Raking the leaf litter. Reference. Löding, 1945 (biology). Eucheila (Inna) boyeri (Solier, 1835). Ecology. Lowlands and uplands. A deciduous forest. A thorn wood. A tropical scrub, in bromeliads growing on trees (Mexico). Nocturnal. Biology. Seasonality: January-August (USA, Mexico, Central America). Dispersal power. Macropterous, probably capable of flight. References. Ball and Shpeley, 1983 (ecology); Shpeley and Ball, 2000 (taxonomy, ecology). Euphorticus occidentalis G.H. Horn, 1891 Ecology. Lowlands and mountains. Stream margins. Moist, sandy soil. Biology. Seasonality: May, August. Dispersal power. Wing-condition unknown. Moderate runner. Euphorticus pubescens (Dejean, 1831) Ecology. Lowlands. Borders of small ponds, pools, and waterways; edges of ditches near artesian wells. Open ground; moist or wet, peaty or sandy soil covered with some vegetation (e.g., grass). Diurnal; active in the sunshine; sheltering on cloudy or cool days among the vegetation and in burrows dug in the soil. Biology. Seasonality: January-November. Adults found overwintering on higher and drier ground, in the upper zone of sand pits; in the soil at the base of plants (Gramineae). Defense-mechanism: body color matching the soil. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Leng, 1915 (ecology); Kirk, 1969 (ecology, dispersal power); Liebherr, 1988c (ecology). Euproctinus (Neoeuproctus) abjectus (Bates, 1883) Ecology. On mimosa shrubs and bromeliads growing on oak-trees (Quercus; Mexico) or on cotton (TX). Biology. Seasonality: January-December. Dispersal power. Wing-condition unknown. Frequent climber (on trees and plants). Collecting technique. Beating or sweeping the vegetation. Euproctinus (Neoeuproctus) balli Shpeley, 1986 Ecology. Forests (mostly Quercus; also Arbutus, Juniperus, Pinus). Occurring in tree bromeliads. Biology. Seasonality: March-October. Dispersal power. Wing-condition unknown. Frequent climber (on trees). Collecting technique. Beating trees.

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Euproctinus (Neoeuproctus) trivittatus (LeConte, 1878) Ecology. Lowlands. Biology. Seasonality: February-May. Dispersal power Unknown. Euryderus grossus (Say, 1830) Ecology. Lowlands and uplands. Prairies, cultivated fields (e.g., corn, oat), rangelands, vacant lots, dunes, blowouts, orchards (e.g., peach), and salt flats. Open ground; dry, sandy soil covered with some vegetation (e.g., grass). Mostly nocturnal; active on cloudy days; sometimes active in the sunshine; usually sheltering in burrows dug in the soil, under dry cattle dung, boards, and logs. Larval habitat: in the soil. Biology. Seasonality: May-September. Tenerals: late May (BC), August (AB). Adult food, in the field and in captivity: grass seeds (e.g., Bromus). Predators: toads and crows. Dispersal power. Macropterous. Frequent flier (often swarming to artificial lights at night). Slow runner. Occasional climber (on plants). Strong burrower. Collecting techniques. Pitfall trapping; light trapping; turning debris; sweeping plants. References. Emden, 1942 (biology; as zabroides LeConte); Chu, 1945 (biology; as zabroides LeConte); Lindroth, 1968 (natural history); Kirk, 1975c (ecology, dispersal power); Lavigne, 1977 (ecology, biology). Evolenes exarata (Dejean, 1831) Ecology. Lowlands. Macrohabitat unknown. Nocturnal; sheltering during the day under pieces of wood. Biology. Seasonality: January, March-April, June, December. Tenerals: April (DC). Adults found overwintering in tree hollows. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Reference. Bousquet, 1996a (taxonomy, ecology). Galerita (Galerita) aequinoctialis Chaudoir, 1852 Ecology. Lowlands. Macrohabitat unknown. Nocturnal; sheltering during the day under stones and logs. Gregarious. Biology. Seasonality: unknown. Adult food, in the field: small insects. Defense-mechanism: when alarmed, the adult may emit a smoky burning liquid from the pygidial glands. Dispersal power. Macropterous, probably capable of flight. Fast runner. Galerita (Progaleritina) atripes LeConte, 1858 Ecology. Lowlands and mountains. Mostly deciduous forests (e.g., Quercus, Carya); also cultivated fields (e.g., cotton). Shaded ground; moderately moist soil, with thick leaf litter. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: June-July, October. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous, probably capable of flight. Fast runner. Occasional climber (on plants). Collecting techniques. Pitfall trapping; raking the leaf litter; sweeping plants. Reference. Ball and Nimmo, 1983 (ecology).

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Galerita (Progaleritina) bicolor (Drury, 1773) Ecology. Lowlands and mountains. Deciduous forests (mostly), coniferous forests (e.g., Pinus), forest clearings, and cultivated fields (e.g., peanut, alfalfa). Shaded ground: moderately moist soil. Nocturnal; sheltering during the day under the loose bark of rotten logs, in rotten tree-stumps, under logs, pieces of wood, and stones. Gregarious. Biology. Seasonality: January-December. Copulating pairs: late May (IN). Eggs are laid in purse-shaped mud cells attached to the underside of smooth leaves (Impatiens). Adults found overwintering in woods, at the edge of woods, and in vacant lots; under fallen trees and pieces of fallen bark, stones, and dead leaves. Adult food, in the field: lepidopterous caterpillars and other insects. Defense-mechanism: when alarmed, the adult may emit a Brachinus-like smoke from the pygidial glands. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in beach drift material, which indicates flight ability. Fast runner. Frequent climber (on plants and trees; up to 20 m above ground). Collecting techniques. Pitfall trapping; light trapping; baiting traps with carrion; lifting loose bark; beating trees. References. Blatchley, 1910 (ecology); King, 1919 (biology); Löding, 1945 (ecology); Erwin, 1981 (natural history); Ball and Nimmo, 1983 (ecology); Anderson et al. 1995 (ecology, biology). Galerita (Progaleritina) forreri Bates, 1883 Ecology. Mountains. A deciduous forest (Quercus; AZ). A thorn scrub-forest (Mexico). Shaded ground; dry or moderately moist soil. Nocturnal; sheltering during the day under dry cow dung. Biology. Unknown. Dispersal power. Wing-dimorphic: macropterous (mostly; probably capable of flight) or brachypterous (incapable of flight). Reference. Ball and Nimmo, 1983 (ecology). Galerita (Progaleritina) janus (Fabricius, 1792) Ecology. Lowlands and mountains. Deciduous forests, their clearings and edges, hedges, orchards, and adjacent cultivated fields (e.g., corn, alfalfa, wheat, soybean, tobacco), meadows, pastures, vacant lots, relict prairies, wastelands, fencerows, and gravel pits. Shaded or open ground; moderately moist soil, often covered with thick leaf litter. Nocturnal; sheltering during the day under the loose bark of logs and fallen trees, under logs, stones, pieces of wood, in leaf litter, and under other debris. Gregarious (in winter). Larval habitat: in woods, under old leaves and in rotten logs. Larvae are nocturnal. Biology. Seasonality: January-December. Tenerals: August (NY). Adults found overwintering in woods and at the edge of woods; under the loose bark of logs, fallen trees, stones, and in the soil; sometimes hibernating in colonies with Brachinus, Chlaenius, and Dicaelus. Adult food: lepidopterous caterpillars and grass seeds (in the field); lepidopterous caterpillars and meat (in captivity). Economic importance: may play an important role in the control of lepidopterous caterpillars. Predators: frogs, raccoons, bluebirds, crows, starlings, and woodpeckers. Parasites: Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult may stridulate or emit a pungent, smoky, audible Brachinus-like formic acid secretion from the pygidial glands. Dispersal power. Macropterous. Frequent flier (to artificial

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lights at night). Frequent in lakeshore drift material, which indicates flight ability. Frequent climber (adult and larva; on logs and trees). Fast runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; baiting traps with syrup; lifting loose bark; turning logs and stones; light trapping; beating trees; turning drift material; sugaring trees. References. Schaupp, 1879b (biology); Blatchley, 1910 (ecology); Leng, 1915 (dispersal power); Wade, 1921 (biology); Löding, 1945 (ecology); Kirk, 1969 (ecology); Lindroth, 1969a (ecology, dispersal power); Larochelle, 1974a (ecology); Kirk, 1975c (ecology); Erwin, 1981 (natural history); Ball and Nimmo, 1983 (ecology); Lesiewicz et al. 1983 (ecology, biology); Anderson et al. 1995 (ecology, biology). Galerita (Progaleritina) lecontei lecontei Dejean, 1831 Ecology. Lowlands. Borders of rivers, brooks, marshes, and swamps; pastures and grasslands. Open ground; wet or moist soil. Nocturnal; sheltering during the day under the loose bark of logs or under logs. Larval habitat: in the soil. Biology. Seasonality: January, March-December. Adults found overwintering at the edge of woods; under the loose bark of fallen trees. Predators: crows. Parasites: Laboulbeniales (Fungi). Defensemechanism: when alarmed, the adult may emit a spray from the pygidial glands. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Fast runner. Frequent climber (on logs and trees). Collecting techniques. Pitfall trapping; light trapping; lifting loose bark; beating trees; sugaring trees. References. Leng, 1915 (collecting technique); Löding, 1945 (ecology); Kirk, 1969 (ecology) and 1975c (ecology, dispersal power); Ball and Nimmo, 1983 (ecology). Galerita (Progaleritina) mexicana Chaudoir, 1872 Ecology. Lowlands and mountains. Forest clearings, palm forests, thorn forests, and rainforests. Open or slightly shaded ground; wet to dry soil covered with thick leaf litter. Nocturnal; sheltering during the day in leaf litter and under other debris. Biology. Seasonality: June. Defense-mechanism: when alarmed, the adult may emit an acrid smell from the pygidial glands. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Collecting techniques. Raking the leaf litter; pitfall trapping. References. McCullough, 1971 (biology); Ball and Nimmo, 1983 (ecology). Galerita (Progaleritina) reichardti Ball and Nimmo, 1983 Ecology. Unknown. Biology. Seasonality: March. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Reference. Ball and Nimmo, 1983 (dispersal power). Gastrellarius blanchardi (G.H. Horn, 1891) Ecology. Mountains. Forests (e.g., Picea, Fagus). Shaded ground; moderately moist soil covered with thick leaf litter. Nocturnal; sheltering during the day among dead leaves

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around rotten logs (mostly) and under stones. Biology. Seasonality: April-August, October, December. Dispersal power. Brachypterous, incapable flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the loose leaf litter around rotten logs. References. Barr, 1969 (ecology); Hoffman, 1998 (ecology, biology). Gastrellarius honestus (Say, 1823) Ecology. Lowlands and mountains. Deciduous (mostly), mixed, and coniferous forests (e.g., Picea, Abies); thickets (Rhododendron). Shaded ground; moderately moist soil covered with thick leaf litter and scattered fallen trees. Nocturnal; sheltering during the day mostly under the loose bark of trees (fallen and standing) and logs, under and in rotten logs, under pieces of wood and stones, in leaf litter, and in mushrooms. Larval habitat: in decaying wood. Biology. Seasonality: January-November. Gravids: May (QU). Tenerals: May (rarely), July-September. Adults found overwintering in woods; in fallen tree trunks. Adult food: Gypsy Moth immature stages (in the field); mealworms, lepidopterous cocoons, meat, and apple (in captivity). Larval food, in captivity: mealworms. Parasites: nemathelminth worms and ascid mites. Phoretic mites found on adult body. Defense-mechanism: when alarmed, the adult may emit a strong smell from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Excellent climber (on trees, tree-stumps, and logs; up to 2 m above ground). Collecting techniques. Lifting the loose bark of trees; pitfall trapping; night collecting with a headlamp. References. Blatchley, 1910 (ecology); Lindroth, 1966 (natural history); Barr, 1969 (ecology); Erwin, 1981 (natural history); Reeves et al. 1983 (biology); Levesque and Levesque, 1986 (biology); Bousquet, 1986b (biology). Gastrellarius unicarum (Darlington, 1931) Ecology. Mountains. Coniferous forests (e.g., Picea): often along slopes. Shaded ground; moderately moist soil covered with thick leaf litter. Nocturnal; sheltering during the day among loose dead leaves lying around logs. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the loose leaf litter along rotten logs. Reference. Barr, 1969 (ecology). Gehringia olympica Darlington, 1933 Ecology. Mountains. Banks and bars of cold, swift, clear brooks (mostly) and moderately large rivers. Close to water. Open ground; sloping, moist soil consisting of very fine, clean gravel and coarse sand, bare or sparsely covered with tiny moss. Nocturnal; sheltering during the day under stones and among gravel. The adult seldom comes to the soil surface. Gregarious. Biology. Seasonality: June-August. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting techniques. Turning stones; pouring water over the ground; raking the loose gravel. References. Lindroth, 1960 and 1961 (natural history).

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Geopinus incrassatus (Dejean, 1829) Ecology. Lowlands and uplands. Dunes, hills, blowouts and sand pits: often along rivers and brooks. Sandy roads and cultivated fields (e.g., wheat, cabbage, flax, corn, oat). Frequently near water. Open ground; well-drained, often sloping, moderately dry soil consisting of very loose, fine sand, covered with sparse vegetation. Nocturnal; active between sunset and midnight; sheltering during the day in burrows dug in the soil (as deep as 30 cm), and under well-embedded logs, pieces of wood, and stones. Gregarious. Biology. Seasonality: February-December. Tenerals: mid-June (QU), August (SA), September (QU). Adults found overwintering (SC). Adult food: seedlings and cutworms (in the field); a dead fly (in captivity). Economic importance: sometimes a pest of seed beds. Predators: toads (regularly), crows, flycatchers, kites, owls, robins, mice, and raccoons. Defensemechanism: body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (sometimes swarming by the hundreds to artificial lights at night). Slow runner. Strong burrower. Strongly favored by human activities. Collecting techniques. Pitfall trapping; night collecting with a headlamp; light trapping. References. Ulke, 1903 (dispersal power); Blatchley, 1910 (dispersal power); Everly and Everly, 1936 (ecology); Lindroth, 1968 (natural history); Kirk, 1969 (ecology, dispersal power), 1970 (dispersal power), and 1975c (ecology, dispersal power); Landry, 1980a (ecology, biology); Erwin, 1981 (natural history); Johnson and Clark, 1989 (natural history). Halocoryza arenaria (Darlington, 1939) Ecology. Coastal lowlands. Sea beaches: intertidal area (FL), at or near the high tide line (outside the U.S.A.). Open ground; wet soil consisting of coquina-coral cemented by very fine silt or sand, covered with seaweed mats. Associated with the centipede Pectiniunguis halirrhytus Crabill. Biology. Seasonality: July. Dispersal power. Wing-polymorphic: brachypterous (FL; incapable of flight) to macropterous (outside the U.S.A.; probably capable of flight). Slow runner. References. Whitehead, 1966 and 1969 (ecology, dispersal power). Harpalus (Pseudoophonus) actiosus Casey, 1914 Ecology. Lowlands and uplands. Cultivated fields (e.g., wheat, alfalfa), roadsides, edges of forests, open deciduous and coniferous forests. Open or shaded ground; moderately dry soil composed or sand or sandy loam, covered with some vegetation. Nocturnal; sheltering during the day under stones and dead leaves. Biology. Seasonality: January, April-July. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones; raking the leaf litter. Reference. Kirk, 1975c (ecology). Harpalus (Harpalus) affinis (Schrank, 1781) Ecology. Lowlands, mountains, subalpine and alpine zones. Vacant lots, pastures, meadows, cultivated fields and their borders (e.g., corn, cabbage, asparagus, barley, wheat, rye, alfalfa, hay, potato, carrot, raspberry), lawns, golf courses, gardens, farmyards, sand

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pits, gravel pits, sand dunes, ski fields, roads, streets, sidewalks, orchards (e.g., apple, peach), forest clearings, and open forests. Open ground; more or less dry, sandy soil covered with weeds. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days in the soil at the base of plants (e.g., Solidago, Verbascum), under stones, dead fallen plants, pieces of wood, and dead leaves. Gregarious. Biology. Seasonality: January-December. Copulating pairs: May to early August (QU); coitus lasting 68 minutes on average. Gravids: mid-April to late August (QU). Tenerals: early May to October; mostly from mid-July on. Adults found overwintering on higher and drier ground, in vacant lots, pastures, meadows, on hills and hillocks, in the upper zone of sand pits and gravel pits, and at the edge of woods; in the soil, under plant rosettes (Verbascum), stones, dead leaves, and pieces of cardboard. Adult food: seeds, grass tissue, dipterous larvae and pupae, lepidopterous caterpillars, and wireworms (in the field); seeds, lepidopterous caterpillars and pupae, dipterous eggs and pupae, aphids, scarabaeid larvae, dead carabids, and earthworms (in captivity). Larval food, in the field: curculionid larvae and pupae. Predators: toads (frequently), frogs (regularly), bluebirds, and crows. Parasites: Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult may emit a malodorous liquid from the pygidial glands. Dispersal power. Macropterous. Frequent flier (in the spring sunshine; to artificial lights at night; to artifical lights in captivity). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants and trees). Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Pitfall trapping; digging at the base of plants; turning stones and vegetal debris; light trapping; turning drift material; sweeping plants. References. Lindroth, 1945, 1955a, 1963a, and 1968 (natural history); Briggs, 1965 (biology; as aeneus (Fabricius)); Finlayson and Campbell, 1976 (biology; as aeneus (Fabricius)); Holliday and Hagley, 1978 (ecology, biology); Tyler and Ellis, 1979 (biology); Lindroth, 1986 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology); Levesque and Levesque, 1994 (biology); Will et al. 1995 (dispersal power). Harpalus (Harpalomerus) amputatus amputatus Say, 1830 Ecology. Lowlands, mountains, subalpine and alpine zones. Grasslands, prairies, pastures, meadows, cultivated fields and their borders (e.g., wheat, barley, corn, alfalfa, sainfoin), roadsides, shrub-deserts, open forests; borders of sloughs, rivers, reservoirs, cattle water holes, irrigation ditches, and seepages. Greenhouses (occasionally). Open ground; dry, often sandy soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day under dry cattle dung, stones, mats of dead plants, and other debris. Biology. Seasonality: February-November. Copulating pairs: May (ID). Gravids: early May, mid-July, and August (AB). Tenerals: April-November; mostly July-August. Adult food: wheat kernels (in the field); lepidopterous eggs and larvae (in captivity). Predators: crows. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning debris; using malaise traps. References. Bugbee, 1942 (biology); Lindroth, 1968 (natural history); Frank, 1971b (biology); Kirk, 1975c (ecology, dispersal power); Lester and Morrill, 1989 (ecology); Noonan, 1991 (natural history); Kataev, 1997 (taxonomy); Ball and Bousquet, 2001 (taxonomy).

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Harpalus (Euharpalops) animosus Casey, 1924 Ecology. Mountains, subalpine and alpine zones. Meadows, pastures, moraines, and forest clearings. Borders of brooks, small lakes, and seepages. Open ground; dry soil covered with some vegetation (e.g., grass). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under stones, logs, pieces of wood, and dry cow dung. Biology. Seasonality: April-September. Tenerals: April-August; mostly August. Dispersal power. Macropterous. Occasional flier (to artificial lights at light). Moderate runner. Collecting techniques. Pitfall trapping; turning debris. References. Lindroth, 1968 (natural history); Noonan, 1991 (natural history). Harpalus atrichatus Hatch, 1949 Ecology. Lowlands and mountains. Biology. Seasonality: May-August. Dispersal power. Macropterous, probably capable of flight. Harpalus balli Noonan, 1991 Ecology. Lowlands. Biology. Seasonality: May. Dispersal power. Macropterous. Occasional in seashore drift material, which indicates flight ability. Harpalus (Megapangus) caliginosus (Fabricius, 1775) Ecology. Lowlands and uplands. Relict prairies, grasslands, meadows, pastures, cultivated fields (e.g., corn, oat, wheat, barley, tobacco, cotton, soybean, peanut, alfalfa, asparagus, cabbage, flax), stubble fields, old fields, vacant lots, golf courses, lawns, fencerows, roadsides, sand pits, gravel pits, orchards (e.g., plum, citrus), and open forests. Greenhouses (occasionally). Open ground; rather dry, usually sandy soil covered with sparse vegetation (e.g., grass). Diurnal and nocturnal; active on plants and on the ground; otherwise sheltering during the day in burrows dug in the soil (mostly; as deep as 8.5 cm), under stones, straw mulch, plant debris, logs, pieces of wood and cardboard. Gregarious. Larval habitat: in burrows (15 cm in length on average). Biology. Seasonality: February-December. Copulating pairs: early July (once), September (mostly). Gravids: mid-September (QU). Tenerals: April-September, mostly July-August. Adults found overwintering on higher and drier ground, on hills; under stones. Adult food: seeds and pollen of plants (e.g., Ambrosia, Helianthus, Fragaria, Trifolium, Phleum), lepidopterous caterpillars, tenebrionids, coccinellids, chrysomelids, scarabaeids, flies, acridids, ant pupae and adults, and mites (in the field); plant seeds (e.g., Triticum, Cassia, Ambrosia, Amaranthus, Datura), rose haws, berries (Polygonatum), apple, peach, banana, boiled wheat, mealworms, meat, and cheese (in captivity). Larval food, in the field: curculionid adults and larvae. Economic importance: may play an important role in the control of lepidopterous caterpillars; sometimes a severe pest of strawberries. Predators: frogs, toads (regularly), quails, cardinals, crows, ducks, grackles, grouses, robins, starlings, badgers, foxes, raccoons, and spiders. Often crushed by pedestrians and cars. Parasites: paramegistid mites. Defense-mechanism: when alarmed, the adult may stridulate or

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emit a defensive fluid accompanied with a pungent vinegar-like smell and a large amount of Brachinus-like smoke. Dispersal power. Macropterous. Frequent flier (sometimes swarming by the thousands to artificial lights at night). Moderate runner. Frequent climber (on plants and shrubs). Strong burrower. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning debris; light trapping; sweeping or beating the vegetation. References. Forbes, 1880 (biology); Hamilton, 1888a (ecology, biology); Horn, 1888 (biology); Ulke, 1890 (biology, dispersal power) and 1903 (dispersal power); Webster, 1900 (biology); Blatchley, 1910 (natural history); Bryson, 1939 (ecology); Fattig, 1933 (biology); McCullough, 1966c (biology); Lindroth, 1968 (natural history); Kirk, 1969, 1970 (dispersal power), and 1975c (ecology, dispersal power); Erwin, 1981 (natural history); Lesiewicz et al. 1983 (ecology, biology); Noonan, 1991 (natural history); Will et al. 1995 (dispersal power). Harpalus cautus Dejean, 1829 Ecology. Lowlands and mountains. Meadows, pastures, roadsides, ridges, forest clearings, and open forests. Greenhouses (occasionally). Open ground; moderately dry, often sandy soil covered with sparse to moderate vegetation (e.g., grass, herbs, shrubs). Nocturnal; sheltering during the day under dry cow dung, stones, and in decaying sagebrush trunks. Biology. Seasonality: February-November. Tenerals: April-September; mostly August. Adult food, in the field and captivity: Pseudotsuga-seeds. Predators: spiders (up to 134 individuals found as prey of the Black Widow Spider, Latrodectus mactans (Fabricius)). Dispersal power. Wing-dimorphic: brachypterous or macropterous. Occasional in seashore drift material, which indicates flight ability. Moderate runner. Moderately favored by human activities. Collecting techniques. Pitfall trapping; turning debris. References. Exline and Hatch, 1934 (biology); Dick and Johnson, 1958 (biology); Lindroth, 1968 (natural history); Noonan, 1991 (natural history). Harpalus (Glanodes) cohni Ball, 1972 Ecology. Uplands. Grasslands. Open ground; dry, clayish, muddy or sandy soil. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Reference. Ball, 1972 (natural history). Harpalus (Pseudoophonus) compar LeConte, 1848 (Harpalus bicolor (Fabricius, 1775)) Ecology. Lowlands and mountains. Grasslands, relict prairies, pastures, cultivated fields (e.g., corn, flax, tobacco, asparagus, alfalfa, cabage, carrot), vacant lots, golf courses, fencerows, sand pits, gravel pits, roadsides, orchards (e.g., apple), forest edges and clearings, open forests, and tree plantations (e.g., Pinus). Caves (occasionally). Open ground; well-drained, moderately dry, often sandy soil covered with some vegetation (e.g., grass). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under plant debris, logs, and pieces of wood. Biology. Seasonality: January-December. Copulating pairs: May and August (QU). Gravids: late July to September (ON, QU). Ten-

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erals: July-September. Adults found overwintering on higher and drier ground, in vacant lots, fencerows, roadsides, on hills, and in the upper zone of sand pits; in the soil, under logs, stones, and pieces of cardboard. Adult food, in the field: plant seeds (Ambrosia) and lygaeids. Predators: toads (frequently), frogs, sparrows, and starlings. Defense-mechanism: when alarmed, the adult may emit a Brachinus-like smoke from the pygidial glands. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Occasional climber (on plants). Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning vegetal debris; light trapping; baiting traps with carrion; sweeping plants. References. Ball and Anderson, 1962 (ecology); Rivard, 1964c (biology) and 1965 (dispersal power); Lindroth, 1968 (natural history; as bicolor (Fabricius)); Kirk, 1975c (ecology, dispersal power); Will et al. 1995 (dispersal power). Harpalus (Opadius) cordifer Notman, 1919 Ecology. Lowlands and uplands. Mostly forests and their clearings; sometimes thickets or bushes (in fields). Shaded or open ground; moderately dry soil covered with thick leaf litter and humus. Nocturnal; sheltering during the day in leaf litter, humus, moss, under stones and logs, or in the soil. Biology. Seasonality: January-November. Tenerals: May, July-September. Adult food, in captivity: dipterous eggs. Dispersal power. Wingdimorphic: macropterous (probably capable of flight) or brachypterous (incapable of flight). Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Lindroth, 1968 (natural history); Noonan, 1991 (natural history). Harpalus (Glanodes) corpulentus (Casey, 1914) Ecology. Uplands. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Harpalus (Glanodes) cunctipeps Ball, 1973 Ecology. Uplands. Grasslands. Open ground; dry, clay soil (gravel-mixed) covered with sparse vegetation (e.g., grass, cacti). Nocturnal; sheltering during the day under dry cattle and horse dung, stones, and heaps of dead plants. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Reference. Ball, 1972 (natural history; as puncticeps Ball). Harpalus (Pharalus) cyrtonotoides Notman, 1919 Ecology. Uplands. Biology and Dispersal power. Unknown. Reference. Ball and Bousquet, 2001 (taxonomy). Harpalus (Pharalus) desertus LeConte, 1859 Ecology. Uplands, mountains, and subalpine zone. Grasslands, prairies, meadows, rangelands, pastures, cultivated fields and their borders (e.g., wheat, barley, oat, corn), road-

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sides, deserts, canyons, forest clearings, open forests, and vicinity of lakes, streams, and cattle water holes. Open ground; dry, sandy soil covered with sparse vegetation (e.g., grass). Mostly nocturnal; quite active in the spring sunshine; usually sheltering during the day under stones, dry cow dung, pieces of wood, other debris, and in ground tunnels. Biology. Seasonality: March-October, December. Tenerals: April-May, July-October; mostly July-August. Adult and larval food, in captivity: grasshopper eggs. Predators: spiders. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (to artificial lights at night). Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning debris. References. Lindroth, 1968 (natural history); Kirk, 1975c (ecology, dispersal power); Lavigne, 1977 and 1978 (ecology, biology); Noonan, 1991 (natural history). Harpalus ellipsis LeConte, 1848 Ecology. Uplands and mountains. Rangelands, pastures, meadows, gravel pits; borders of rivers, drainage ditches, and cattle water holes; open forests. Open ground; moderately dry, sandy or silty soil covered with sparse vegetation (e.g., grass, herbs). Nocturnal; sheltering during the day under dry cow and buffalo dung, stones, and pieces of wood. Biology. Seasonality: January-November. Predators: bluebirds and woodpeckers (flickers). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning debris. References. Lindroth, 1968 (natural history); Kirk, 1975c (ecology); Noonan, 1991 (natural history). Harpalus (Plectralidus) eraticus Say, 1823 Ecology. Lowlands and mountains. Meadows, pastures, grasslands, cultivated fields (e.g., corn, oat, flax, soybean, cabbage, asparagus), wastelands, dunes, sand pits, gravel pits, river banks, orchards (e.g., apple), and open forests (e.g., Pinus). Open ground; dry soil consisting of fine sand, bare or covered with sparse vegetation. Mostly nocturnal; sometimes active in the sunshine; sheltering during the day in burrows dug in the ground (mostly; as deep as 15-60 cm), cracks in the soil, under logs, stones, boards, dead leaves, pieces of wood, dry cow dung, and other debris. Gregarious. Larval habitat: in a burrow (as deep as 60 cm). Biology. Seasonality: April-November. Copulating pairs: August-September (SD, IN). Gravids: July-October (ON), late August to October (SD). Eggs are laid below the entrance of a burrow (5-12 cm deep). Tenerals: May (rarely), July-September; mostly August. Adults found overwintering in fields (mostly) and in woods; in burrows dug in the soil (mostly; as deep as 30-45 cm) and under stones. Life cycle: 2 years. Adult food: corn (stalks, kernels, roots), chrysomelids, scarabaeid eggs, and other insects (living, injured or dead) (in the field); chrysomelid larvae (in captivity). Larval food, in the field: plant seeds (e.g., Setaria) previously stored and pressed into the walls of its burrow. Predators: toads (frequently), crows, pheasants, starlings, raccoons, and skunks. Parasites: rickettsias. Defense-mechanism: body color matching the soil. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strong burrower. Strongly favored by human activities. Collecting technique. Pitfall trapping. References. Rivard,

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1964c (biology) and 1965 (dispersal power); Lindroth, 1968 (natural history); Kirk, 1972, 1974 (biology), and 1975c (ecology); Noonan, 1991 (natural history). Harpalus (Pseudoophonus) erythropus Dejean, 1829 Ecology. Lowlands and mountains. Meadows, pastures, relict prairies, wastelands, cultivated fields (e.g., corn, soybean, tobacco, alfalfa), semi-open fields, fencerows, gardens, gravel pits, roadsides, forest edges, and open forests (e.g., Quercus). Open ground; rather dry, sandy or clayish soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day under stones, dead leaves and plants, and pieces of wood. Biology. Seasonality: January-October. Copulating pairs: late September (IN). Tenerals: June-October. Adults found overwintering in open woods; under stones. Adult food: scarabaeid larvae (in the field); chrysomelid prepupae and pupae (in captivity). Predators: frogs, toads, and starlings. Parasites: Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult may emit a Brachinus-like smoke from the pygidial glands. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; to artifical lights in captivity). Moderate runner. Occasional climber (on trees). Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning debris; light trapping. References. Blatchley, 1910 (biology); Ball and Anderson, 1962 (ecology); Lindroth, 1968 (natural history); Kirk, 1970 and 1975c (dispersal power); Erwin, 1981 (natural history). Harpalus (Pseudoophonus) faunus Say, 1823 Ecology. Lowlands and mountains. Meadows, pastures, relict prairies, cultivated fields (e.g., oat, wheat, tobacco, cabbage, alfalfa, asparagus, flax), old fields, vacant lots, fencerows, roadsides, sand pits, gravel pits, and open forests. Open ground; dry, sandy soil covered with some vegetation (e.g., grass). Mostly nocturnal; sheltering during the day under fallen pieces of bark, boards, logs, stones, dead leaves, and in the soil at the base of plants. Semi-gregarious. Biology. Seasonality: January-November. Gravids: September (QU). Tenerals: April (rarely), late June to August. Adults found overwintering (SC). Adult food, in the field: plant seeds (Ambrosia) and lepidopterous caterpillars. Predators: toads, woodpeckers, and starlings. Parasites: tapeworms. Defense-mechanism: body color matching the soil. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; to artifical lights in captivity). Moderate runner. Occasional climber (on plants). Strongly favored by human activities. Collecting techniques. Pitfall trapping; lifting wood debris; light trapping; sweeping plants. References. Larochelle, 1974a (ecology); Kirk, 1975c (natural history); Erwin, 1981 (natural history); Anderson et al. 1995 (ecology). Harpalus (Euharpalops) fraternus LeConte, 1852 Ecology. Lowlands, mountains, and subalpine zone. Grasslands, meadows, rangelands, pastures, cultivated fields (e.g., corn, hay), roadsides, open forests; borders of seepages and cattle water holes. Greenhouses (occasionally). Open ground; dry, often sandy soil covered with sparse vegetation (e.g., grass). Mostly nocturnal; sheltering during the day under dry

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cow and bison dung (mostly), stones, pieces of wood, logs, and fallen sagebrush trunks. Biology. Seasonality: March-October. Tenerals: April-October; mostly August. Adult food, in the field: strawberries, tenebrionids, and grasshoppers. Adult and larval food, in captivity: grasshopper eggs. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Occasional climber (on plants). Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning dry dung; sweeping plants. References. Lindroth, 1968 (natural history); Kirk, 1975c (ecology, dispersal power); Parmenter and MacMahon, 1988 (biology); Noonan, 1991 (natural history). Harpalus fulgens Csiki, 1932, see Trichotichnus (Iridessus) fulgens (Csiki, 1932). Reference. Ball and Bousquet, 2001 (taxonomy). Harpalus (Euharpalops) fulvilabris Mannerheim, 1853 Ecology. Lowlands, montains, subalpine and alpine zones. Forests, their clearings and edges; pastures, vacant lots, moraines, gravel pits and sand pits near forests; thickets (e.g., Alnus); coastal tundra. Ground shaded by bushes, shrubs or deciduous trees; moderately dry soil covered with thick leaf litter and humus; open ground (seacoast and above timber limit). Nocturnal; sheltering during the day among dead leaves, in humus, under pieces of wood, logs, stones, and pieces of cardboard. Biology. Seasonality: May-October. Gravids: early July to August (AB, QU). Tenerals: May-October; mostly June. Adult food, in the field: dipterous eggs. Adult and larval food, in captivity: mealworms. Predators: toads. Phoretic mites found on adult body. Dispersal power. Wingdimorphic: brachypterous (mostly) or macropterous. Occasional in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Lindroth, 1955a, 1963a, and 1968 (natural history); Noonan, 1991 (natural history). Harpalus (Harpalobius) fuscipalpis Sturm, 1818 (Harpalellus basilaris (Kirby, 1837)) Ecology. Lowlands, mountains, subalpine and alpine zones. Grasslands, meadows, moraines, cultivated fields and their borders (e.g., corn, wheat, barley, alfalfa, cabbage), vacant lots, abandoned fields, dunes, and gravel pits.Open ground; dry, sandy soil covered with sparse vegetation. Nocturnal; sheltering during the day under stones, plant rosettes (Verbascum), dead leaves, and pieces of cardboard. Biology. Seasonality: April-October. Tenerals: mid-May (BC), late June (AK). Adults found overwintering in vacant lots; under plant rosettes (Verbascum). Adult food, in the field: plant seeds (Descurainia). Predators: crows. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Occasional climber (on plants, e.g., Asclepias). Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; sweeping plants. References. Lindroth, 1968 (natural history; as basilaris (Kirby)); Kirk, 1975c (ecology, dispersal power).

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Harpalus (Pharalus) gravis LeConte, 1858 Ecology. Lowlands. Pastures, meadows, cultivated fields (e.g., cotton, corn, soybean), orchards (e.g., peach); borders of rivers and brooks in flood plains. Open ground. Nocturnal; sheltering during the day under dry cattle dung, pieces of wood, and dead leaves. Biology. Seasonality: January, March-December. Tenerals: April, June-October; mostly August. Adults found overwintering in meadows; under heaps of hay. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; light trapping; turning dry cattle dung. References. Kirk, 1969 (ecology, dispersal power) and 1970 (dispersal power); Noonan, 1991 (natural history). Harpalus (Pseudoophonus) hatchi Ball and Anderson, 1962 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Harpalus herbivagus Say, 1823 Ecology. Lowlands, mountains, subalpine and alpine zones. Grasslands, pastures, meadows, shrub-steppes, cultivated fields and their borders (e.g., corn, wheat, alfalfa, barley, asparagus, clover, oat, flax, soybean, cotton, cabbage, carrot), lawns, vacant lots, roadsides, fencerows, sand pits, gravel pits, sand dunes, hills, ski fields, orchards (e.g., peach, apple), forest clearings, and open forests (e.g., Quercus); borders of lakes, marshes, and streams. Open ground; dry, sandy soil covered with sparse vegetation (e.g., grass, herbs). Mostly nocturnal; active in the spring sunshine; usually sheltering during the day under dead leaves, stones, dead plants, reindeer moss (Cladonia), plant rosettes (Verbascum), in the soil at the base of plants (e.g., Solidago), under boards, logs, and dry dung (cow, bison). Biology. Seasonality: January-December. Copulating pairs: April-June (QU, SD); coitus lasting 7-15 minutes. Gravids: April-May and mid-July (QU). Tenerals: March-December; mostly August-October. Adults found overwintering on higher and drier ground, in vacant lots, meadows, pastures, in the upper zone of sand pits and gravel pits, at the edge of woods, and in woods; in the soil at the base of plants, under plant rosettes (Verbascum), in leaf litter, and under stones. Adult food: grass shoots (Poa), grass seeds, fungi, lepidopterous caterpillars, scarabaeids, flies, and cabbage worms (in the field); earthworms and meat (in captivity). Larval food, in the field: grasshopper eggs. Predators: toads (frequently), frogs, bluebirds, crows, woodpeckers (flickers), pheasants, robins, starlings, and thrushes. Parasites: podapolipid mites and tapeworms. Dispersal power. Macropterous. Frequent flier (in the spring sunshine; at dusk; to artificial lights at night). Frequent in lakeand seshore drift material, which indicates flight ability. Moderate runner. Occasional climber. Favored by human activities (strongly) and spring floods along large rivers. Collecting techniques. Pitfall trapping; turning debris; light trapping; turning drift material. References. Webster, 1880 (biology); Lindroth, 1968 (natural history); Kirk, 1975c (natural history); Doane, 1981 (ecology, biology); Erwin, 1981 (natural history); Reeves et al. 1983 (biology); Noonan, 1991 (natural history).

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Harpalus (Glanodes) huachuca Ball, 1972 Ecology. Uplands. Biology and Dispersal power. Unknown. Harpalus (Pharalus) indianus Csiki, 1932 Ecology. Lowlands. Sand dunes, sand hills, sand blowouts, wastelands, pastures, and cultivated fields (e.g., corn, cucumber). Open ground; dry, sandy soil covered with sparse vegetation. Nocturnal; sheltering during the day under dead leaves, pieces of wood, and other debris. Biology. Seasonality: June-October. Defense-mechanism: body color matching the soil. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pitfall trapping. References. Blatchley, 1910 (ecology; as testaceus (LeConte)); Lindroth, 1968 (natural history); Kirk, 1975c (ecology); Noonan, 1991 (natural history). Harpalus (Euharpalops) indigens Casey, 1924 Ecology. Lowlands and mountains. Grasslands, relict prairies, meadows, pastures, vacant lots, cultivated fields (e.g., cabbage), hills, roadsides, sand pits, gravel pits, orchards (e.g., apple), forest clearings, and open forests. Open ground; soft, dry, sandy soil covered with sparse vegetation (e.g., grass). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under dead leaves, stones, reindeer moss (Cladonia), pieces of wood or cardboard, and in the soil at the base of plants (e.g., Rumex). Semigregarious. Biology. Seasonality: May-October. Tenerals: late June to August; mostly after mid-July. Adults found overwintering on hills, in meadows, and wood clearings; in leaf litter and under stones. Adult food, in the field: lepidopterous caterpillars. Predators: toads (frequently). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Collecting technique. Raking the leaf litter. References. Lindroth, 1968 (dispersal power); Larochelle, 1971d (ecology); Noonan, 1991 (natural history). Harpalus innocuus LeConte, 1863 Ecology. Lowlands, mountains, subalpine and alpine zones. Sagebrush-steppes, rangelands, vacant lots, and meadows. Open ground; sandy or gravelly soil covered with sparse vegetation (e.g., grass, herbs). Nocturnal; sheltering during the day under stones. Biology. Seasonality: January, March-October. Tenerals: May-August. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Turning stones. References. Lindroth, 1971a (natural history); Parmenter and MacMahon, 1988 (biology); Noonan, 1991 (natural history). Harpalus (Megapangus) katiae Battoni, 1985 Ecology. Lowlands. Fields, sand dunes, and sandy alluvial habitats. Open ground; dry, sandy soil. Mostly nocturnal; usually sheltering during the day. Biology. Seasonality:

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March, June-October. Dispersal power. Macropterous. Regular flier (to artificial lights at night). Moderate runner. Collecting technique. Light trapping. References. Nunn, 1986 (dispersal power); Will, 1997 (ecology, dispersal power). Harpalus (Euharpalops) laevipes Zetterstedt, 1828 (Harpalus quadripunctatus Dejean, 1829; Harpalus egregius Casey, 1914) Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines covered with scattered bushes (e.g., Alnus); old fields, meadows, vacant lots, cultivated fields (e.g., raspberry), sand pits, gravel pits, forest clearings and edges, open forests, and burnt forests (from soon after fires up to several years after). Ground shaded by bushes (often; e.g., Rubus) or open; rather dry, often sandy soil (often humus-mixed) covered with sparse vegetation (bushes, grass). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under stones, pieces of wood, logs, pieces of cardboard, dead leaves, and dry cow dung. Biology. Seasonality: April-October. Tenerals: late May-August; mostly May. Predators: toads. Defense-mechanism: body luster matching charred wood. Dispersal power. Macropterous. Regular in sea- and lakeshore drift material, which indicates flight ability. Moderate runner. Favored by forest fires. Collecting techniques. Pitfall trapping; turning stones; turning drift material. References. Lindroth 1945, 1955a (as baergi Csiki), 1963a (as egregius Casey), 1968 (as egregius Casey), and 1986 (natural history); Koch, 1989 (ecology); Holliday, 1991 (ecology, dispersal power; as egregius Casey); Noonan, 1991 (natural history); Levesque and Levesque, 1994 (biology; as quadripunctatus Dejean). Harpalus (Euharpalops) laticeps LeConte, 1850 Ecology. Lowlands, mountains, subalpine and alpine zones. Open forests, burnt forests (soon after fire up to several years after), forest edges and adjacent fields, vacant lots, and sand pits; moraines and meadows. Often in the vicinity of bushes and trees. Open or shaded ground; dry, sandy (often) or gravelly soil covered with sparse vegetation (e.g., grass, herbs). Nocturnal; sheltering during the day under stones, logs, pieces of wood, dead leaves, and pieces of cardboard. Gregarious. Biology. Seasonality: AprilOctober, December. Gravids: mid-July and early August (QU). Tenerals: June. Defensemechanism: body luster matching charred wood. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Long-distance disperser. Favored by forest fires. Collecting techniques. Pitfall trapping; light trapping; turning stones. References. Blatchley, 1910 (ecology); Lindroth, 1963a and 1968 (dispersal power); Larochelle, 1974a (ecology); Richardson and Holliday, 1982 (ecology); Holliday, 1984 (ecology) and 1991 (biology); Noonan, 1991 (natural history). Harpalus (Harpalobrachys) leiroides Motschulsky, 1844 Ecology. Sandy areas near rivers and lakes. Biology. Seasonality: May-August. Tenerals: July to mid-August (AK). Dispersal power. Macropterous, probably capable of

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flight. Moderate runner. References. Lindroth, 1968 (natural history); Ball and Bousquet, 2001 (ecology). Harpalus (Euharpalops) lewisii LeConte, 1865 Ecology. Lowlands, mountains, subalpine and alpine zones. Open forests, burnt forests (soon after fires), forest clearings and edges, vacant lots, old fields, sand pits, pastures, meadows, cultivated fields, and roadsides. Open ground; soft, dry, sandy soil covered with sparse vegetation (e.g., grass). Nocturnal; sheltering during the day in the soil at the base of plants, under plant rosettes (Verbascum), reindeer moss (Cladonia), stones, dead leaves, fallen tree bark, and pieces of wood. Gregarious. Biology. Seasonality: MayAugust. Copulating pairs: late June (QU). Gravids: June and mid-August (QU). Tenerals: May-August; mostly August. Adult food, in captivity: meat and a dead zygopteran. Adults are cannibalistic in captivity. Predators: toads and crows. Defense-mechanism: body luster matching charred wood. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Strong burrower. Long-distance disperser. Favored by forest fires. Collecting techniques. Pitfall trapping; light trapping. References. Lindroth, 1968 (biology, dispersal power); Larochelle, 1972a (ecology, biology); Noonan, 1991 (natural history). Harpalus (Pseudoophonus) liobasis Chaudoir, 1868 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Harpalus (Pseudoophonus) longicollis LeConte, 1848 Ecology. Lowlands and mountains. Pastures, vacant lots, cultivated fields (e.g., tobacco, asparagus, cabbage, alfalfa), roadsides, sand pits, gravel pits, forest edges and clearings, open forests; upper zone of river and brook banks. Open ground; moderately dry or moist, often sandy soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day under stones (often), dead leaves, and dead fallen plants. Biology. Seasonality: January-February, April-November. Tenerals: July-August. Adults found overwintering in woods; under the loose bark of fallen trees and under stones. Adult food, in the field: scarabaeid larvae. Predators: frogs, toads, bluebirds, and crows. Defense-mechanism: when alarmed, the adult may emit a Brachinus-like smoke from the pygidial glands. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Occasional climber (on trees; up to 3 m above ground). Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; night collecting with a headlamp. References. Ball and Anderson, 1962 (ecology); Lindroth, 1968 (natural history); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Harpalus martini Van Dyke, 1926 Ecology. Uplands. Biology. Seasonality: May-June. Dispersal power. Macropterous, probably capable of flight.

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Harpalus megacephalus LeConte, 1848 Ecology. Lowlands. Deciduous (e.g., Populus) or coniferous forests. Shaded ground. Biology. Seasonality: June-August. Tenerals: late August (MB). Dispersal power. Macropterous. Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting technique. Pitfall trapping. References. Lindroth, 1968 (natural history); Noonan, 1991 (natural history). Harpalus nigritarsis C.R. Sahlberg, 1827 Ecology. Lowlands, mountains, subalpine and alpine zones. Meadows, moraines, pastures, shrub-steppes, vacant lots, roadsides, sand pits, and forest clearings. Open ground; firm, rather dry, usually gravelly soil covered with short dense vegetation (e.g., grass). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under stones (mostly), pieces of wood, fallen trees, pieces of cardboard, and dry cow dung. Biology. Seasonality: May-October. Tenerals: June-October; mostly July. Dispersal power. Macropterous. Frequent flier (to artificial lights in captivity). Frequent in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Turning stones; pitfall trapping; turning drift material. References. Lindroth, 1955a, 1963a, and 1968 (natural history); Noonan, 1991 (natural history). Harpalus (Glanodes) obliquus G.H. Horn, 1880 Ecology. Uplands. A grassland with damp, clay soil, under dry cattle dung and stones. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Reference. Ball, 1972 (natural history). Harpalus obnixus Casey, 1924 Ecology. Lowlands and mountains. Grasslands, pastures, sagebrush communities, hills, forest clearings, and open forests. Open ground; firm, dry soil consisting of silt (sometimes gravel-mixed) or loam, covered with sparse to moderate vegetation (e.g., grass, herbs). Nocturnal; sheltering during the day under pieces of wood, stones, and dry cow dung. Biology. Seasonality: March-September. Dispersal power. Wing-dimorphic: brachypterous (incapable of flight) or macropterous (probably capable of flight). Moderate runner. Collecting techniques. Pitfall trapping; turning debris. References. Lindroth, 1968 (natural history); Noonan, 1991 (natural history). Harpalus ochropus Kirby, 1837 Ecology. Lowlands and uplands. Moraines. Open ground; rather dry, sandy soil covered with sparse vegetation. Nocturnal; sheltering during the day under debris. Biology. Seasonality: May-August. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning debris. References. Lindroth, 1968 (natural history); Kirk, 1975c (ecology). Noonan, 1991 (natural history).

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Harpalus opacipennis (Haldeman, 1843) Ecology. Lowlands and mountains. Grasslands, meadows, pastures, cultivated fields and their borders (e.g., corn, wheat, cabbage), old fields, vacant lots, roadsides, gravel pits, sand pits, sandy ridges, hills, forest clearings, and open forests. Open ground; dry sandy or gravelly soil covered with sparse vegetation (e.g., grass). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under well-embedded stones, reindeer moss (Cladonia), dry cow or bison dung, dead leaves, pieces of wood (sometimes well-embedded) and cardboard. Gregarious (in winter). Biology. Seasonality: April-November. Copulating pairs: mid-July (QU). Tenerals: May-August, October-November; mostly from July on. Adults found overwintering on higher and drier ground, in pastures, on hills and hillocks, at the edge of woods, and in woods; under well-embedded stones (mostly) and plant rosettes (Verbascum). Predators: toads and frogs. Dispersal power. Macropterous. Occasional flier (in the sunshine). Moderate runner. Strong burrower. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning embedded stones; using interception traps. References. Lindroth, 1968 (natural history); Kirk, 1975c (ecology). Erwin, 1981 (natural history); Epstein and Kulman, 1990 (ecology, biology); Noonan, 1991 (natural history). Harpalus (Pseudoophonus) paratus Casey, 1924 Ecology. Lowlands and uplands. Grasslands, cultivated fields (e.g., corn), fencerows, and blowouts. Open ground; rather dry soil composed of sand or sandy clay, covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: June-August. Tenerals: July (SD). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; turning vegetal debris. References. Ball and Anderson, 1962 (ecology); Lindroth, 1968 (natural history); Kirk, 1975c (natural history). Harpalus (Pseudoophonus) pensylvanicus (DeGeer, 1774) Ecology. Lowlands and mountains. Cultivated fields and their borders (e.g., corn, tobacco, cotton, asparagus, cabbage, tomato, oat, soybean, beans, peanut, alfalfa, hop, strawberry, carrot), stubble fields, pastures, grasslands, rangelands, meadows, old fields, vacant lots, lawns, gardens, fencerows, sand pits, sand dunes, gravel pits, roadsides, ravines; borders of ponds, lakes, and rivers; orchards (e.g., apple, peach), forest edges, and open forests. Greenhouses (occasionally). Open ground; dry soil consisting of sand or sandy loam, covered with moderately dense but frequently tall vegetation. Mostly nocturnal; sometimes active in the sunshine and before sunset; usually sheltering during the day mostly in cracks in the soil, entrances to mouse burrows, cavities dug by field crickets, holes dug by Harpalus pensylvanicus adults, also under clods of earth, in the soil at the base of plants, under dead leaves, plant debris, pieces of wood, stones, and reindeer moss (Cladonia). Gregarious. Larval habitat: in a burrow (30 cm deep on average). Biology. Seasonality: January-December. Copulating pairs: June, August-September. Gravids: JulyOctober. Eggs are laid singly beneath the soil surface (5-15 cm deep). Tenerals: mostly

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early July to mid-August. Adults found overwintering in cultivated fields, vacant lots, fencerows, in the upper zone of sand pits and gravel pits, at the edge of woods, and in wood clearings; in cells and burrows (mostly; 8-20 cm deep) dug in the soil, under logs, stones, and in leaf litter. Adult food: plant seeds, tissue, and pollen (e.g., Ambrosia, Echinochloa, Fragaria, Phleum, Poa, Rumex, Setaria, Triticum, and Zea), fungi, lepidopterous caterpillars, curculionid larvae, chrysomelid eggs and adults, coccinellids, scarabaeid larvae and adults, carabids, grasshoppers, honey bees, and mites (in the field); aphids (in captivity). Larval food, in the field: seeds, chrysomelids, scarabaeid larvae, and other insects. The larva stores plant seeds (e.g., Setaria) in a burrow and presses them into its walls. Economic importance: may play an important role in the control of crop pests; sometimes a pest of strawberries. Predators: toads (frequently), frogs, quails, catbirds, crows, woodpeckers (flickers), flycatchers, grackles, grouses, pheasants, sparrows, starlings, foxes, and spiders. Parasites: hymenopterans, mites, tapeworms, gregarines, Laboulbeniales (Fungi), and ricksettsias. Defense-mechanism: when alarmed, the adult may stridulate or emit a Brachinus-like smoke from the pygidial glands. Dispersal power. Macropterous. Frequent flier (often swarming to artificial lights at night, sometimes to the point of becoming a public nuisance). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Frequent climber (on plants and trees). Strong burrower. Strongly favored by human activities. Collecting techniques. Pitfall trapping; light trapping; turning debris; sweeping or beating the vegetation; using emergence traps; baiting traps with carrion. References. Forbes, 1880 (biology); Webster, 1880 (biology); Hamilton, 1888a (ecology, biology); Blatchley, 1910 (biology, dispersal power); McDermott, 1916 (dispersal power); Fattig, 1937 (biology); Chu, 1945 (biology); Ball and Anderson, 1962 (ecology); Rivard, 1964c (biology), 1965 (dispersal power), and 1966 (biology); Lindroth, 1968 (natural history); Kirk, 1972, 1973, and 1975c (natural history); Lund, 1975 (biology); Tomlin, 1975a (biology); Monteith, 1976 and 1977 (biology); Best and Beegle, 1977 (biology); Best et al. 1981 (biology, dispersal power); Erwin, 1981 (natural history); Lesiewicz et al. 1983 (ecology, biology); Los and Allen, 1983 (biology); Ferguson and McPherson, 1985 (ecology, biology); Barney and Pass, 1986a (biology); Perfecto et al. 1986 (ecology, dispersal power); Morrill, 1992 (biology); Anderson et al. 1995 (ecology); Riddick and Mills, 1995 (biology); Will et al. 1995 (dispersal power). Harpalus plenalis Casey, 1914 Ecology. Lowlands and mountains. Hills, pastures, vacant lots, roadsides, sand pits, and gravel pits. Open ground; dry, usually sandy soil covered with sparse vegetation (e.g., grass). Mostly nocturnal; sometimes active in the spring sunshine; usually sheltering during the day under dead leaves, stones, well-embedded pieces of wood, reindeer moss (Cladonia), plant rosettes (Verbascum), and in the soil at the base of plants (e.g., Selene). Gregarious. Biology. Seasonality: March-November. Copulating pairs: April and mid-July (QU). Tenerals: April-October. Adults found overwintering in sand pits, gravel pits, pastures, and on hills; under stones and plant rosettes (Verbascum). Adult food, in captivity: corn kernels. Dispersal power. Macropterous. Occasional flier (in the sunshine). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strong burrower. Strongly favored by human activities. Collecting techniques.

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Pitfall trapping; raking the leaf litter. References. Lindroth, 1955a, 1963a, and 1968 (natural history); Noonan, 1991 (biology). Harpalus (Pseudoophonus) poncei Will, 2001 Ecology. Lowlands. Biology. Seasonality: December. Dispersal power. Unknown. Reference. Will, 2001 (taxonomy). Harpalus (Pseudoophonus) protractus Casey, 1914 Ecology. Lowlands. A meadow with rather dry clay soil covered with some vegetation (e.g., grass, shrubs). Nocturnal; sheltering during the day under stones. Biology. Seasonality: April-July, September-October. Tenerals: early June (AL). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Reference. Ball and Anderson, 1962 (ecology). Harpalus (Euharpalops) providens Casey, 1914 (Harpalus viduus LeConte, 1865) Ecology. Lowlands and mountains. Open or light deciduous (e.g., Acer) forests, their edges and clearings, thickets (e.g., Juniper), and adjacent roadsides, sand pits, gravel pits, hills, and vacant lots. Ground shaded by trees and shrubs or open; soft, rather dry sandy soil covered with sparse vegetation (e.g., grass) or thick leaf litter. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in burrows or cavities dug in the soil (mostly; as deep as 7.5 cm), under dead leaves, stones, reindeer moss (Cladonia), at the base of plants (e.g., Oenothera), and under pieces of cardboard. Gregarious. Biology. Seasonality: January, March-December. Tenerals: March-November; mostly August-September. Adults found overwintering on higher and drier ground, in woods, wood clearings, vacant lots, in the upper zone of roadsides, and on hillocks; under stones and in leaf litter. Adult food: Gypsy Moth lepidopterous caterpillars (in the field); meat (in captivity). Predators: toads (frequently). Dispersal power. Macropterous. Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Strong burrower. Collecting techniques. Pitfall trapping; digging into the sand; night collecting with a headlamp. References. Hamilton, 1886 (ecology, biology; as viduus LeConte); Lindroth, 1968 (natural history; as viduus LeConte); Larochelle, 1974a (ecology; as viduus LeConte); Erwin, 1981 (natural history; as viduus LeConte); Noonan, 1991 (natural history; as viduus LeConte). Harpalus (Ophonus) puncticeps (Stephens, 1828), see Ophonus puncticeps Stephens, 1828. References. Kataev, 1996 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Harpalus (Glanodes) regressus (Casey, 1914) Natural History. Unknown.

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Harpalus (Plectralidus) retractus LeConte, 1863 Ecology. Uplands. Grasslands, pastures, corrals, roadsides; borders of cattle water holes; open forests. Open ground; dry, sandy soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day under plant debris. Biology. Seasonality: JuneSeptember. Tenerals: July-September; mostly August. Defense-mechanism: body color matching the soil. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; turning plant debris. Reference. Noonan, 1991 (natural history) Harpalus (Euharpalops) reversus Casey, 1924 (Harpalus funerarius Csiki, 1932) Ecology. Lowlands and mountains. Grasslands, rangelands, pastures, cultivated fields and their borders (e.g., wheat, barley, oat, corn, cabbage, flax, potato), vacant lots, sand pits, and open forests. Open ground; very soft, dry soil consisting of very fine sand, covered with sparse vegetation (e.g., grass). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in the soil (mostly), under plant rosettes (Verbascum), stones, logs, dead leaves, and dry cow dung. Gregarious. Biology. Seasonality: April-October. Tenerals: May-September; mostly July-August. Adult food: chrysomelid pupae (in the field); lepidopterous eggs and larvae (in captivity). Predators: toads (regularly). Dispersal power. Macropterous. Frequent flier (sometimes swarming to artificial lights at night). Moderate runner. Occasional climber (on plants). Strong burrower. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Pitfall trapping; sweeping plants. References. Lindroth, 1968a and 1971a (natural history; as funerarius Csiki); Kirk, 1975c (ecology, dispersal power; as funerarius Csiki); Noonan, 1991 (natural history; as funerarius Csiki). Harpalus rewolinskii Noonan, 1991, see Harpalus somnulentus Dejean, 1829. Reference. Ball and Bousquet, 2001 (taxonomy). Harpalus (Harpalus) rubripes (Duftschmid, 1812) Ecology. Lowlands. Cultivated fields (e.g., hay), vacant lots, and roadsides. Europe: sandy pastures and fields, gravel pits, fallow fields, and ruderal places. Open ground; dry, sandy soil covered with sparse vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in the soil and under stones. Biology. Seasonality: April-September. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Frequent in seashore drift material, which indicates flight ability. Moderate runner. Strong burrower. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; turning drift material. References. Lindroth, 1945 and 1986 (natural history); Krinsky and Oliver, 1988 (ecology); Koch, 1989 (ecology).

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Harpalus (Ophonus) rufibarbis (Fabricius, 1792), see Ophonus rufibarbis (Fabricius, 1792). References. Kataev, 1996 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Harpalus (Pseudoophonus) rufipes (DeGeer, 1774) Ecology. Lowlands and mountains. Cultivated fields (e.g., potato, corn, hay, oat, strawberry, dry bean), pastures, meadows, old fields, vacant lots, roadsides, parks, lawns, courts, yards, sand dunes, gravel pits, dumps, orchards, forest edges, and open forests. Greenhouses and human habitations (occasionally). Open ground; moderately dry, clay soil covered with some vegetation (e.g., grass, weeds). Mostly nocturnal, with an activity peak after midnight; sometimes active in the sunshine; usually sheltering during the day under stones, pieces of wood, heaps of hay, pieces of cardboard, and in burrows dug in the soil. Semi-gregarious. First and second instar larvae are active on the ground surface while third instar larvae remain in their burrows to feed and complete their development. Biology. Seasonality: April-August, October. Copulating pairs: July-August (QU). Gravids: early July to mid-August (NB, QU). Tenerals: July-August. Adult food: strawberries and lepidopterous caterpillars (North America), seeds of cereals, strawberries, and conifers (Europe) (in the field); wheat kernels (in captivity). Larval food, in captivity: seeds of Chenopodium album (Linné). Economic importance: sometimes a pest of strawberries. Predators: foxes. Adults often crushed by cars and pedestrians. Defense-mechanism: when alarmed, the adult may emit a spray from the pygidial glands. Dispersal power. Macropterous. Frequent flier (mostly tenerals; sometimes swarming to artificial lights at night). Frequent in lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Dispersal by flight occurs early in the season and dispersal by running on the ground surface, later in the season. Clearly effective as a colonist. Strongly favored by human activities. Sometimes invading homes in large numbers, to the point of becoming a nuisance. Collecting techniques. Pitfall trapping; light trapping; turning stones or pieces of wood; turning drift material; collecting at night around street lights; sweeping plants. References. Morrison, 1941 (ecology, biology); Lindroth, 1945, 1955a, 1963a, and 1968 (natural history); Briggs, 1965 (biology); Greenslade, 1965 (biology); Lindroth, 1986 (natural history); Wallin and Ekblom, 1988 (dispersal power); Koch, 1989 (ecology, biology); Dearborne and Donahue, 1993 (biology); Luff, 1993 (biology); Zhang et al. 1997a, 1997b, and 1998 (biology, dispersal power); Hartke et al. 1998 (biology, dispersal power). Harpalus solitaris Dejean, 1829 (Harpalus fuliginosus (Duftschmidt, 1812)) Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, meadows, vacant lots, abandoned fields, sand pits, gravel pits, roadsides, thickets (e.g., Alnus), and open forests. Open ground; dry sandy soil covered with sparse low vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under stones (mostly), dead leaves, pieces of wood and cardboard. Biology. Seasonality: April-

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August. Tenerals: late June to August. Predators: toads. Dispersal power. Macropterous. Occasional flier (to artificial lights in captivity). Frequent in seashore drift material, which indicates flight activity. Moderate runner. Collecting techniques. Turning stones; pitfall trapping; turning drift material. References. Lindroth, 1945, 1955a, 1963a, 1968 (as fuliginosus Duftschmid), and 1986 (natural history); Koch, 1989 (ecology); Noonan, 1991 (biology; as fuliginosus Duftschmid). Harpalus somnulentus Dejean, 1829 (Harpalus pleuriticus Kirby, 1837; Harpalus fallax LeConte, 1859) Ecology. Lowlands, mountains, subalpine and alpine zones. Grasslands, relict prairies, meadows, rangelands, pastures, cultivated fields and their borders (e.g., corn, wheat, barley, asparagus, cabbage, flax, raspberry), moraines, vacant lots, ski fields, hills, roadsides, sand pits, gravel pits, hedges, thickets (e.g., Alnus), forest edges and clearings, open or light forests, and vicinity of human habitations. Ground open (mostly) or shaded by bushes (e.g., Alnus), shrubs or trees; rather firm, well-drained, moderately dry, often gravelly or sandy soil covered with some vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under stones, dead leaves, heaps of hay, reindeer moss (Cladonia), pieces of wood, bark, logs, pieces of cardboard and paper. Biology. Seasonality: March-December. Copulating pairs: July (AB, QU). Gravids: mid-July to August (AB). Tenerals: March-December, mostly July-August. Adults found overwintering on higher and drier ground, in vacant lots, pastures, on hills, in cultivated fields, in the upper zone of sand pits, at the edge of woods, and in woods; under stones and dead leaves (mostly), in the soil, under plant rosettes (Verbascum), heaps of hay, and in rotten tree-stumps. Adult food, in the field: lepidopterous caterpillars and grasshopper eggs. Predators: toads (frequently), frogs, woodpeckers (flickers), robins, and foxes. Parasites: dipterous larvae and Laboulbeniales (Fungi). Dispersal power. Macropterous. Occasional flier (to lights in captivity; into interception traps). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning stones; turning drift material; using interception traps; sweeping plants. References. Lindroth, 1955a (as pleuriticus Kirby), 1963a (as pleuriticus Kirby), and 1968 (natural history; as somnulentus Dejean, pleuriticus Kirky, carbonatus LeConte, intactus Casey, uteanus Casey, fallax LeConte); Frank, 1971b (biology; as pleuriticus Kirby); Levesque et al. 1979 (biology; as fallax LeConte); Reeves et al. 1983 (biology; as pleuriticus Kirby); Levesque and Levesque, 1987 (as pleuriticus Kirby) and 1994 (biology); Noonan, 1991 (taxonomy; natural history); Kavanaugh, 1992 (natural history); Ball and Bousquet, 2001 (taxonomy). Harpalus (Euharpalops) spadiceus Dejean, 1829 (Harpalus carolinae Schaeffer, 1910) Ecology. Lowlands and mountains. Deciduous, mixed and coniferous forests: often along gorges. Ground shaded by trees or shrubs; rather dry soil covered with thick leaf litter. Nocturnal; sheltering during the day in leaf litter, under pieces of wood, and under

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stones. Biology. Seasonality: January-October, November. Tenerals: March-October; mostly June and September. Adults found overwintering (SC). Adult food, in the field: lepidopterous caterpillars. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Barr, 1969 (ecology; as carolinae Schaeffer); Kirk, 1970 (biology); Erwin, 1981 (natural history); Noonan, 1991 (natural history). Harpalus (Glanodes) stephani Ball, 1972 Ecology. Uplands. A grassland (in desert). Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Reference. Ball, 1972 (ecology). Harpalus (Opadius) tadorcus Ball, 1972 Ecology. Uplands. Grasslands, pastures, roadsides; borders of ponds and streams. Open ground; dry soil composed of sandy silt or loam (sometimes gravel-mixed), covered with sparse vegetation (e.g., grass). Nocturnal; sheltering during the day under dry cow dung or stones, and in burrows dug in the soil. Biology. Seasonality: June-September; mostly August. Tenerals: June; August-September (mostly). Adult food, in the field: grass seeds. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on plants). Collecting techniques. Pitfall trapping; turning dry cow dung; sweeping plants. Reference. Noonan, 1991 (natural history). Harpalus (Pseudoophonus) texanus Casey, 1914 Ecology. Lowlands. A scrub oak area, under stones (TX). Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Ball and Anderson, 1962 (ecology). Harpalus (Euharpalops) ventralis LeConte, 1848 Ecology. Uplands. Grasslands, rangelands, pastures, cultivated fields (e.g., corn, wheat), old fields, roadsides, gravel pits, hills, and open forests; borders of marshes and sloughs. Open ground; dry, sandy or silty soil, with some vegetation (e.g., grass). Nocturnal; sheltering during the day under dry cow dung, stones, logs, and fallen dead plants. Biology. Seasonality: April-November. Tenerals: June-August. Dispersal power. Submacropterous. Occasional flier (to artificial lights at night). Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning dry cow dung. References. Lindroth, 1968 (natural history); Kirk, 1975c (ecology, dispersal power); Noonan, 1991 (natural history). Harpalus vittatus alaskensis Lindroth, 1968 Ecology. Unknown. Biology. Seasonality: June. Dispersal power. Macropterous, probably capable of flight. Reference. Kataev, 1990 (taxonomy).

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Hartonymus alternatus (LeConte, 1863) Ecology. Uplands. Sand dunes. At a distance from water. Open ground; sandy soil. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Hartonymus hoodi Casey, 1914 Ecology. Lowlands. Sand bars and sand dunes: along streams. Open ground; sandy soil. Nocturnal; sheltering during the day under pieces of wood and other debris. Biology. Seasonality: March-May, October. Tenerals: October (IL). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; turning debris. Reference. Hood, 1952 (ecology). Helluomorphoides clairvillei (Dejean, 1831) Ecology. Lowlands. Forests (e.g., Pinus). Sand hills (occasionally). Nocturnal; sheltering during the day under pieces of wood. Biology. Seasonality: January-March, May-August, November. Adult and larval food, in captivity: mealworms. Defense-mechanism: when alarmed, the adult may emit an acrid repugnant spray from the pygidial glands. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Occasional climber. Collecting techniques. Pitfall trapping; turning pieces of wood; baiting traps with turpentine. References. Kirk, 1970 (dispersal power); Attygalle et al. 1992 (biology). Helluomorphoides ferrugineus (LeConte, 1853) Ecology. Lowlands and mountains. Open forests (e.g., Pinus). Nocturnal; sheltering during the day under logs and rotten pieces of wood. Biology. Seasonality: March, June, August-September. Tenerals: late July (TX). Adult food: adult Army ants (in the field); all development stages of Army ants (in captivity). Adult beetles follow columns of Army ants in search of food. Defense-mechanism: when alarmed, the adult may emit an acrid repugnant spray from the pygidial glands. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; turning logs. References. Leng, 1915 (ecology); Eisner et al. 1968 (biology); Topoff, 1969 (biology). Helluomorphoides latitarsis (Casey, 1913) Ecology. Mountains. Open forests (e.g., Pinus). Biology. Seasonality: August-September. Adult food: larvae and pupae of Army ants (in the field); all development stages of Army ants (in captivity). Adult beetles follow columns of Army ants in search of food. Defense-mechanism: when alarmed, the adult may emit an acrid repugnant spray from the pygidial glands. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pitfall trapping. References. Eisner et al. 1968 (biology); Topoff, 1969 (biology).

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Helluomorphoides nigripennis (Dejean, 1831) Ecology. Lowlands. Deciduous forests (e.g., Quercus), low wet situations (e.g., swamps), and vicinity of cultivated fields (e.g., cotton). Wet or moist soil. Nocturnal; sheltering during the day in leaf litter and in clumps of broomsedges. Biology. Seasonality: January-September, November-December. Adults found overwintering in wood debris. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter; using tanglefoot screens. References. Löding, 1945 (ecology); Kirk, 1969 (natural history). Helluomorphoides papago (Casey, 1913) Ecology. Mountains. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Helluomorphoides praeustus bicolor (T.W. Harris, 1828) Ecology. Uplands. Forests. Dry soil. Nocturnal; sheltering during the day under stones and logs. Biology. Seasonality: March-September. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; turning stones and logs. References. Blatchley, 1910 (ecology); Kirk, 1969 and 1975c (dispersal power). Helluomorphoides praeustus floridanus Ball, 1956 Ecology. Lowlands. Biology. Seasonality: March. Dispersal power. Macropterous, probably capable of flight. Helluomorphoides praeustus praeustus (Dejean, 1825) Ecology. Lowlands. Biology. Seasonality: June. Dispersal power. Macropterous, probably capable of flight. Helluomorphoides texanus (LeConte, 1853) Ecology. Lowlands and mountains. Forests. Biology. Seasonality: July. Adult food: adult Army ants (in the field); all development stages of Army ants (in captivity). Adult beetles follow columns of Army ants in search of food. Defense-mechanism: when alarmed, the adult may emit a spray from the pygidial glands. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Pitfall trapping. Reference. Plsek et al. 1969 (biology).

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Horologion speokoites Valentine, 1932 Ecology. A cave. Wet soil. Occurring on small pieces of wet, rotten wood. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Hyboptera dilutior Oberthür, 1884 Natural History. Unknown. Reference. Ball and Bousquet, 2001 (taxonomy). Infernophilus castaneus (G.H. Horn, 1882) Ecology. Deserts. Occurring on the ground, under the leaves of decaying yucca plants, or active on low vegetation. Biology. Seasonality: May and July. Tenerals: early July (CA). Dispersal power. Macropterous, probably capable of flight. Climber. Reference. Larson, 1969 (biology, dispersal power). Inna boyeri (Solier, 1835), see Eucheila (Inna) boyeri (Solier, 1835). Reference. Shpeley and Ball, 2000 (taxonomy). Lachnocrepis parallela (Say, 1830) Ecology. Lowlands and uplands. Borders of eutrophic marshes, ponds, lakes, and swamps. Somewhat shaded ground; soil inundated, covered with dense tall vegetation (e.g., Typha), or, when non-inundated, soft, very wet, muddy, rich in organic debris, and covered with similar vegetation. Diurnal and nocturnal; active under water along plant stems or sheltering during the day among floating vegetal debris (e.g., dead Typha-stems). Biology. Seasonality: January, March-December. Gravids: late June (VT). Tenerals: July-August, October. Adults found overwintering on higher and drier ground, in the upper banks of marshes, on hillocks, in vacant lots and hedges, at the edge of woods, and in woods; in the soil (as deep as 15 cm), under stones, logs, dead leaves, pieces of fallen bark, under the loose bark of fallen trees, in rotten logs, and in clumps of moss. Adult food, in the field: plant seeds. Predators: robins. Phoretic mites found on adult body. Defense-mechanism: when alarmed, the adult escapes by swimming and diving into the water (able to remain submerged at the base of Typha-stems for about 20 minutes). Dispersal power. Macropterous. Regular flier (to artificial lights at night; to artifical lights in captivity). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Frequent climber (on plants, e.g., Typha). Excellent swimmer and diver. Collecting techniques. Treading the dead floating vegetation into the water; sweeping the vegetation; light trapping. References. Ulke, 1903 (ecology); Lindroth, 1969a (natural history); Landry, 1976 (biology); Bousquet, 1996a (taxonomy, ecology).

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Lachnophorus elegantulus Mannerheim, 1843 Ecology. Lowlands and mountains. Margins of rivers and brooks. Open ground; moist or wet, gravelly soil. Diurnal; active in the sunshine. Biology. Seasonality: April, August. Tenerals: August (TX). Defense-mechanism: body color-pattern matching the soil. Dispersal power. Wing-condition unknown. Fast runner. Collecting technique. Hunting with a butterfly net. Reference. Ball and Bousquet, 2001 (ecology). Laemostenus (Laemostenus) complanatus (Dejean, 1828) Ecology. Lowlands. Ports, human habitations (e.g., cellars), greenhouses, gardens, stables, alleys, fields, and sandy epilittoral zone. Shaded or open ground; moderately moist soil, bare or covered with some vegetation. Nocturnal; sheltering during the day under pieces of wood. Biology. Seasonality: January-March, May, July, December. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Clearly effective as a colonist. Strongly favored by human activities. Collecting technique.Turning pieces of wood. Reference. Lindroth, 1966 (natural history). Laemostenus (Pristonychus) terricola terricola (Herbst, 1784) Ecology. Lowlands and mountains. Human habitations (e.g., cellars), stables, barns, and forests. Also nests of mammals and caves (Europe). Shaded or open ground; moderately moist soil, bare or covered with some vegetation. Nocturnal; sheltering during the day under pieces of wood and stored vegetables. Gregarious. Biology. Seasonality: April-November. Tenerals: July, September (NF). Adult food, in the field (Europe): insects, isopods, and guano. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Clearly effective as a colonist. Strongly favored by human activities. Collecting technique. Turning boards and vegetables stored in house cellars. References. Lindroth, 1945, 1955a, 1963a, 1966, and 1986 (natural history); Koch, 1989 (ecology). Lebia (Lebia) abdita Madge, 1967 Ecology. Mountains. Biology. Seasonality: August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Lebia (Lebia) abdominalis Chaudoir, 1843 Ecology. Lowlands. Pastures and cultivated fields (e.g., cotton, soybean, corn, tobacco). Open (mostly) or slightly shaded ground; rather dry soil. Mostly diurnal; active in the sunshine on plants (mostly) and trees; sheltering on cloudy or cool days under plant rosettes (Verbascum) and debris. Biology. Seasonality: January-July, November. Adults found overwintering in woods; in clumps of broomsedges and under debris. Predators: blackbirds. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner.

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Frequent climber (on plants and trees). Strongly favored by human activities. Collecting techniques. Sweeping or beating the vegetation. Lebia (Lebia) analis Dejean, 1825 Ecology. Lowlands and mountains. Abandoned fields, cultivated fields (e.g., soybean (mostly), oat, cotton, alfalfa, lespedeza, corn, cowpea, gardenpea, vetch, mustard green), roadsides, and orchards (e.g., plum, peach). Open ground; rather dry soil. Mostly diurnal and crepuscular; active in the sunshine on plants (mostly), bushes, trees, and on the ground. Adults descend to the lower parts of plants and to the ground during the hottest part of the day. Associated with chrysomelids (Capraita obsidiana (Fabricius), Disonycha glabrata Fabricius), which is a case of Batesian mimicry. Biology. Seasonality: January-November. Adults found overwintering in woods; in clumps of broomsedges, under dead leaves, pieces of wood, and in tree hollows. Adult food: lepidopterous immature stages (mostly), nabid and pentatomid nymphs (in the field); lepidopterous immature stages (e.g., eggs of Heliothis zea (Boddie)), adult cicadellids, and chrysomelid larvae (in captivity). Economic importance: may play an important role in the control of Bollworm eggs on cotton. Predators: ducks. Dispersal power. Macropterous. Frequent flier (in the sunshine; to artificial lights at night). Moderate runner. Frequent climber (on plants, bushes, and trees). Strongly favored by human activities. Collecting techniques. Pitfall trapping; sweeping or beating the vegetation; light trapping; sifting dead leaves. References. Blatchley, 1910 (ecology); Whitcomb and Bell, 1960 (biology); Hemenway and Whitcomb, 1967 (ecology, biology); Kirk, 1969 (biology, dispersal power) and 1975c (dispersal power); Lindroth, 1971b (ecology, biology); Erwin, 1981 (natural history). Lebia (Lebia) arizonica Schaeffer, 1910 Ecology. Mountains. Biology. Seasonality: April. Dispersal power. Macropterous, probably capable of flight. Lebia (Loxopeza) atriceps LeConte, 1863 Ecology. Uplands. Biology. Seasonality: May-October. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Reference. Kirk, 1975c (dispersal power). Lebia (Loxopeza) atriventris Say, 1823 Ecology. Lowlands and mountains. Abandoned fields, meadows, grasslands, cultivated fields (e.g., cotton, corn, soybean, alfalfa), roadsides, forest edges and clearings, and open forests. Open or slightly shaded ground; rather dry soil. Mostly diurnal; active in the sunshine on plants (e.g., Solidago (mostly), Heterotheca, Ambrosia, Gossypium, Zea, Chenopodium, Rumex, grass) and trees; sheltering on cloudy or cool days under stones and in leaf litter. Associated with chrysomelids (Zygogramma heterothecae Lameere). Biology. Seasonality: January-December. Gravids: mid-June and early July (QU). Tener-

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als: June-August. Adults found overwintering on higher and drier ground, in vacant lots, on hillocks, in roadsides, at the edge of woods, in clearings and woods; under stones and dead leaves (mostly), logs, pieces of wood, moss, clumps of grass, under the loose bark of tree-stumps, logs, and fallen trees, and under plant rosettes (Verbascum). Adult food, in the field: immature stages of chrysomelids (Leptinotarsa, Zygogramma), fungi, pollen and anthers of weeds. Predators: toads and frogs. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Frequent in lakeshore drift material, which indicates flight ability. Moderate runner. Frequent climber (mostly on plants; also on trees). Strongly favored by human activities. Collecting techniques. Sweeping or beating the vegetation; pitfall trapping; light trapping; turning drift material. References. Forbes, 1880 (biology); Smith, 1882 (ecology); Lovell, 1915 (ecology); Lindroth, 1969a (natural history); Kirk, 1969, 1970, and 1975c (dispersal power); Erwin,1981 (natural history); Will et al. 1995 (dispersal power). Lebia (Lebia) bilineata Motschulsky, 1859 Ecology Uplands. Biology. Seasonality: March, June-August. Dispersal power. Macropterous, probably capable of flight. Lebia (Lebia) bitaeniata Chevrolat, 1834 Ecology. Lowlands. On shrubs and weeds. Biology. Seasonality: April-June. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Sweeping or beating the vegetation. Lebia (Lebia) bivittata (Fabricius, 1798) Ecology. Lowlands and mountains. Cultivated fields (e.g., corn, potato) and open forests. Open ground. Mostly diurnal; active in the sunshine on plants; sheltering on cloudy or cool days at the base of plants and under debris. Biology. Seasonality: January, MarchAugust, October, December. Adults found overwintering under pieces of wood and other debris. Dispersal power. Macropterous. Occasional in seashore drift material,which indicates flight ability. Moderate runner. Climber (on plants). Collecting techniques. Pitfall trapping; sweeping plants. Lebia (Lebia) bumeliae Schaeffer, 1910 Ecology. In an oak scrub, on bromeliads. Biology. Seasonality: April-May, July, December. Dispersal power. Macropterous, probably capable of flight. Lebia (Lebia) calliope Bates, 1883 Ecology. Unknown. Biology. Seasonality: January, March, May, July. Dispersal power. Macropterous, probably capable of flight.

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Lebia (Lebia) collaris Dejean, 1826 Ecology. Lowlands. In a deciduous forest (Quercus), on Spanish moss (growing on trees) and on plants (e.g., Solidago). Biology. Seasonality: January-November. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Climber (on trees and plants). Collecting technique. Sweeping or beating the vegetation. Reference. Leng, 1915 (ecology). Lebia (Lebia) cyanipennis Dejean, 1831 Ecology. Lowlands and mountains. Macrohabitat unknown. On plant leaves (Artemisia). Biology. Seasonality: April-June, August-September. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Frequent climber (on plants). Lebia (Loxopeza) deceptrix Madge, 1967 Ecology. Uplands. Biology. Seasonality: July-August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Lebia (Lamprias) divisa LeConte, 1850 Ecology. Edge of a slough, on weeds. Biology. Seasonality: April, July. Dispersal power. Macropterous, probably capable of flight. Regular climber (on plants). Collecting technique. Sweeping plants. Lebia (Lebia) esurialis Casey, 1920 Ecology. Unknown. Biology. Seasonality: May-August. Dispersal power. Macropterous, probably capable of flight. Lebia (Lebia) fuscata Dejean, 1825 Ecology. Lowlands and mountains. Light or open forests, their edges and clearings, and adjacent meadows, pastures, vacant lots, sand pits, gravel pits, roadsides, and ski fields. On shaded or open ground; moderately moist soil. Mostly diurnal; active in the sunshine on trees (e.g., Amelanchier, Betula, Carya, Ostrya, Picea, Pinus, Populus, Prunus, Quercus, Tilia,Tsuga,Ulmus), flowering bushes and shrubs (e.g., Alnus, Clematis, Crataegus, Physocarpus, Rhododendron, Rhus, Rubus, Salix, Viburnum, Vitis), and flowering plants (e.g., Solidago, Spiraea, Taraxacum). Occurring preferably on Populus, Salix, and Solidago. Sheltering on cloudy or cool days under dead leaves. Biology. Seasonality: January, March-September, November-December. Copulating pairs: May (QU). Tenerals: JulyAugust. Adults found overwintering on higher and drier ground, in woods, at the edge of woods, and on hillocks; in leaf litter (mostly), under the loose bark of tree trunks and stumps, and in the soil. Adult food, in the field: chrysomelid pupae and tenerals (Galerucella cavicollis LeConte, Trirhabda borealis Blake, T. virgata LeConte). Predators: toads.

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Dispersal power. Macropterous. Frequent flier (to artificial lights at night; into rotary traps). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (on trees, shrubs, bushes, and plants). Collecting techniques. Sweeping or beating the vegetation; light trapping; using rotary traps. References. Lovell, 1915 (ecology); Cushman and Isely, 1916 (biology); Hemenway and Whitcomb, 1967 (ecology, biology); Kirk, 1970 (dispersal power); Larochelle, 1972a (ecology). Lebia (Loxopeza) grandis Hentz, 1830 Ecology. Lowlands and mountains. Abandoned fields, cultivated fields (e.g., potato, corn, soybean), orchards (e.g., peach), and open forests. Open ground; rather dry soil. Mostly diurnal; active in the sunshine on plants (e.g., Solanum, Zea, Solidago, Asclepias) and trees (e.g., Pinus); sheltering on cloudy or cool days under stones and dead leaves. Associated with chrysomelids (Leptinotarsa). Biology. Seasonality: January-December. Eggs are laid singly in the soil and covered with particles of sand or humus. Adults found overwintering on higher and drier ground, at the edge of woods and in woods; under logs, pieces of wood, stones, dead leaves, and the loose bark of dead trees. Adult food: chrysomelid eggs, larvae, and pupae (Leptinotarsa decemlineata (Say)); eggs and larvae of soft-bodied insects, and young worms (in the field); chrysomelid immature stages (Leptinotarsa, Crioceris) (in captivity). The larva is an ectoparasite of chrysomelid larvae and pupae (Leptinotarsa). Economic importance: plays an important role in the control of the Colorada Potato Beetle (Leptinotarsa). Predators: crows, flycatchers, and starlings. Defense-mechanism: when alarmed, the adult may emit a Brachinus-like smoke from the pygidial glands. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (on plants and trees). Strongly favored by human activities. Collecting techniques. Sweeping or beating the vegetation; light trapping; sugaring tree trunks; using tanglefoot screens. References. Riley, 1871 (biology); Smith, 1882 (ecology); Blatchley, 1910 (ecology, biology); Chaboussou, 1938 and 1939 (biology); Hemenway and Whitcomb, 1967 (ecology, biology); Kirk, 1969 and 1970 (dispersal power); Lindroth, 1969a (natural history); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Lebia (Lebia) guttula LeConte, 1851 Ecology. Uplands. Forest edges and vineyards. Rather dry soil. Diurnal; active in the sunshine on trees and plants. Gregarious (in winter). Biology. Seasonality: January-August, October-December. Adults found overwintering near fields; in moss and lichens growing on trees (Quercus) and under the loose bark of trees (Eucalyptus). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Frequent climber (on trees and plants). Collecting technique. Sweeping or beating the vegetation. Lebia (Lebia) histrionica Bates, 1883 Ecology. Mountains. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight.

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Lebia (Lebia) insulata Madge, 1967 Ecology. Unknown. Biology. Seasonality: May, August. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Lebia (Polycheloma) lecontei Madge, 1967 Ecology. Unknown. Biology. Seasonality: March, May. Dispersal power. Macropterous, probably capable of flight. Lebia (Lebia) lecta G.H. Horn, 1885 Ecology. Lowlands. Borders of an hillock, in the curled leaves of cut branches (FL). Biology. Seasonality: March. Dispersal power. Macropterous, probably capable of flight. Reference. Leng, 1915 (ecology). Lebia (Lebia) lobulata LeConte, 1863 Ecology. Lowlands and mountains. Abandoned fields, gardens, forest edges, light or open forests (e.g., Quercus, Carya). Open or slightly shaded ground; rather dry soil covered with some vegetation (e.g., weeds). Mostly diurnal; active in the sunshine on plants (e.g., weeds); sheltering on cloudy or cool days in leaf litter. Gregarious. Biology. Seasonality: FebruaryNovember. Adults found overwintering under the loose bark of trees (e.g., Carya) and among vegetal debris. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Climber (on plants). Collecting techniques. Sweeping plants; sifting leaf litter. References. Leng, 1915 (ecology); Löding, 1945 (biology); Erwin, 1981 (natural history). Lebia (Lebia) marginicollis Dejean, 1825 Ecology. Lowlands and mountains. Grassy meadows, cultivated fields (e.g., cotton, corn, alfalfa, soybean), roadsides, hillocks in marshy areas, light or open forests. Open or slightly shaded ground; rather dry soil covered with some vegetation (e.g., grass, weeds). Diurnal; active in the sunshine on plants (mostly), bushes, low shrubs, and trees (e.g., Quercus, Roystonea). Biology. Seasonality: January-November. Adults found overwintering under the loose bark of dead trees (Pinus). Adult food, in the field: small insects. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Frequent climber (on plants, bushes, and trees). Strongly favored by human activities. Collecting techniques. Sweeping or beating the vegetation; pitfall trapping. References. Leng, 1915 (ecology); Kirk, 1970 (dispersal power). Lebia (Lebia) miranda (G.H. Horn, 1872) Ecology. Mountains. Biology. Seasonality: June-August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night).

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Lebia (Lebia) moesta LeConte, 1850 Ecology. Lowlands, mountains, subalpine and alpine zones. Meadows, abandoned fields, roadsides, ski fields, forest edges and clearings, and light forests. Diurnal; active in the sunshine on flowering plants and shrubs (mostly Cornus and Viburnum; also Taraxacum, Kalmia, Salix, Myrica), and trees (e.g., Ulmus). Biology. Seasonality: March-September, November. Tenerals: late August (SA). Adults found overwintering on higher and drier ground, at the edge of woods and on hillocks; in leaf litter. Dispersal power. Macropterous. Occasional flier (to artificial lights at night; into rotary traps). Moderate runner. Frequent climber (on plants, shrubs, and trees). Collecting techniques. Sweeping and beating the vegetation; using rotary traps. References. Lovell, 1915 (ecology); Lindroth, 1969a (natural history). Lebia (Lebia) nigricapitata Madge, 1967 Ecology. Mountains. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Lebia (Lebia) ornata Say, 1823 Ecology. Lowlands, mountains, subalpine and alpine zones. Light and open forests (mostly deciduous; mixed), their clearings, edges, roadsides, and adjacent ski fields; hedges, wooded river banks, and swamps. Open or slightly shaded ground; moderately moist soil. Mostly diurnal; active in the sunshine on trees, shrubs, and plants (mostly Solidago, Prunus, and Salix; also Acer, Amelanchier, Betula, Juglans, Ostrya, Prunus, Quercus, Tilia, Ulmus, Aronia, Ceanothus, Cornus, Corylus, Crataegus, Rhus, Rubus, Viburnum, Eupatorium, Melilotus), especially when they are flowering; sheltering on cloudy or cool days in leaf litter at the base of trees. Gregarious (in winter). Biology. Seasonality: February-December. Copulating pairs: May (QU). Tenerals: May (rarely), July-September. Adults found overwintering in woods and at the edge of woods; among dead leaves, under stones and logs. Adult food, in captivity: chrysomelid larvae and pupae (Haltica chalybea Illiger). Predators: toads, grouses, and ducks. Dispersal power. Macropterous. Frequent flier (in the bright sunshine; to artificial lights at night). Moderate runner. Frequent climber (on trees, shrubs, and plants). Collecting techniques. Sweeping or beating the vegetation; light trapping; baiting traps with carrion. References. Smith, 1882 (ecology); Blatchley, 1910 (ecology); Lovell, 1915 (ecology); Cushman and Isely, 1916 (biology); Isely, 1920 (biology); Madge, 1967 (biology); Kirk, 1969 (natural history) and 1970 (ecology, dispersal power); Lindroth, 1969a (natural history); Larochelle, 1974a (ecology); Erwin, 1981 (natural history). Lebia (Lebia) pectita G.H. Horn, 1885 Ecology. Lowlands. Meadows. Open ground; rather dry soil. Diurnal; active in the sunshine on flowering plants (e.g., Solidago, Achillea). Gregarious. Biology. Seasonality: April-

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October. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Regular climber (on plants). Collecting technique. Sweeping plants. References. Lovell, 1915 (ecology); Erwin, 1981 (natural history). Lebia (Lebia) perita Casey, 1920 Ecology. Uplands. Biology. Seasonality: June-September. Tenerals: September (BC). Dispersal power. Macropterous, probably capable of flight. Reference. Lindroth, 1969a (biology, dispersal power). Lebia (Lebia) perpallida Madge, 1967 Ecology. Mountains. Biology. Seasonality: August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Lebia (Loxopeza) pimalis (Casey, 1920) Ecology. Mountains. Biology. Seasonality: June, September. Dispersal power. Macropterous, probably capable of flight. Lebia (Lebia) pleuritica LeConte, 1848 Ecology. Lowlands. Meadows, city parks, and open forests. Open or slightly shaded ground; rather dry soil. Mostly diurnal; active in the sunshine on flowering plants (e.g., Solidago) and trees (e.g., Quercus); sheltering on cloudy or cool days in leaf litter. Biology. Seasonality: March-October. Adults found overwintering in leaf litter. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Frequent climber (on plants and trees). Collecting techniques. Sweeping and beating the vegetation; light trapping; sifting leaf litter. Reference. Kirk, 1975c (ecology, dispersal power). Lebia (Lebia) pulchella Dejean, 1826 Ecology. Lowlands and mountains. Abandoned fields, pastures, cultivated fields (e.g., soybean, cotton, corn, alfalfa, oat), and forest edges. Open ground; rather dry soil. Mostly diurnal; active in the sunshine on plants, especially when they are flowering; sheltering on cloudy or cool days under stones, in leaf litter, and in clumps of broomsedges. Biology. Seasonality: March-October. Adults found overwintering in pastures and woods; in clumps of broomsedges, among pieces of wood, and under the loose bark of trees. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (on plants). Strongly favored by human activities. Collecting techniques. Sweeping plants; pitfall trapping. References. Löding, 1945 (biology); Hemenway and Whitcomb, 1967 (ecology); Kirk, 1969 (biology, dispersal power) and 1970 (dispersal power).

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Lebia (Lebia) pumila Dejean, 1831 Ecology. Lowlands and mountains. Meadows, weedy fields, cultivated fields (e.g., cotton, alfalfa), vacant lots, sand pits, gravel pits, roadsides, forest paths and clearings. Open ground; rather dry soil. Mostly diurnal; active in the sunshine on flowering plants (e.g., Achillea, Apocynum, Asclepias, Aster, Chamaerops, Eupatorium, Gossypium, Origanum, Pastinacea, Poa, Solidago (mostly)), bushes (e.g., Rubus, Spiraea, Vitis), and trees (e.g., Acer, Celtis, Pinus); sheltering on cloudy or cool days mostly under dead leaves (e.g., under Alnus-bushes), also under stones, in the soil at the base of plants (e.g., Rumex), and under reindeer moss (Cladonia). Associated with chrysomelids (Lexiphanes saponatus (Fabricius)). Gregarious. Biology. Seasonality: January-November. Gravids: late June (QU). Tenerals: May (rarely), July-August. Adults found overwintering on higher and drier ground, in sand pits and woods; in leaf litter. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; into rotary traps). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Frequent climber (on plants, bushes, and trees). Collecting techniques. Sweeping and beating the vegetation; light trapping; using rotary traps; raking the leaf litter; pitfall trapping; using an aspirator. References. Smith, 1882 (ecology); Blatchley, 1910 (ecology; as rhodopus Schwarz); Lovell, 1915 (ecology); Lindroth, 1969a (natural history); Kirk, 1975c (ecology, dispersal power); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Lebia (Lebia) rufopleura Schaeffer, 1910 Ecology. Uplands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Lebia (Lebia) scalpta Bates, 1883 Ecology. Mountains. Biology. Seasonality: March, May-August, October. Dispersal power. Macropterous, probably capable of flight. Lebia (Lebia) scapula G.H. Horn, 1885 Ecology. Mountains. Biology. Seasonality: June-August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Lebia (Lebia) solea Hentz, 1830 Ecology. Lowlands and mountains. Open or light deciduous forests; forest roads, clearings, and edges; orchards (plum, peach), meadows, pastures, vacant lots, and cultivated fields (e.g., corn, hay). Slightly shaded or open ground; moderately moist soil. Mostly diurnal; active in the sunshine on the flowers of plants (e.g., Anthriscus, Artemisia, Chenopodium, Solidago caesia Linné (mostly), Taraxacum), bushes (e.g., Alnus, Ceanothus, Rubus, Sambucus), and trees (e.g., Abies, Carya, Liriodendron, Ostrya, Prunus, Quercus,

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Ulmus), or, on foliage (e.g., corn, hay); sheltering on cloudy or cool days in leaf litter and at the base of plants. Biology. Seasonality: January, March-November. Tenerals: August (MD). Adults found overwintering in woods and at the edge of woods; in leaf litter. Adult and larval food, in the field: chrysomelid immature stages (Xanthogalerucella luteola (Müller)). The larva is ectoparasitic on chrysomelid pupae. Defense-mechanism: when alarmed, the adult escapes by flying. Dispersal power. Macropterous. Frequent flier (when alarmed; in the sunshine; to artificial lights at night; into rotary traps; into window traps). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Frequent climber (on plants, bushes, shrubs, and trees). Collecting techniques. Sweeping and beating the vegetation; light trapping; using rotary traps; using window traps. References. Blatchley, 1910 (ecology); Lindroth, 1969a (dispersal power); Kirk, 1970 (dispersal power) and 1975c (ecology, dispersal power); Larochelle, 1974a (ecology); Erwin, 1981 (natural history). Lebia (Loxopeza) subdola Madge, 1967 Ecology. Mountains. Biology. Seasonality: June-August. Dispersal power. Macropterous, probably capable of flight. Lebia (Loxopeza) subgrandis Madge, 1967 Ecology. Mountains. Biology. Seasonality: August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Lebia (Lebia) subrugosa Chaudoir, 1871 Ecology. Mountains. Biology. Seasonality: June-August. Dispersal power. Macropterous, probably capable of flight. Lebia (Loxopeza) tricolor Say, 1823 Ecology. Lowlands and mountains. Light or open forests (mostly deciduous), their edges and clearings; thickets, orchards (e.g., peach), hedges (Crataegus), meadows, and abandoned fields. Slightly shaded ground; rather dry soil. Mostly diurnal; active in the sunshine on the flowers of plants (e.g., Echinocystus, Solidago (mostly)), bushes (e.g., Cornus, Spiraea, Salix), and trees (e.g., Malus), or, on the foliage of trees (e.g., Ostrya, Populus, Ulmus); sheltering on cloudy or cool days under dead leaves and stones. Biology. Seasonality: January, March-October. Tenerals: August-September (QU). Adults found overwintering on higher and drier ground, in hawthorn hedges, at the edge of woods, and in woods; in leaf litter, under stones, and moss. Predators: toads. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Frequent climber (on plants, bushes, and trees). Collecting techniques. Sweeping or beating the vegetation; light trapping; baiting traps with molasses and fermented sugar. References. Lindroth, 1969a (natural history); Erwin, 1981 (natural history).

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Lebia (Lebia) tuckeri (Casey, 1920) Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Lebia (Lebia) viridipennis Dejean, 1826 Ecology. Lowlands and mountains. Meadows, vacant lots, cultivated fields (e.g., cotton), river banks, lake shores, swampy areas, light or open forests. Open or slightly shaded ground; moderately moist soil. Mostly diurnal; active in the sunshine on flowering plants (e.g., Solidago), bushes, and trees; sheltering on cloudy or cool days in leaf litter and under plant rosettes (e.g., Verbascum). Gregarious (in winter). Biology. Seasonality: January-December. Tenerals: mid-July (WV), September (MD). Adults found overwintering in vacant lots and woods; under dead leaves, logs, and plant rosettes (Verbascum). Dispersal power. Macropterous. Frequent flier (in the sunshine; to artificial lights at night). Moderate runner. Frequent climber (on flowering plants, bushes, trees). Collecting techniques. Sweeping or beating the vegetation; light trapping; using tanglefoot screens. References. Blatchley, 1910 (biology); Leng, 1915 (dispersal power); Lovell, 1915 (ecology); Hemenway and Whitcomb, 1967 (ecology, biology); Kirk, 1969 and 1975c (dispersal power); Erwin, 1981 (natural history). Lebia (Lebia) viridis Say, 1823 Ecology. Lowlands, mountains, subalpine and alpine zones. Grasslands, meadows, abandoned fields (e.g., corn, alfalfa, clover, vetch, soybean, hay, fescue, lespedeza), lawns, gardens, hedges, fencerows, roadsides, sand pits, gravel pits, orchards (e.g., plum, peach), forests (deciduous, mixed or coniferous), their edges and clearings, marshes, swamps, borders of rivers, ponds, pools, and irrigation canals. Open or slightly shaded ground; dry to wet soil. Mostly diurnal; active in the sunshine on the flowers of plants (e.g., Ambrosia, Asclepias, Daucus, Eupatorium, Fragaria, Oenothera, Salvia, Steironema, Taraxacum, Thelesperma), shrubs (e.g., Amelanchier, Ceanothus, Cornus, Crataegus, Parthenocissus, Rhus, Rubus, Salix, Spiraea, Viburnum, Vitis), and trees (e.g., Acer, Betula, Fraxinus, Juglans, Pinus, Populus, Prunus, Quercus, Ulmus), or, on plant foliage (e.g., Andropogon, Carex, Juncus, ferns, grass). Preferably on Solidago, Salix, Cornus, Vitis, and Parthenocissus. Sheltering on cloudy or cool days under dead leaves, stones, other debris, in the soil at the base of plants (Solidago), and in clumps of broomsedges. Associated with chrysomelids (Altica ambiens LeConte, A. chalybaea Illiger, A. foliacea LeConte, A. subplicata LeConte, A. ulmi Woods), which constitutes a case of Batesian mimicry. Semi-gregarious. Biology. Seasonality: JanuaryDecember. Tenerals: July-September. Adults found overwintering on higher and drier ground, in pastures, on hills and hillocks, in hawthorn hedges, at the edge of woods, in woods, and in swamps; in leaf litter, clumps of moss, grass and broomsedges, under stones, and in wood debris. Adult food: chrysomelid eggs, larvae, and pupae (Altica chalybaea, A. foliacea, A. ulmi, Leptinotarsa decemlineata (Say)) (in the field); lepidopterous eggs (in captivity). The larva is ectoparasitic on chrysomelid pupae (Altica, Leptinotarsa). Defensemechanism: when alarmed, the adult may emit a Brachinus-like smoke from the pygidial

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glands; green color of adult body matching the surrrounding vegetation. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; into rotary traps). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (on flowering plants, bushes, trees). Collecting techniques. Sweeping and beating the vegetation; pitfall trapping; light trapping; using rotary traps; using malaise traps; using tanglefoot screens; pyrethrum spraying; hanging yellow sticky boards in vineyards. References. Smith, 1882 (ecology); Blatchley, 1910 (ecology); Lovell, 1915 (ecology); Isely, 1920 (biology); Hemenway and Whitcomb, 1967 (ecology, biology); Madge, 1967 (biology); Kirk, 1969 (natural history), 1970 (ecology), and 1975c (ecology, dispersal power); Lindroth, 1969a (natural history) and 1971b (ecology, biology); Erwin, 1981 (natural history); Capogreco, 1989 (biology); Will et al. 1995 (dispersal power). Lebia (Lebia) vittata (Fabricius, 1777) Ecology. Lowlands and mountains. Borders of standing (e.g., pools and ponds in sand pits) and running waters; moist situations in abandoned fields, grasslands, cultivated fields (e.g., corn), meadows, gardens, sand pits, and gravel pits. Open ground; moist, often sandy soil. Diurnal; active in the sunshine on the foliage of bushes (e.g., Alnus, Sabal, Salix) and on plant flowers (e.g., Amaranthus, Solidago, Spiraea). Preferably on Solidago and Salix. Also found in clumps of broomsedges. Associated with chrysomelids (Disonycha alternata (Illiger), D. procera Casey). Gregarious. Biology. Seasonality: January-December. Adults found overwintering in pastures and near bodies of water; in clumps of broomsedges, and under pieces of wood. Adult food, in the field: chrysomelid larvae (Disonycha alternata). The larva is an ectoparasite of chrysomelid pupae (Dysonycha alternata). Predators: toads. Defense-mechanism: when alarmed, the adult escapes by flying. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Frequent climber (on bushes, shrubs and flowers). Collecting techniques. Sweeping or beating the vegetation; light trapping. References. Lovell, 1915 (ecology; as furcata LeConte); Balsbaugh, 1967 (ecology, biology); Kirk, 1969 (biology, dispersal power) and 1975c (ecology, dispersal power); Lindroth, 1969a (natural history) and 1971b (ecology, biology); Bell, 1971 (ecology, biology). Leistus (Leistus) ferrugineus (Linné, 1758) Ecology. Lowlands. A mixed forest (Abies, Alnus, Prunus), with moist gravelly soil. Nocturnal; sheltering during the day under stones, in leaf litter and in humus. Grassy areas, hedges, and open woodlands (Europe). Biology. Seasonality: August-October. Adult food, in the field (Europe): insects. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Pitfall trapping. References. Lindroth, 1945 and 1985 (natural history); Larson, 1978 (ecology). Leistus (Neoleistus) ferruginosus Mannerheim, 1843 Ecology. Lowlands. Deciduous forests: along streams. Half-shaded ground, often under Alnus- and Salix-bushes; moderately moist soil. Nocturnal; sheltering during the

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day in leaf litter, under stones and under loose bark. Biology. Seasonality: May-August. Tenerals: June. Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Fast runner. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Lindroth, 1961 (natural history); Erwin, 1970a (biology); Kavanaugh, 1992 (natural history). Leistus (Neoleistus) longipennis Casey, 1920 Ecology. Lowlands. Coniferous forests (Sequoia). Biology. Seasonality: June-July. Tenerals: June-July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Pitfall trapping. Leistus (Neoleistus) madmeridianus Erwin, 1970 Ecology. Coniferous forests (Sequoia). Biology. Seasonality: June-July. Tenerals: June-July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Reference. Erwin, 1970a (biology). Leptotrachelus depressus Blatchley, 1923 Ecology. Lowlands. An flooded lake shore, between the leaves and stems of sawgrass (Cladium) growing in the open water. Biology. Seasonality: March. Dispersal power. Unknown. Leptotrachelus dorsalis (Fabricius, 1801) Ecology. Lowlands and mountains. Mostly borders of eutrophic marshes (e.g., Typha), pools, ponds, lakes, and slow rivers; swamps. Also open wet forests, roadside ditches, wet meadows, and cultivated fields (e.g., sugarcane, wheat). Open ground; soil inundated, covered with dense tall vegetation (mostly Typha; also Polygonum, Saccharum, grass, herbs), or, when not inundated, soft, wet, often muddy or clayish, rich in organic debris, covered with similar vegetation. Nocturnal and diurnal; sheltering during the day between the layers of Typha-stems or active on their leaves. Gregarious (in winter). Biology. Seasonality: January-December. Adults found overwintering in swamps and woods; mostly between the layers or sheaths of cattail (Typha) (once found by the hundreds), in leaf litter, and under pieces of wood. Adult food, in the field: lepidopterous eggs and hymenopterous larvae. Larval food, in the field: hymenopterous larvae. Defense-mechanism: when alarmed, the adult emits a strong smell from the pygidial glands. Dispersal power. Macropterous. Frequent flier (to artificial lights at light). Frequent climber (on plants). Collecting techniques. Sweeping or by treading the Typha-vegetation into the the water; sweeping or beating the vegetation; light trapping. References. Webster, 1894 (biology); Blatchley, 1910 (ecology); Grossbeck, 1913 (biology); Leng, 1915 (ecology, dispersal power); Kirk, 1969 (biology, dispersal power), 1970, and 1975c (dispersal power); Lindroth, 1969a (natural history); Will et al. 1995 (dispersal power).

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Leptotrachelus pallidulus Motschulsky, 1864 Ecology. Lowlands. Biology and Dispersal power. Unknown. Reference. Bousquet, 1997a (taxonomy). Licinus silphoides (Rossi, 1790) Ecology. Dry, sandy hills and river banks (Europe). Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Reference. Ball and Bousquet, 2001 (doubtfully established). Lophoglossus gravis LeConte, 1873 Ecology. Lowlands. Cypress-swamps and flood-plain forests. Shaded ground; wet, clayish or muddy soil covered with thick leaf litter. Nocturnal; sheltering during the day under logs and pieces of wood. Biology. Seasonality: March-May, July, October-November. Adults found overwintering in the upper banks of swamps and in forests; under the loose bark of fallen trees and in rotten standing tree-stumps. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Occasional climber (on fallen trees and tree-stumps). Collecting techniques. Pitfall trapping; turning logs. Lophoglossus haldemanni (LeConte, 1848) Ecology. Lowlands. Cypress-swamps. Shaded ground, wet clayish or muddy soil. Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: April-September. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Blatchley, 1910 (ecology); Leng, 1915 (ecology). Lophoglossus scrutator (LeConte, 1848) Ecology. Lowlands. Margins of eutrophic marshes, pools, and ponds; swamps and floodplain forests. Close to water. Open ground; soft, very wet, clayish or muddy soil, rich in organic debris, covered with dense vegetation (e.g., Typha, Salix, Naumbergia, Solanum, grass, Carex, Juncus). Nocturnal; sheltering during the day under large stones, pieces of wood, logs, fallen trees, heaps of grass, or in leaf litter and moss. Biology. Seasonality: March-December. Gravids: May-June and mid-September (QU). Tenerals: May (rarely), August, November. Adults found overwintering on higher and drier ground, in the upper banks of swamps and marshes, and on hills; under stones, in moss, and in the soil (as deep as 37 cm). Adult food, in captivity: scarabaeid larvae, mealworms, meat, and apple. Larval food, in captivity: mealworms. Parasites: ascid mites. Phoretic mites found on adult body. Dispersal power. Macropterous, probably capably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning large stones. References. Blatchley, 1910 (ecology); Lindroth, 1966 (ecology, dispersal power); Will et al. 1995 (dispersal power); Bousquet, 1999 (biology).

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Lophoglossus substrenuus (Csiki, 1930) Ecology. Lowlands. Swamps, flood-plain forests, wet forests, and pond margins in low forests. Shaded ground; wet, clayish or muddy soil, rich in organic debris. Nocturnal; sheltering during the day under stones and logs. Gregarious (in winter). Biology. Seasonality: March-July, September. Tenerals: July (PA). Adults found overwintering in the upper zone of flood-plain forests; in standing, rotten tree-stumps and under dead leaves. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones or logs. Reference. Löding, 1945 (ecology; as strenuus (LeConte)). Lophoglossus tartaricus (Say, 1823) Ecology. Lowlands. Sandy localities near water. A wet forest. Margins of a large manmade canal. Shaded ground; wet, sandy soil covered with thick leaf litter. Nocturnal; sheltering during the day under logs and pieces of wood. Biology. Seasonality: AprilDecember. Adults found overwintering (IN). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning logs; light trapping. References. Leng, 1915 (ecology); Löding, 1945 (ecology). Lophoglossus vernix Casey, 1913 Ecology. Lowlands. Biology. Seasonality: July-August, October. Dispersal power. Macropterous, probably capable of flight. Moderate runner (inferred from the shape of legs). Loricera decempunctata Eschscholtz, 1833 Ecology. Lowlands, mountains, subalpine and alpine zones. Borders of ponds, lakes, marshes, slow rivers and brooks, and seepages; cultivated fields and gardens. Close to water or at some distance from it. Open or shaded ground; moist, often muddy soil, rich in organic debris, covered with some vegetation (e.g., grass, moss). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under pieces of wood, in leaf litter, and in the soil. Biology. Seasonality: February, May, July-August. Tenerals: August. Adults found overwintering in moss. Dispersal power. Macropterous. Occasional flier (at midday). Fast runner. Collecting techniques. Pitfall trapping; lifting pieces of wood; raking the leaf litter. References. Lindroth, 1961 (natural history); Kavanaugh, 1992 (natural history). Loricera foveata LeConte, 1851 Ecology. Unknown. Biology. Seasonality: January, May-June. Dispersal power. Macropterous, probably capable of flight.

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Loricera pilicornis congesta Mannerheim, 1853 Ecology. Coastal lowlands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loricera pilicornis pilicornis (Fabricius, 1775) Ecology. Lowlands and mountains. Borders of eutrophic marshes, ponds, lakes, pools (often polluted by refuse), slow rivers, brooks, and irrigation ditches; fens, swamps, low forests, flood-plain forests, mud flats, wet meadows and pastures, cultivated fields (e.g., barley, dry bean). Ground shaded by bushes and tall plants; soft, wet, often muddy soil, rich in organic debris, covered with sparse vegetation. Diurnal and nocturnal; active in the sunshine or sheltering during the day at the base of plants, under plant mats, heaps of grass, dead leaves, stones, pieces of wood, and in moss. Gregarious (in winter). Biology. Seasonality: January-December. Copulating pairs: May and July (QU). Tenerals: mid-June to September; mostly from mid-July on. Adults found overwintering on higher and drier ground, in the upper banks of marshes and swamps, in woods, at the edge of woods, in vacant lots, and on hillocks; under dead leaves, stones, fallen trees, fallen pieces of bark, under the loose bark of fallen trees, in rotten standing treestumps, and in the soil (as deep as 20 cm). Adult food: springtails (mostly), insect larvae, and mites (in the field, Europe); mealworms and ham (in captivity). Adults use the strong long setae on the basal segments of their antennae as an enclosing trap to capture springtails. Larval food: springtails (in the field); mealworms (in captivity). Larvae use the apical part of their galea as a sticky trap to capture springtails. Predators: frogs and toads. Phoretic mites found on adult body. Defense-mechanism: when alarmed, the adult may slowly emit a discharge from the pygidial glands (at a range less than its body length). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Frequent in lake- and seashore drift material, which indicates flight ability. Fast runner. Occasional climber (on plants and trees). Moderately favored by human activities. Collecting techniques. Pitfall trapping; light trapping; turning drift material; sweeping plants. References. Lindroth, 1945, 1955a, 1961, and 1963a (natural history); Greenslade, 1965 (ecology, biology); Sharova, 1981 (ecology, biology); Bauer, 1982 (biology); Lindroth, 1985 (natural history); Loreau, 1985 (biology); Hintzpeter and Bauer, 1986 (biology); Levesque and Levesque, 1987 (biology); Bauer and Kredler, 1988 (biology); Koch, 1989 (ecology); Luff, 1993 (biology); Will et al. 1995 (dispersal power). Loxandrus accelerans Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus aduncus Allen, 1972 Ecology. Lowlands. Biology. Seasonality: January-February, May-June. Dispersal power. Macropterous, probably capable of flight.

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Loxandrus agilis (Dejean, 1828) Ecology. Lowlands. Pond margins. Nocturnal; sheltering during the day under pieces of wood. Biology. Seasonality: February-April, June, August, December. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; baiting traps with syrup. References. Blatchley, 1910 (ecology); Leng, 1915 (ecology). Loxandrus algidus Allen, 1972 Ecology. Lowlands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Loxandrus brevicollis (LeConte, 1848) Ecology. Border of streams and other water bodies. Flood-plain forests. Moist soil covered with dead leaves. Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: February-August. Defense-mechanism: when alarmed, the adult escapes by running. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Leng, 1915 (dispersal power); Allen, 1972 (natural history); Will and Liebherr, 1997 (ecology). Loxandrus breviusculus Casey, 1924. Ecology. Lowlands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus calathinus LeConte, 1878 Ecology. Lowlands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus celeris (Dejean, 1828) Ecology. Lowlands. Mexico: borders of marshes (Typha, Scirpus). Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: January-December. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Leng, 1915 (ecology); Allen, 1972 (natural history). Loxandrus cervicalis Casey, 1918 Ecology. Lowlands. Pond margins and lake shores. Nocturnal; sheltering during the day. Biology. Seasonality: January, March, August, October. Dispersal power. Macropterous, probably capable of flight. Moderate runner.

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Loxandrus cincinnati Casey, 1924 Ecology. Lowlands. Biology. Seasonality: March-June, November. Dispersal power. Macropterous, probably capable of flight. Loxandrus circulus Allen, 1972 Ecology. Lowlands. Pond margins. Nocturnal; sheltering during the day in leaf litter and in mud. Biology. Seasonality: April, July-August. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter; pouring water over the ground; washing the leaf litter in water. Reference. Erwin, 1981 (natural history). Loxandrus collucens Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus comptus Casey, 1918 Ecology. Lowlands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus concinnus Casey, 1918 Ecology. Lowlands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus contumax Casey, 1918 Ecology. Lowlands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus crenatus LeConte, 1852 Ecology. Lowlands. Biology. Seasonality: February-June, August-September, November. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Collecting technique. Sifting the leaf litter. Reference. Leng, 1915 (ecology). Loxandrus crenulatus Chaudoir, 1868 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight.

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Loxandrus cursitans Casey, 1918 Ecology. Lowlands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus duryi Wright, 1939 Ecology. Lowlands. Biology. Seasonality: January-March, May-August. Dispersal power. Macropterous, probably capable of flight. Loxandrus erraticus (Dejean, 1828) Ecology. Lowlands. Margins of streams and small pools. Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: April-June, December. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. Reference. Allen, 1972 (ecology, biology). Loxandrus extendus Allen, 1972 Ecology. Lowlands. Biology. Seasonality: January, May. Dispersal power. Macropterous, probably capable of flight. Loxandrus floridanus LeConte, 1878 Ecology. Lowlands. Biology. Seasonality: February-July, October-December. Dispersal power. Macropterous, probably capable of flight. Loxandrus fulgens Casey, 1918 Ecology. Lowlands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus gibbus Allen, 1972 Ecology. Lowlands. Biology. Seasonality: February-March, May, July-September. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Loxandrus icarus Will and Liebherr, 1997 Ecology. Lowlands. Along a brook in a deciduous flood-plain forest. Wet sandy soil covered with dead leaves. Nocturnal; active during the night among dead leaves. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Reference. Will and Liebherr, 1997 (taxonomy, ecology, dispersal power).

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Loxandrus inferus Allen, 1972, see Loxandrus velocipes Casey, 1918. Reference. Will and Liebherr, 1997 (taxonomy). Loxandrus infimus Bates, 1882 Ecology. Lowlands. Pond and marsh edges. Nocturnal; sheltering during the day in mud cracks. Biology. Seasonality: June. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; pouring water over the ground; treading the soil with the feet. Reference. Allen, 1972 (biology, dispersal power). Loxandrus lateralis Casey, 1918 Ecology. Lowlands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Loxandrus limatus Casey, 1918 Ecology. Lowlands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Loxandrus lucens Chaudoir, 1868 Ecology. Lowlands. Biology. Seasonality: February-March, May, July-August. Dispersal power. Macropterous, probably capable of flight. Loxandrus ludovicianus Casey, 1918 Ecology. Lowlands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus micans Chaudoir, 1868 Ecology. Lowlands. Macrohabitat unknown. Nocturnal; sheltering during the day in leaf litter and under logs. Biology. Seasonality: March, May, December. Dispersal power. Macropterous, probably capable of flight. Collecting techniques. Pitfall trapping; raking the leaf litter; turning logs Loxandrus minor (Chaudoir, 1843) Ecology. Lowlands. Sandy sites near water. Flood-plain forests. Moist soil covered with dead leaves. Biology. Seasonality: January-February, April, June, December. Adults found overwintering in wood debris. Dispersal power. Macropterous. Occasional in lakeshore

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drift material, which indicates flight ability. Moderate runner. Collecting technique. Raking the leaf litter. References. Blatchley, 1910 (ecology); Lindroth, 1966 (dispersal power); Kirk, 1969 (biology); Will and Liebherr, 1997 (ecology). Loxandrus minutus Allen, 1972 Ecology. Lowlands. Biology. Seasonality: January-June, October-November. Dispersal power. Macropterous, probably capable of flight. Loxandrus mundus Casey, 1918 Ecology. Lowlands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus nitidulus (LeConte, 1848) Ecology. Lowlands. Biology. Seasonality: March-July, October. Dispersal power. Macropterous, probably capable of flight. Loxandrus pactinullus Allen, 1972 Ecology. Damp pastures (Mexico). Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Loxandrus parallelus Casey, 1918 Ecology. Lowlands. Flood-plain forests. Moist soil covered with dead leaves. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: February, April, June-August. Adults found overwintering in wood debris. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Kirk, 1969 (biology); Will and Liebherr, 1997 (ecology). Loxandrus parvicollis Casey, 1918 Ecology. Lowlands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus parvulus Chaudoir, 1868 Ecology. Lowlands. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus piceolus Chaudoir, 1868 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight.

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Loxandrus piciventris (LeConte, 1848) Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus pravitubus Allen, 1972 Ecology. Lowlands. Biology. Seasonality: April-July, October, December. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Loxandrus proximus Chaudoir, 1868 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus rectangulus LeConte, 1878 Ecology. Lowlands. Borders of swamps and marshes. Biology. Seasonality: February, April, September. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Allen, 1972 (ecology, biology). Loxandrus rectus (Say, 1823) Ecology. Lowlands. Biology. Seasonality: January-August, October-December. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Loxandrus reflexus LeConte, 1878 Ecology. Lowlands. Pond margins. Nocturnal; sheltering during the day under logs. Biology. Seasonality: February-March. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning logs. Loxandrus robustus Allen, 1972 Ecology. Lowlands. Flood-plain forests. Shaded ground; moist soil covered with dead leaves. Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: May, July-August. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Allen, 1972 (ecology, biology); Will and Liebherr, 1997 (ecology). Loxandrus rossi Allen, 1972 Ecology. Unknown. Biology. Seasonality: June. Dispersal power. Macropterous, probably capable of flight.

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Loxandrus saccisecundaris Allen, 1972 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus saphyrinus (Chaudoir, 1843) Ecology. Lowlands. Stream margins. Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: January-September, November-December. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. Reference. Allen, 1972 (natural history). Loxandrus sculptilis Bates, 1884 Ecology. Pastures, meadows, and roadsides (Mexico). Biology. Seasonality: June. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Loxandrus spinilunatus Allen, 1972 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus straneoi Will and Liebherr, 1997 Ecology. Lowlands. Flood-plain forests. Moist soil covered with dead leaves. Nocturnal; sheltering during the day under debris. Biology. Seasonality: March. Dispersal power. Wing-dimorphic: macropterous (mostly; probably capable of flight) or brachypterous (incapable of flight). Moderate runner. Collecting technique. Night collecting with a headlamp. Reference. Will and Liebherr, 1997 (taxonomy, ecology, dispersal power). Loxandrus suturalis Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Loxandrus taeniatus LeConte, 1852 Ecology. Lowlands. Flood-plain forests. Moist soil covered with dead leaves. Nocturnal; sheltering during the day under logs. Biology. Seasonality: March-June, August, December. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; turning logs. Reference. Will and Liebherr, 1997 (ecology). Loxandrus uniformis Allen, 1972 Ecology. Lowlands. Biology. Seasonality: March, November. Dispersal power. Macropterous, probably capable of flight.

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Loxandrus unilobus Allen, 1972 Ecology. Lowlands. Biology. Seasonality: June. Dispersal power. Macropterous, probably capable of flight. Loxandrus velocipes Casey, 1918 Ecology. Lowlands. Margins of brooks and ponds; borders of marshes (in low deciduous forests and flood-plain forests). Wet soil covered with dead leaves. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: January, March-July, October. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Lindroth, 1966 (ecology, dispersal power); Allen, 1972 (natural history; as inferus Allen); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power); Will and Liebherr, 1997 (taxonomy). Loxandrus velox (Dejean, 1828) Ecology. Unknown. Biology. Seasonality: February. Adults found overwintering in wood debris. Dispersal power. Macropterous, probably capable of flight. Reference. Kirk, 1969 (biology). Loxandrus vitiosus Allen, 1972 Ecology. Lowlands. River banks. Wet soil. Nocturnal; sheltering during the day under debris. Biology. Seasonality: January, March-July, September, December. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning debris. Loxandrus vulneratus Casey, 1918 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Metacolpodes buchannani (Hope, 1831). Ecology. A forest (BC). Under a pile of dry cow manure (OR). Biology. Seasonality: June. Dispersal power. Macropterous. References. Liebherr, 1998 (taxonomy); Ball and Bousquet, 2001 (taxonomy); Jarrett and Scudder, 2001 (ecology, distribution). Metrius contractus contractus Eschscholtz, 1829 Ecology. Lowlands, mountains, subalpine and alpine zones. Mixed forests; coniferous forests (e.g., Pseudotsuga). Shaded or half-shaded ground; moderately moist, sandy soil. Nocturnal; sheltering during the day in the soil, under stones and fallen trees. Associat-

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ed with the millipede Xystocheir francisca (Chamberlin). Larval habitat: in rotten logs. Biology. Seasonality: March-September. Gravids: mid-March (CA). Tenerals: March and May (CA). Adult and larval food, in the field: millipedes. Adult food, in captivity: meat. Dispersal power. Brachypterous, incapable of flight. Slow runner. Collecting techniques. Pitfall trapping; turning stones and fallen trees. References. MacSwain and Garner, 1956 (biology); Lindroth, 1961 (natural history); Bousquet, 1986a (biology); Weber and Kavanaugh, 1992 (ecology). Metrius contractus planatus Van Dyke, 1925 Ecology. Alpine zone. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Reference. Van Dyke, 1925 (ecology). Metrius contractus sericeus Rivers, 1900 Ecology. Mountains. Slopes. Biology. Seasonality: March-April, July. Dispersal power. Brachypterous, incapable of flight. Metrius explodens Bousquet and Goulet, 1990 Ecology. Mountains. A partially logged coniferous forest (Pinus, Abies). Biology. Seasonality: July. Predators: spiders. Defense-mechanism: when alarmed, the adult emits a repugnant audible spray from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Micranillodes depressus Jeannel, 1963 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Micratopus aenescens (LeConte, 1848) Ecology. Lowlands and mountains. Margins of lakes and swamps; ravines and sinkholes. Soil covered with thick leaf litter. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in leaf litter and under old logs. Biology. Seasonality: January, March-July, September, December. Dispersal power. Macropterous. Occasional flier (during the day; to artificial lights at night). Collecting techniques. Raking and sifting the leaf litter; turning old logs. Reference. Barr, 1971b (natural history). Micrixys distincta (Haldeman, 1852) Ecology. Lowlands and mountains. Grasslands (mostly) and open forests. Open ground; dry soil. Nocturnal; sheltering during the day under stones and at the base of plants. Biology. Seasonality: March-May, July-August, October. Defense-mechanism: body color-pattern offers a striking protective resemblance to Velvet Ants (Hymenoptera: Mutillidae). Dispersal power. Brachypterous, incapable of flight. Moderate runner.

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Collecting techniques. Pitfall trapping; turning stones. Reference. Ball and Bousquet, 2001 (ecology). Microlestes brevilobus brevilobus Lindroth, 1969 Ecology. Lowlands. Biology. Seasonality: February, June. Dispersal power. Macropterous. Occasional in lakeshore drift material, which indicates flight ability. References. Lindroth, 1969a (dispersal power); Will et al. 1995 (dispersal power). Microlestes curtipennis (Casey, 1920) Ecology. Lowlands and uplands. Grasslands. Open ground; dry soil composed of fine sand, covered with sparse vegetation (e.g., grass). Mostly diurnal; active in the bright sunshine; sheltering on cloudy or cool days under stones. Biology. Seasonality: April-June. Dispersal power. Wing-polymorphic: brachypterous (incapable of flight) to macropterous (capable of flight). Moderate runner. Collecting techniques. Hunting with a butterfly net; turning stones. References. Lindroth, 1969a (dispersal power); Bell, 1971 (ecology). Microlestes lindrothi Mateu, 1994 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably, capable of flight. Reference. Mateu, 1994 (taxonomy). Microlestes linearis (LeConte, 1851) Ecology. Lowlands and mountains. Grasslands, relict prairies, meadows, cultivated fields and their borders (e.g., corn, wheat, soybean), abandoned fields, hills, fencerows, forest clearings, and the upper zone of river banks. Open ground; dry soil consisting of fine sand, covered with sparse vegetation (e.g., grass, weeds). Mostly diurnal; active in the bright sunshine; sheltering on cloudy days under stones, dead leaves and plants, and at the base of plants. Biology. Seasonality: January-December. Copulating pairs: May (QU). Tenerals: August (ON). Adults found overwintering on higher and drier ground, in abandoned fields, at the edge of woods, and in woods; under dead leaves, plant rosettes (Verbascum), and fallen trees. Adult food, in captivity: lygaeids, a piece of earthworm, and corn kernels. Dispersal power. Wing-polymorphic: macropterous (mostly) to brachypterous (incapable of flight). Occasional flier (to artificial lights at light). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Strongly favored by human activities. Collecting techniques. Pitfall trapping; hunting with a butterfly net; turning stones; raking the leaf litter. References. Lindroth, 1969a (natural history); Lavigne, 1978 (ecology, biology); Hsin et al. 1979 (biology). Microlestes lucidus lucidus (LeConte, 1851) Ecology and Biology. Unknown. Dispersal power. Submacropterous, probably incapable of flight.

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Microlestes major Lindroth, 1969 Ecology. Unknown. Biology. Seasonality: May. Dispersal power. Macropterous, probably capable of flight. Microlestes nigrinus (Mannerheim, 1843) Ecology. Lowlands and uplands. Mostly cultivated fields (e.g., corn, oat, alfalfa); also river banks, lake shores, and marsh borders. Open ground; dry, sandy-loamy soil. Diurnal; active in the sunshine. Biology. Seasonality: January-December. Adults found overwintering in dry cow dung. Dispersal power. Wing-polymorphic: brachypterous (mostly; incapable of flight) to macropterous (rarely). Occasional flier (to artificial lights at night). Moderate runner. Occasional climber (on plants). Collecting techniques. Pitfall trapping; sweeping plants. References. Lindroth, 1969a (dispersal power); Kirk, 1975c (natural history); Lavigne, 1977 (ecology). Microlestes pusio (LeConte, 1863) Ecology. Lowlands. Cultivated fields (e.g., corn). Edge of a lake (SD). Open ground; sandy-loamy soil. Mostly diurnal; active in the bright sunshine; sheltering on cloudy days at the base of plants and under debris. Biology. Seasonality: May-August, October-December. Adult food, in captivity: lygaeid bugs. Dispersal power. Wing-dimorphic: macropterous (mostly) or brachypterous (incapable of flight). Occasional flier (at dusk; to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Hunting with a butterfly net; pitfall trapping. References. Lindroth, 1969a (dispersal power); Kirk, 1975c (ecology). Mioptachys flavicauda (Say, 1823) Ecology. Lowlands. Forested areas. Nocturnal; sheltering during the day under the loose bark of dead or dying trees (mostly deciduous, e.g., Acer, Ulmus, Populus, Juglans), and of decaying logs; also in wood chips or sawdust. Biology. Seasonality: January-December. Tenerals: October (MI). Adults found overwintering at the edge of woods; under dead leaves. Adult food, in the field: mites and springtails. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Frequent climber (on trees). Collecting techniques. Lifting the loose bark of trees; sifting (e.g., with Berlese funnels) decaying wood chips or sawdust; baiting traps with carrion. References. Ulke, 1903 (ecology); Blatchley, 1910 (ecology); Leng, 1915 (ecology); Lindroth, 1966 (natural history); Kirk, 1975c (ecology); Erwin, 1981 (natural history); Nelson, 1991 (ecology). Miscodera arctica (Paykull, 1798) Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, meadows, sand pits, gravel pits, roadsides, forest edges and clearings, light forests (coniferous, mixed). Ground open or shaded by trees and bushes (e.g., Alnus); rather dry, mostly sandy soil

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(sometimes gravel-mixed) covered with short vegetation (e.g., moss, lichens). Nocturnal; sheltering during the day in burrows dug in the soil, under dead leaves, stones, pieces of wood and cardboard. Associated with byrrhids (Byrrhus, Cytilus). Larval habitat: in the soil and under stones (Europe). Biology. Seasonality: June-September, November. Tenerals: early July to mid-August (NS, QU). Adult food, in the field (Europe): byrrhid larvae. Adult and larval food, in captivity: mealworms. Predators: toads. Defense-mechanism: when alarmed, the adult feigns death. Dispersal power. Macropterous. Flight records (Europe). Occasional in lake- and seashore drift material, which indicates flight ability. Slow runner. Strong burrower. Collecting techniques. Pitfall trapping; raking the leaf litter; baiting bottles with sugar mixture. References. Larsson, 1939 (biology); Lindroth, 1945, 1955a, 1961, and 1963a (natural history); Andersen, 1968 (biology); Luff, 1978 (ecology, biology); Sharova, 1981 (ecology, biology); Lindroth, 1985 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology). Mochtherus tetraspilotus (W.S. MacLeay, 1825) Ecology. Lowlands. Forests. On trunks and branches of standing or fallen trees. Biology. Unknown. Dispersal power. Unknown. Reference. Ball and Bousquet, 2001 (ecology). Morion aridus Allen, 1968 Ecology. Macrohabitat unknown. Found in the stems of saguaro-type cacti. Biology. Seasonality: May, August, November-December. Dispersal power. Unknown. Reference. Ball and Bousquet, 2001 (ecology). Morion monilicornis (Latreille, 1806) Ecology. Lowlands. Forests. Nocturnal; sheltering during the day under the loose bark of fallen trees and logs (mostly; e.g., Pinus), also in rotten logs, under stones, in leaf litter, and in ant nests. Larval habitat: under the loose bark of pine logs. Biology. Seasonality: January-December. Adults found overwintering (SC). Parasites: Laboulbeniales (Fungi). Dispersal power. Wing-condition unknown. Moderate runner. Frequent climber (on fallen trees and logs). Collecting techniques. Lifting the loose bark of trees; using tanglefoot screens; light trapping. References. Schwarz, 1884 (ecology); Leng, 1915 (ecology); Löding, 1945 (ecology); Kirk, 1969 (ecology, biology). Myas coracinus (Say, 1823) Ecology. Lowlands and mountains. Deciduous forests (e.g., Quercus). Shaded ground; loose, moderately moist soil covered with thick leaf litter, logs, pieces of wood, and stones. Nocturnal; sheltering during the day under dead leaves, logs, and stones. Biology. Seasonality: April-October. Tenerals: mid-June (MI) to late July (WI). Predators: grouses. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; baiting traps with molasses; raking the leaf litter. References. Blatchley, 1910 (ecology); Lindroth, 1966 (natural history); Erwin, 1981 (natural history).

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Myas cyanescens Dejean, 1828 Ecology. Lowlands and mountains. Forests (mostly deciduous, e.g., Acer, Quercus; also mixed or coniferous), tree plantations (e.g., Pinus); forest roadsides, clearings, slopes, hillocks, and edges; thickets. Shaded ground; moderately moist soil consisting of humus, gravel or stones, covered with thick leaf litter. Nocturnal; sheltering during the day in leaf litter, under logs, fallen branches and pieces of bark, stones, mushrooms, reindeer moss (Cladonia), and in the soil. Biology. Seasonality: March-October. Copulating pairs: late June (QU). Adults found overwintering on higher and drier ground, in woods, and at the edge of woods; in leaf litter and in the soil. Adult food: Gypsy Moth immature stages and sawfly cocoons (in the field); mealworms (in captivity). Predators: toads and grouses. Defense-mechanism: when alarmed, the adult escapes by burrowing deep into the leaf litter. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter; light trapping. References. Lindroth, 1966 (natural history); Larochelle, 1974a (ecology); Dearborne and Donahue, 1993 (biology); Bousquet, 1999 (biology). Myas lindrothi (Allen, 1980) Ecology. Mountains. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Neaphaenops tellkampfii henroti Jeannel, 1949 Ecology. Caves. Biology. Seasonality: August. Adult food, in the field: small invertebrates. Dispersal power. Brachypterous, incapable of flight. Fast runner. Neaphaenops tellkampfii meridionalis Barr, 1959 Ecology. Caves: on the ground (well above stream and pool banks), walls or ledges. Wet, silty soil. Biology. Seasonality: January, March. Tenerals: early July. Adult food: a small chrysomelid. Dispersal power. Brachypterous, incapable of flight. Fast runner. Good climber. Neaphaenops tellkampfii tellkampfii (Erichson, 1844) Ecology. Caves: mostly along streams; on ledges or sheltering under stones. Dry to wet, silty soil. Biology. Seasonality: January-December. Copulating pairs: December. Tenerals: all months, except May; mostly late spring to early autumn. Adult food, in the field: cavecricket eggs and nymphs (mostly), dipteran larvae, spiders, worms, flies, springtails, and other small arthropods. Adults and larvae are cannibalistic. Parasites: Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult may emit a faint smell from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Fast runner. Good climber. References. Kane, 1975 (ecology, biology); Kane et al. 1975 (biology); Norton et al. 1975 (biology); Turanchik and Kane, 1979 (ecology); Kane and Ryan, 1983 (biology).

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Neaphaenops tellkampfii viator Barr, 1979 Ecology. Caves. Biology. Seasonality: September. Adult food, in the field: small invertebrates. Dispersal power. Brachypterous, incapable of flight. Fast runner. Nebria (Reductonebria) acuta acuta Lindroth, 1961 Ecology. Lowlands, mountains, subalpine and alpine zones. Banks of cold, swift small brooks (originating from snowfields). Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: June-August. Tenerals: late August (AK). Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Fast runner. References. Lindroth, 1961 (natural history); Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) acuta quileute Kavanaugh, 1979 Ecology. Mountains. Banks of rivers and springs. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: May-August. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) acuta sonorae Kavanaugh, 1981 Ecology. Mountains. Stream banks. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: June, August. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1981 and 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Nipponebria) altisierrae Kavanaugh, 1984, see Nipponebria (Vancouveria) altisierrae (Kavanaugh, 1984). Reference. Kavanaugh, 1995 (taxonomy). Nebria (Reductonebria) appalachia Darlington, 1931 Ecology. Mountains. Banks of small forest streams. Wet, gravelly soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: May-September. Tenerals: early July (TN). Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones. References. Barr, 1969 (ecology); Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) arkansana arkansana Casey, 1913 Ecology. Mountains, subalpine and alpine zones. Margins of cool swift streams (e.g., small brooks). Wet, stony-gravelly soil. Nocturnal; sheltering during the day under stones.

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Biology. Seasonality: June-August. Tenerals: late July to early August. (AB, BC). Adult and larval food, in captivity: mealworms. Dispersal power. Wing-polymorphic: brachypterous (incapable of flight) to macropterous. Fast runner. Collecting technique. Turning stones. References. Lindroth, 1961 (natural history); Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) arkansana edwardsi Kavanaugh, 1979 Ecology. Mountains. Margins of cool streams. Wet soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: June-August. Copulating pairs: early August (MT). Dispersal power. Macropterous. Fast runner. Collecting technique. Turning stones. References. Kavanaugh, 1979 and 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) arkansana fragilis Casey, 1924 Ecology. Mountains. Margins of cool streams. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: July-September. Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) arkansana oowah Kavanaugh, 1979 Ecology. Mountains. Margins of cool streams. Wet soil. Biology. Seasonality: August. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1979 and 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Nebria) brevicollis (Fabricius, 1792) Ecology. Lowlands. Deciduous forests (mostly) and parks (Europe). Biology. Seasonality: unknown. Adult food, in the field (Europe): mostly insects. Dispersal power. Macropterous. Flight records (Europe). Fast runner. References. Lindroth, 1945, 1963a, and 1985 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology). Comments. Doubtfully established in North America. Nebria (Catonebria) calva Kavanaugh, 1984 Ecology. Mountains. A coniferous forest (Picea, Abies). Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Nipponebria) campbelli Kavanaugh, 1984, see Nipponebria (Vancouveria) campbelli (Kavanaugh, 1984). Reference. Kavanaugh, 1995 (taxonomy).

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Nebria (Catonebria) carri Kavanaugh, 1979 Ecology. Mountains. Margins of cool springs. Wet soil. Biology. Seasonality: July-August. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) catenata Casey, 1913 Ecology. Mountains. Banks of rivers and springs. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: June-August. Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) charlottae Lindroth, 1961 Ecology. Coastal lowlands. Upper zone of sea beaches. Partially shaded ground; wet, cobbly soil, with freshwater seepages running through the gravel-sand layer underneath the cobbles. Nocturnal; sheltering during the day under cobbles; active at night on logs. Biology. Seasonality: July-August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Climber (on logs). Collecting technique. Turning stones. References. Kavanaugh, 1985 (dispersal power) and 1992 (natural history); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) coloradensis Van Dyke, 1943 Ecology. Mountains. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: June-August. Dispersal power. Brachypterous, incapable of flight. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Boreonebria) crassicornis crassicornis Van Dyke, 1925 Ecology. Mountains. Forests, bog edges, and mountain slopes near snow patches. Moist or wet soil. Biology. Seasonality: July. Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Fast runner. References. Van Dyke, 1925 (ecology); Lindroth, 1961 (natural history); Kavanaugh, 1985 (dispersal power). Nebria (Boreonebria) crassicornis intermedia Van Dyke, 1949 Ecology. Mountains and alpine zone. Dry forests, moist alpine meadows, and edges of snowfields. Nocturnal; sheltering during the day under stones and logs. Biology. Seasonality: June-August. Tenerals: mid-July (MT) to early August (AB). Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Fast runner. Collecting technique. Turning stones and logs. References. Lindroth, 1961 (natural history); Kavanaugh, 1985 (dispersal power).

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Nebria (Reductonebria) danmanni Kavanaugh, 1981 Ecology. Mountains. Macrohabitat unknown. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: July, September. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Kavanaugh, 1981 and 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) darlingtoni Kavanaugh, 1979 Ecology. Mountains. River banks and vicinity of waterfalls. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: July-September. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) desolata Kavanaugh, 1971 Ecology. Mountains. Base of steep sandstone cliffs. Shaded ground; cool, wet soil. Nocturnal; sheltering during the day in debris and under blocks of sandstone. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Kavanaugh, 1971 (ecology, dispersal power) and 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) diversa LeConte, 1863 Ecology. Coastal lowlands. Sea beaches (above the high tidemark). Open ground; wet, sandy, bare soil. Nocturnal; sheltering during the day under driftwood, other debris, and in cliff bank crevices at some distance from water. Semi-gregarious. Larval habitat: under driftwood. Biology. Seasonality: May-August, November. Tenerals: May-June (BC), late July (OR). Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning driftwood. References. Lindroth, 1961 (natural history); Kavanaugh, 1985 (biology) and 1992 (natural history); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) eschscholtzii Ménétriés, 1843 Ecology. Mountains. Stream banks. Wet soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: June-July. Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Fast runner. Collecting technique. Turning stones. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Boreonebria) frigida R.F. Salhberg, 1844 Ecology. Mountains. River banks. Ground shaded by bushes and trees; rather wet soil. Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: June-

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July. Tenerals: early July (AK). Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Fast runner. Collecting technique. Raking the leaf litter. References. Lindroth, 1961 (natural history); Kavanaugh, 1985 (dispersal power). Nebria (Catonebria) gebleri albimontis Kavanaugh, 1984 Ecology. Mountains. Wet soil. Biology. Seasonality: July. Dispersal power. Macropterous. References. Kavanaugh, 1984 and 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) gebleri cascadensis Kavanaugh, 1979 Ecology. Lowlands and mountains. River banks, brook margins, and lake shores. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: June-October. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) gebleri fragariae Kavanaugh, 1979 Ecology. Mountains. Brook margins. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: May. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) gebleri gebleri Dejean, 1831 Ecology. Mountains. Margins of streams. Close to water. Wet, stony, bare soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: May-August. Tenerals: August (AB). Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Fast runner. Collecting technique. Turning stones. References. Lindroth, 1961 (natural history); Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) gebleri rathvoni LeConte, 1853 Ecology. Mountains. River banks. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: May-June, August. Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) gebleri siskiyouensis Kavanaugh, 1979 Ecology. Mountains. River banks and brook margins. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: June-August. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1979 and 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy).

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Nebria (Boreonebria) gouleti Kavanaugh, 1979 Ecology. Mountains. Banks of rivers, brooks, and waterfalls. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: May-September. Dispersal power. Macropterous. Fast runner. Reference. Kavanaugh, 1985 (dispersal power). Nebria (Reductonebria) gregaria Fischer von Waldheim, 1820 Ecology. Lowlands. Banks of rivers and brooks; upper zone of sea beaches. Not restricted to the vicinity of water. Open ground; moderately wet, stony, gravelly or peaty soil covered with some vegetation. Nocturnal; sheltering during the day. Biology. Seasonality: July, October. Tenerals: from late July (AK). Adult and larval food, in captivity: mealworms. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Lindroth, 1961 (natural history); Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Boreonebria) gyllenhali castanipes (Kirby, 1837) Ecology. Lowlands, mountains, subalpine and alpine zones. Banks of cold rivers, brooks, and rivulets; seashore. Sometimes at some distance from water. Ground shaded by bushes (e.g., Alnus) or open (tundra and mountain tops); wet stony, gravelly soil or moderately moist silty soil sometimes mixed with sand or detritus. Nocturnal; sheltering during the day under stones (mostly), logs, dead leaves, and fallen branches. Biology. Seasonality: June-September. Copulating pairs: July (QU). Gravids: late June (QU). Tenerals: early summer (mostly); autumn (NH). Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Fast runner. Collecting technique. Turning stones. References. Lindroth, 1945, 1955a, 1961, 1963a, and 1985 (natural history); Kavanaugh, 1985 (dispersal power). Nebria (Boreonebria) gyllenhali lassenensis Kavanaugh, 1979 Ecology. Mountains. Lake shores and brook margins. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: July-August, October. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1979 and 1985 (dispersal power). Nebria (Boreonebria) gyllenhali lindrothi Kavanaugh, 1979 Ecology. Mountains. River and brook margins; lake shores. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: June-August. Tenerals: July. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Kavanaugh, 1979 and 1985 (dispersal power). Nebria (Reductonebria) haida Kavanaugh, 1984 Ecology. Alpine zone. Summits and ridges (far from snowfields and water). Open ground; moist soil covered with sparse vegetation. Nocturnal; sheltering during the

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day under stones. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones. References. Kavanaugh, 1985 (dispersal power) and 1992 (natural history); Ball and Bousquet, 2001 (taxonomy). Nebria (Boreonebria) hudsonica LeConte, 1863 Ecology. Mountains, subalpine and alpine zones. Banks of rivers and brooks. Open ground; wet, stony, bare soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: June-August. Tenerals: May to early July (BC). Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Fast runner. Collecting technique. Turning stones. References. Lindroth, 1961 (natural history); Kavanaugh, 1985 (dispersal power). Nebria (Catonebria) ingens ingens G.H. Horn, 1870 Ecology. Mountains. Lake shores. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: August. Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) ingens riversi Van Dyke, 1925 Ecology. Mountains. Margins of glacial streams. Wet soil. Nocturnal; sheltering during the day in rocky crevices. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Van Dyke, 1925 (ecology); Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) jeffreyi Kavanaugh, 1984 Ecology. Mountains. Banks of rivers and brooks. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: July. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) kincaidi balli Kavanaugh, 1979 Ecology. Mountains. Lake shores; borders of waterfalls. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: June-September. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) kincaidi kincaidi Schwarz, 1900 Ecology. Lowlands and mountains. Margins of small brooks crossing forests. Shaded ground; wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: May-June,

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August. Tenerals: early August (WA). Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Lindroth, 1961 (natural history); Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) labontei Kavanaugh, 1984 Ecology. Mountains. Macrohabitat unknown. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: July, September. Dispersal power. Macropterous. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Boreonebria) lacustris bellorum Kavanaugh, 1979 Ecology. Mountains. Banks of rivers and brooks. Shaded ground; wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: May-September. Dispersal power. Macropterous. Fast runner. Reference. Kavanaugh, 1985 (dispersal power). Nebria (Boreonebria) lacustris lacustris Casey, 1913 Ecology. Lowlands and mountains. Banks of large rivers and brooks (with water often silty and slower than in the habitat of Nebria pallipes). Shaded or open ground; wet, clayish, sandy, gravelly or stony, bare soil. Nocturnal; sheltering during the day under stones (mostly) and pieces of wood. Semi-gregarious. Biology. April-October. Tenerals: April to late July. Adult food, in the field: dead or dying insects. Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Fast runner. Occasional climber (on plants). Collecting techniques. Turning stones; night collecting with a headlamp. References. Lindroth, 1961 (natural history); Spence, 1979 (biology); Erwin, 1981 (natural history); Kavanaugh, 1985 (dispersal power); Will et al. 1995 (ecology). Nebria (Reductonebria) lituyae Kavanaugh, 1979 Ecology. Alpine zone. Hill summits. Moist soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) louiseae Kavanaugh, 1984 Ecology. Coastal lowlands. Upper zone of sea beaches. Partially shaded ground; wet, cobbly soil, with freshwater seepages running through the gravel-sand layer underneath the cobbles. Nocturnal; sheltering during the day; active at night on logs. Biology. Seasonality: June-August. Copulating pairs: August (BC). Dispersal power. Brachypterous, incapable of flight. Fast runner. Climber (on logs). References. Kavanaugh, 1985 (dispersal power) and 1992 (natural history); Ball and Bousquet, 2001 (taxonomy).

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Nebria (Reductonebria) lyelli Van Dyke, 1925 Ecology. Mountains. Margins of glacier streams. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Van Dyke, 1925 (ecology); Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) mannerheimii Fischer von Waldheim, 1828 Ecology. Lowlands and mountains. Margins of streams. Wet, stony, bare soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: May-September. Tenerals: mid-May (OR) to late June (BC). Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Fast runner. Collecting technique. Turning stones. References. Lindroth, 1961 (natural history); Kavanaugh, 1985 (dispersal power) and 1992 (natural history); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) meanyi giulianii Kavanaugh, 1981 Ecology. Mountains. Brook margins. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: June-July. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1981 and 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) meanyi lamarckensis Kavanaugh, 1979 Ecology. Mountains. Brook margins. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: July-August. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) meanyi meanyi Van Dyke, 1925 Ecology. Lowlands and mountains. Stream margins. Wet soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: June-September. Tenerals: early September (WA). Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Fast runner. Collecting technique. Turning stones. References. Van Dyke, 1925 (ecology); Lindroth, 1961 (natural history); Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) meanyi sylvatica Kavanaugh, 1979 Ecology. Mountains. Vicinity of springs. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: May-August. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy).

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Nebria (Catonebria) metallica Fischer von Waldheim, 1820 Ecology. Lowlands. Stream margins, at some distance from water, on ground shaded by bushes and trees. Open country (Alaska and its islands); not associated with water. Nocturnal; sheltering during the day under stones and dead leaves. Biology. May-August. Tenerals: late August (AK). Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Fast runner. Collecting techniques. Turning stones; raking the leaf litter. References. Lindroth, 1961 (natural history); Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) navajo Kavanaugh, 1979 Ecology. Mountains. Macrohabitat unknown. Wet soil. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. References. Kavanaugh, 1979 and 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Boreonebria) nivalis gaspesiana Kavanaugh, 1979 Ecology. Lowlands, mountains, and alpine zone. Banks of cold rivers and brooks. Close to snowfields. Open ground; wet, stony, bare soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: June-September. Copulating pairs: late July (QU). Tenerals: early July (NF) to mid-August (QU). Dispersal power. Macropterous. Fast runner. Collecting technique. Turning stones. Reference. Kavanaugh, 1985 (dispersal power). Nebria (Boreonebria) nivalis nivalis (Paykull, 1790) Ecology. Coastal tundra. Margins of snowfields; banks of cold rivers and brooks. Open ground; wet, sandy, muddy or gravelly, bare soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: July-August. Tenerals: early August (QU). Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Fast runner. Collecting technique. Turning stones. References. Lindroth, 1945, 1961, 1963a, and 1985 (natural history); Kavanaugh, 1985 (dispersal power). Nebria (Reductonebria) obliqua chuskae Kavanaugh, 1979 Ecology. Mountains. Brook margins. Wet soil. Biology. Seasonality: July-August. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Kavanaugh, 1979 and 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) obliqua obliqua LeConte, 1866 Ecology. Lowlands, mountains, subalpine and alpine zones. Wet stream banks. Open gravelly or stony ground (at higher elevations); shaded muddy ground (at lower elevations). Nocturnal; sheltering during the day. Biology. Seasonality: May-September. Tenerals: early summer, e.g., from late May. Adult and larval food, in captivity: mealworms.

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Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) ovipennis LeConte, 1878 Ecology. Mountains and subalpine zone. Meadows and snowfields. Moist or wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: July-September. Tenerals: early July to late August. Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) pallipes Say, 1823 Ecology. Lowlands and mountains. Mostly banks of clear, swift rivers and brooks; also lake shores and cave stream banks. Ground shaded by trees and bushes; wet, stonygravelly, bare soil. Nocturnal; sheltering during the day under stones (mostly), pieces of wood, and dead leaves. Semi-gregarious. Biology. Seasonality: April-October. Copulating pairs: late August (VT). Gravids: August (QU). Tenerals: May-August; mostly before July. Adult and larval food, in captivity: mealworms. Parasites: Laboulbeniales (Fungi). Dispersal power. Wing-dimorphic: macropterous or brachypterous (incapable of flight). Fast runner. Collecting technique. Turning stones. References. Blatchley, 1910 (ecology); Spence, 1979 (biology); Erwin, 1981 (natural history); Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Boreonebria) paradisi Darlington, 1931 Ecology. Mountains and subalpine zone. Stream margins. Wet soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: July-August. Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones. Reference. Kavanaugh, 1985 (dispersal power). Nebria (Catonebria) piperi Van Dyke, 1925 Ecology. Lowlands, mountains, and subalpine zone. Banks of cold rivers and brooks. Close to water. Wet, stony, bare soil. Nocturnal; sheltering during the day under large stones. Biology. Seasonality: June-July, October. Tenerals: June (AK, BC). Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Fast runner. Collecting technique. Turning stones. References. Van Dyke, 1925 (ecology); Lindroth, 1961 (natural history); Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) piute piute Erwin and Ball, 1972 Ecology. Mountains. A lake shore. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: September. Dispersal power. Brachypterous, incapable of flight. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy).

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Nebria (Catonebria) piute sevieri Kavanaugh, 1984 Ecology. Mountains. Macrohabitat unknown. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: June-August. Dispersal power. Brachypterous, incapable of flight. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) piute utahensis Kavanaugh, 1979 Ecology. Mountains. Macrohabitat unknown. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. References. Kavanaugh, 1979 and 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) purpurata LeConte, 1878 Ecology. Mountains, subalpine and alpine zones. Margins of cold streams. Wet, sandystony soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: JuneAugust. Copulating pairs: mid-June (NM); at night on emergent rocks in a stream. Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Fast runner. Good swimmer. Collecting technique. Turning stones. References. Kavanaugh and Martinko, 1972 (ecology, biology); Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) sahlbergii modoc Kavanaugh, 1979 Ecology. Mountains. Brook margins. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: May-June. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) sahlbergii sahlbergii Fischer von Waldheim, 1828 Ecology. Lowlands, mountains, and alpine zone. Margins of streams (often small); lake shores (at high elevation). Shaded ground; wet, gravelly-stony soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: May-August. Tenerals: May to mid-July. Adult and larval food, in captivity: mealworms. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Fast runner. Collecting technique. Turning stones. References. Kavanaugh, 1985 (dispersal power) and 1992 (natural history); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) sahlbergii triad Kavanaugh, 1979 Ecology. Mountains. Stream margins. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: June-August. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy).

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Nebria (Catonebria) schwarzi beverlianna Kavanaugh, 1979 Ecology. Mountains. A river bank. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: July-August. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) schwarzi schwarzi Van Dyke, 1925 Ecology. Mountains. Stream margins. Wet bare soil composed of coarse sand. Nocturnal; sheltering during the day under stones. Biology. Seasonality: June-October. Tenerals: mid-July to mid-August (AB, BC). Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Fast runner. Collecting technique. Turning stones. References. Van Dyke, 1925 (ecology); Lindroth, 1961 (natural history); Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) sierrablancae Kavanaugh, 1984 Ecology. Mountains. Macrohabitat unknown. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) spatulata sierrae Kavanaugh, 1979 Ecology. Mountains. Brook margins and lake shores. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: July-September. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Kavanaugh, 1979 and 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) spatulata spatulata Van Dyke, 1925 Ecology. Mountains and alpine zone. Brook margins and lake shores. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: July-September. Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) steensensis Kavanaugh, 1984 Ecology. Mountains. Biology. Seasonality: July-August. Dispersal power. Brachypterous, incapable of flight. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) suturalis LeConte, 1850 Ecology. Lowlands, mountains, subalpine and alpine zones. CO and northern QU: brook margins. NH: below the tree line, along brook margins; above the tree line, along seep-

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ages and trickles running on mountains slopes. VT: in summer, on mountain slopes, in deep crevices; in spring and on cold rainy summer days, on the slopes above the crevices. Open ground; wet or moist soil. Nocturnal; sheltering during the day under stones. Larval habitat: in crevices. Biology. Seasonality: June-August. Tenerals: late July (QU), early August (VT). Adult food, in the field: beetles. Adult and larval food, in captivity: mealworms. Dispersal power. Wing-dimorphic: macropterous or brachypterous (incapable of flight). Fast runner. Collecting technique. Turning stones. References. Lindroth, 1961 (natural history); Bell, 1978 (ecology); Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) trifaria pasquineli Kavanaugh, 1984 Ecology. Mountains. Biology. Seasonality: May-August. Dispersal power. Brachypterous, incapable of flight. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) trifaria trifaria LeConte, 1878 Ecology. Mountains. Snowfields and vicinity. Wet soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: July-August. Adult food, in the field: scarabaeids. Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Boreonebria) turmaduodecima Kavanaugh, 1981 Ecology. Mountains, alpine zone. Lake shores and snowfield edges. Wet soil. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Kavanaugh, 1981 and 1985 (dispersal power). Nebria (Catonebria) vandykei vandykei Bänninger, 1928 Ecology. Mountains. Stream margins. Wet soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: July. Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Catonebria) vandykei wyeast Kavanaugh, 1979 Ecology. Mountains. River and brook margins. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: June-August. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy).

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Nebria (Nipponebria) virescens G.H. Horn, 1870, see Nipponebria (Vancouveria) virescens (G.H. Horn, 1870). Reference. Kavanaugh, 1995 (taxonomy). Nebria (Reductonebria) wallowae Kavanaugh, 1984 Ecology. Mountains. Lake shores and stream margins. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: July, September. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) zioni oasis Kavanaugh, 1979 Ecology. Mountains. Brook margins. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nebria (Reductonebria) zioni zioni Van Dyke, 1943 Ecology. Mountains. A canyon (UT). Biology. Seasonality: May. Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. References. Kavanaugh, 1985 (dispersal power); Ball and Bousquet, 2001 (taxonomy). Nelsonites jonesi Valentine, 1952 Ecology. Caves: vicinity of pools and drips. Standing during the day on moist walls, flowstone, and stalagmites. Biology. Seasonality: January, August-November. Adult food, in the field: worms and dipterous larvae. Dispersal power. Brachypterous, incapable of flight. Fast runner. Frequent climber (on cave walls). Nelsonites walteri Valentine, 1952 Ecology. Caves. Sheltering during the day under stones. Biology. Seasonality: January, June, September-October. Tenerals: June, October. Adult food, in the field: small invertebrates. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones. Nemotarsus elegans LeConte, 1853 Ecology. Lowlands. Light or open forests, orchards (plum, peach), and cultivated fields (e.g., cotton). Slightly shaded ground; rather dry soil. Nocturnal; sheltering during the day mostly under the loose bark of trees (e.g., Quercus), also on plants and in moss. Biology. Seasonality: January-September, November-December. Adults found overwintering under the loose bark of standing trees. Dispersal power. Macropterous. Oc-

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casional flier (to artificial lights at night). Moderate runner. Frequent climber (on trees and plants). Collecting technique. Beating or sweeping the vegetation. References. Leng, 1915 (biology); Löding, 1945 (ecology); Kirk, 1969 (ecology). Nipponebria (Vancouveria) altisierrae (Kavanaugh, 1984) Ecology. Mountains. Macrohabitat unknown. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: July-August, November. Dispersal power. Macropterous. Fast runner. References. Kavanaugh, 1985 (dispersal power) and 1995 (taxonomy). Nipponebria (Vancouveria) campbelli (Kavanaugh, 1984) Ecology. Mountains. Biology. Seasonality: July-August. Dispersal power. Macropterous. References. Kavanaugh, 1985 (dispersal power) and 1995 (taxonomy). Nipponebria (Vancouveria) virescens (G.H. Horn, 1870) Ecology. Lowlands. A meadow, on slightly shaded ground with rather dry soil. Biology. Seasonality: April-October. Adult and larval food, in captivity: mealworms. Predators: spiders (Black Widow). Dispersal power. Macropterous. References. Lindroth, 1961 (natural history); Kavanaugh, 1985 (dispersal power) and 1995 (taxonomy). Nomius pygmaeus (Dejean, 1831) Ecology. Lowlands and mountains. Mostly coniferous forests (e.g., Pseudotsuga); human habitations. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under the loose bark of fallen trees, logs, stones, and in leaf litter. Biology. Seasonality: February-November. Tenerals: October (NC). Adults found overwintering in wood debris. Predators: spiders (frequently). Defense-mechanism: when alarmed, the adult emits a strong fetid smell, reminiscent of some old cheese or dead mouse, which clings deeply to the fingers when the beetle is seized. The smell from one live individual may render a house inhabitable for two weeks. Economic importance: sometimes a pest of stored products. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; to the smoke of forest fires). Occasional in drift material, which indicates flight ability. Moderate runner. Long-distance disperser. Strongly favored by human activities. Collecting techniques. Lifting loose bark; light trapping. References. Schwarz, 1889 (biology); Barrows, 1897 (natural history); Holland, 1907 (biology); Blatchley, 1910 (ecology, biology); Hatch, 1931 (biology); Spencer, 1942 and 1953 (natural history); Lindroth, 1961 (natural history). Notiobia (Anisotarsus) brevicollis (Chaudoir, 1837) Ecology. Mountains. Grasslands. Open forests (e.g., Pinus, Quercus): often along trails. Open or shaded ground; dry soil, with bare spots surrounded by sparse vegetation. Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: January-

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November. Copulating pairs: September (TX). Adult food, in the field: scarabaeids. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter. Reference. Noonan, 1973 (natural history). Notiobia (Anisotarsus) cephalus (Casey, 1914) Ecology. Lowlands. Forests (e.g., Quercus). Nocturnal; sheltering under the bark of a tree and in a rotting tree-stump (FL). Biology. March-June, August-September. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Reference. Shpeley, 2001 (taxonomy). Notiobia (Anisotarsus) maculicornis (Chaudoir, 1843) Ecology. Lowlands. In a thicket. On docks. Among dead leaves. Biology. April-May, August-October. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Notiobia (Anisotarsus) mexicana (Dejean, 1829) Ecology. Mountains. Grasslands; open forests (e.g., Pinus, Quercus). Open or shaded ground. Nocturnal; sheltering during the day under stones. Biology. Seasonality: January-December. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Turning stones. Reference. Noonan, 1973 (natural history). Notiobia (Anisotarsus) nitidipennis (LeConte, 1848). Ecology. Lowlands and mountains. Light or open deciduous forests (e.g., Acer, Quercus, Fagus), their roadsides, clearings, and edges. Grassy meadows (FL). Shaded ground; moderately moist, sandy, gravelly or stony soil covered with dead leaves, moss carpets or grass. Nocturnal; sheltering during the day under stones (mostly) and dead leaves. Biology. Seasonality: January-December. Tenerals: mid-June (rarely), August to early November. Adults found overwintering in woods and at the edge of woods; under stones and in leaf litter. Adult and larval food, in captivity: mealworms. Predators: toads. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; to artifical lights in captivity). Frequent in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Turning stones; pitfall trapping; light trapping; turning drift material. References. Blatchley, 1910 (ecology); Leng,1915 (ecology, dispersal power); Lindroth, 1968 (natural history); Noonan, 1973 (natural history); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power); Shpeley, 2001 (taxonomy). Notiobia (Anisotarsus) purpurascens (Bates, 1882) Ecology. Lowlands and mountains. Vicinity of two ponds (CA): on moist bare clay soil covered with scattered trees (e.g., Salix); in a clump of Populus-trees and Typha.

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Biology. Seasonality: January-December. Dispersal power. Macropterous Occasional flier (to artificial lights at night). Moderate runner. Reference. Noonan, 1973 (natural history). Notiobia (Anisotarsus) sayi (Blatchley, 1910) (Anisotarsus piceus (LeConte, 1848)) Ecology. Lowlands. Sand pits, sandy blowouts, gravel pits, vacant lots, cultivated fields, and rubbish dumps. Open ground; soft, dry soil consisting of fine sand, covered with sparse vegetation. Nocturnal; sheltering during the day under pieces of wood and other debris. Biology. Seasonality: January-October. Tenerals: early June (OH), mid-July (MI). Predators: toads, woodpeckers (flickers), and flycatchers. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Strongly favored by human activities. Collecting techniques. Turning pieces of wood; pitfall trapping; light trapping. References. Lindroth, 1968 (natural history; as piceus (LeConte)); Kirk, 1969 (dispersal power; as piceus (LeConte)); Noonan, 1973 (natural history; as piceus (LeConte)); Kirk, 1975c (dispersal power; as piceus (LeConte)). Notiobia (Anisotarsus) terminata (Say, 1823) Ecology. Vacant lots, meadows, cultivated fields (e.g., corn, tobacco, wheat), golf courses, sand pits, gravel pits, orchards (e.g., peach, plum), and forest clearings. Open ground; dry soil consisting of fine sand, covered with sparse vegetation. Nocturnal; sheltering during the day under dead leaves and plants (e.g., corn, hay, weeds), in clumps of broomsedges, under pieces of wood, stones, pieces of fallen bark or cardboard, and under fallen trees. Semi-gregarious. Biology. Seasonality: February-December. Copulating pairs: August (QU). Gravids: late September (QU). Tenerals: February (AL), June-August, October. Adults found overwintering in vacant lots, meadows, and at the edge of woods; under logs, dead leaves, stones, and in the soil. Adult food: pollen, seeds, and the tissue of grass (e.g., Sorghum) and weeds (e.g., Erechtites) (in the field); grass seeds, beetles (legs removed), curculionid larvae, lepidopterous caterpillars and pupae, and mealworms (in captivity). Larval food, in captivity: mealworms. Economic importance: sometimes causing considerable damage to milo seedlings. Predators: toads (frequently), frogs, and crows. Parasites: mites and tapeworms. Defense-mechanism: when alarmed, the adult may emit a Brachinus-like smoke from the pygidial glands. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Frequent climber (on plants and trees). Strongly favored by human activities. Collecting techniques. Raking dead leaves; pitfall trapping; light trapping; sweeping or beating the vegetation. References. Blatchley, 1910 (ecology, biology); Lindroth, 1968 (natural history); Kirk, 1969 (ecology, dispersal power), 1970 (dispersal power), and 1975c (ecology, dispersal power); Noonan, 1973 (natural history); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power).

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Notiophilus aeneus (Herbst, 1806) Ecology. Lowlands and mountains. Deciduous forests (mostly; e.g., Acer, Quercus, Ostrya), mixed forests, and coniferous forests (e.g., Pinus); forest clearings and roadsides. Shaded ground; moderately moist soil covered with thick leaf litter. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days among dead leaves (mostly) and in moss. Semi-gregarious. Biology. Seasonality: January-December. Tenerals: JulySeptember. Adults found overwintering in woods and at the edge of woods; in leaf litter (mostly), in the soil, and under the loose bark of fallen trees. Adult food: lepidopterous caterpillars (in the field); meat (in captivity). Predators: toads and thrushes. Defense-mechanism: when alarmed, the adult escapes by running. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight activity. Fast runner. Frequent climber (on trees). Collecting techniques. Raking the leaf litter; pitfall trapping; beating trees. References. Lindroth, 1961 (natural history); Larochelle, 1974a (ecology); Erwin, 1981 (natural history); Reeves et al. 1983 (biology); Levesque and Levesque, 1986 (biology). Notiophilus aquaticus (Linné, 1758) Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, fields, meadows, vacant lots, cultivated fields (e.g., corn, barley), roadsides, sand pits, gravel pits, and forest edges. Open ground; well-drained, moderately dry, gravelly or sandy soil covered with sparse (mostly) or moderate vegetation. Mostly diurnal; active in the sunshine (e.g., among grass tufts); sheltering on cloudy or cool days under stones (mostly), pieces of wood and cardboard, or in the soil at the base of plants. Gregarious. Larval habitat: in the soil. Biology. Seasonality: January-December. Tenerals: late June (QU) to late August (NS, QU). Adults found overwintering on higher and drier ground, in fields and at the edge of woods; under stones and in leaf litter. Adult food, in the field (Europe): springtails. Predators: toads and frogs. Defense-mechanism: when alarmed, the adult escapes by running. Dispersal power. Wing-dimorphic: macropterous or brachypterous (incapable of flight). Flight records (Europe). Occasional in drift material, which indicates flight ability. Fast runner. Moderately favored by human activities. Collecting techniques. Turning stones; pitfall trapping. References. Lindroth, 1945, 1955a, 1961, 1963a, and 1985 (natural history); Erikstad, 1989 (ecology); Koch, 1989 (ecology); Luff, 1993 (biology). Notiophilus biguttatus (Fabricius, 1779) Ecology. Lowlands. Mostly forests (deciduous, coniferous): along clearings, roads, and edges; also adjoining cultivated fields and vacant lots. Shaded or open ground; moderately dry, often gravelly, sandy or loamy soil covered with leaf litter (mostly) or sparse vegetation. Mostly diurnal; active in the sunshine, mostly at midday; sheltering on cloudy or cool days under dead leaves or stones, among conifer needles, under pieces of wood and cardboard. Larvae often run on tree trunks in the sunshine (Europe). Gregarious. Biology. Seasonality: May-November. Tenerals: late June, July, and September. Adult

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and larval food, in the field (Europe): springtails, mites, flies, and other arthropods. Larval food, in captivity: springtails. The species has been reared on a diet of springtails. Adults hunt springtails by means of visual cues while larvae detect their aggregations using chemical cues. Defense-mechanism: when alarmed, the adult escapes by running. Dispersal power. Wing-polymorphic: macropterous to brachypterous (incapable of flight). Flight records (Europe). Occasional in drift material, which indicates flight ability. Fast runner. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Raking the leaf litter; pitfall trapping. References. Lindroth, 1945, 1955a, 1961, and 1963a (natural history); Grenslade, 1965 (ecology, biology); Bauer, 1975, 1979, and 1981 (biology); Honek, 1981 (dispersal power); Lindroth, 1985 (natural history); Loreau, 1985 (biology); Koch, 1989 (ecology); Ernsting et al. 1990 (biology); Luff, 1993 (biology). Notiophilus borealis T.W. Harris, 1869 Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, meadows, vacant lots, roadsides, forest clearings, and light forests. Open or slightly shaded ground; rather dry, gravelly or sandy soil covered with sparse or moderate short vegetation (e.g., moss). Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under stones and in moss. Solitary. Biology. Seasonality: June-September. Tenerals: August-September (NH, QU). Defense-mechanism: when alarmed, the adult escapes by running. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Fast runner. Collecting techniques. Turning stones; pitfall trapping; breaking dry moss. References. Lindroth, 1961 and 1963a (natural history). Notiophilus directus Casey, 1920 Ecology. Mountains. Moraines; vicinity of brooks and lakes. Ground open or shaded by shrubs or trees; dry, gravelly soil (sometimes sand-mixed), bare (mostly) or covered with tiny moss. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days among gravel or under pieces of wood. Biology. Seasonality: June-August. Tenerals: late July and early August (BC). Defense-mechanism: when alarmed, the adult escapes by running. Dispersal power. Wing-dimorphic: macropterous or brachypterous (incapable of flight). Fast runner. Collecting techniques. Raking the loose gravel; pitfall trapping. Reference. Lindroth, 1961 (natural history). Notiophilus intermedius Lindroth, 1954 Ecology. Lowlands. Banks of a river. A peaty site. Open ground; well-drained, usually dry, sandy soil covered with sparse vegetation. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under moss and lichen. Biology. Seasonality: MayJuly, September. Defense-mechanism: when alarmed, the adult escapes by running. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting techniques. Turning moss and lichen; pitfall trapping. References. Lindroth, 1955a, 1961, and 1963a (natural history).

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Notiophilus nemoralis Fall, 1906 Ecology. Mountains. Open coniferous forests (Abies, Picea). Open ground; dry soil, with bare spots surrounded by vegetation. Mostly diurnal; active in the sunshine among conifer needles; sheltering on cloudy or cool days in needle litter (mostly), among dead leaves, and in moss. Gregarious. Biology. Seasonality: July-September. Tenerals: August (NH, VT). Defense-mechanism: when alarmed, the adult escapes by running. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting techniques. Raking conifer needles and the leaf litter; pitfall trapping. Notiophilus nitens LeConte, 1857 Ecology. Lowlands. Grasslands, pastures, cultivated fields, and hills. Open ground; dry, sandy, gravelly or rocky soil covered with sparse vegetation. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under dead fallen plants or dry cattle dung. Biology. Seasonality: April-July. Tenerals: first half of May (OR), early June (BC). Defense-mechanism: when alarmed, the adult escapes by running. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Fast runner. Collecting techniques. Turning debris; pitfall trapping. Reference. Lindroth, 1961 (natural history). Notiophilus novemstriatus LeConte, 1848 Ecology. Lowlands and mountains. Light or open forests (deciduous, mixed, coniferous): near streams and ponds. Also forest edges and thickets. Open ground; dry soil, especially with green moss. Diurnal; active in the sunshine on moss, around stones and other debris; sheltering on cloudy or cool days under moss, dead leaves, sticks or bunches of grass. Gregarious. Biology. January-December. Adults found overwintering among moss, dead leaves and sticks. Adult food, in the field: dipterans (gnats). Defense-mechanism: when alarmed, the adult escapes by running and sheltering in soil crevices or beside stones. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (to artificial lights at night). Fast runner. Occasional climber (on trees). Collecting techniques. Turning moss carpets; raking the leaf litter and sticks on the ground; pitfall trapping. References. Frost, 1941 (ecology, biology); Kirk, 1969 and 1970 (ecology); Thompson and Allen, 1974 (biology); Erwin, 1981 (natural history). Notiophilus palustris (Duftschmid, 1812) Ecology. Lowlands. Edges of cultivated fields. Europe: drier zones of swamps (Alnus) covered with dense tall vegetation. Open ground; moderately moist, sandy soil covered with some vegetation. Diurnal; active in the sunshine. Biology. Seasonality: June-July. Tenerals: mid-July (PE). Adult food, in the field (Europe): springtails and mites. Defense-mechanism: when alarmed, the adult escapes by running. Dispersal power. Macropterous, probably capable of flight. Frequent in seashore drift material, which indi-

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cates flight ability. Fast runner. Collecting techniques. Pitfall trapping; turning drift material. References. Lindroth, 1945 and 1985 (natural history); Koch, 1989 (ecology). Notiophilus semiopacus Eschscholtz, 1833 Ecology. Mountains. Ridges and vicinity of snowfields. Open ground. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days at the base of stones and in ant nests. Biology. Seasonality: February, April-June, September, December. Defense-mechanism: when alarmed, the adult escapes by running. Dispersal power. Wing-dimorphic: macropterous (mostly) or brachypterous (incapable of flight). Occasional flier (to artificial lights at night). Fast runner. Collecting technique. Turning stones. Notiophilus semistriatus Say, 1823 Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, hills, abandoned fields, fencerows, roadsides, orchards; forest edges and clearings. Open or half-shaded ground; rather dry, gravelly (mostly), sandy, rocky or peaty soil covered with sparse low thin vegetation or patches of moss and lichen. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under reindeer moss (Cladonia), stones, dead leaves, and in clumps of broomsedges. Gregarious. Biology. Seasonality: January-December. Tenerals: mid-May and mid-June. Adults found overwintering under moss and dead leaves. Predators: thrushes. Defense-mechanism: when alarmed, the adult escapes by running. Dispersal power. Wing-dimorphic: macropterous or brachypterous (incapable of flight). Fast runner. Occasional climber (on trees). Collecting techniques. Pitfall trapping; lifting reindeer moss and stones. References. Lindroth, 1961 (natural history); Kirk, 1969 (ecology); Erwin, 1981 (natural history); Anderson et al. 1995 (ecology). Notiophilus sierranus Casey, 1920 Ecology. Mountains. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Notiophilus simulator Fall, 1906 Ecology. Mountains. Macrohabitat unknown. Open or slightly shaded ground, under bushes (Alnus); sandy bare soil, except for moss patches. Biology. Seasonality: JuneJuly, September. Dispersal power. Wing-dimorphic: macropterous or brachypterous (incapable of flight). Reference. Lindroth, 1961 (natural history). Notiophilus sylvaticus Eschscholtz, 1833 Ecology. Lowlands, mountains, subalpine and alpine zones. Light forests (mostly; e.g., Pinus) and meadows. Greenhouses (occasionally). Slightly shaded (mostly) or open ground; gravelly soil. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under the loose bark of fallen trees, under logs and stones, in moss, and among

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conifer needles. Gregarious. Biology. Seasonality: April, June-September. Tenerals: early June and August (BC). Defense-mechanism: when alarmed, the adult escapes by running. Dispersal power. Wing-dimorphic: macropterous or brachypterous (incapable of flight). Fast runner. Collecting techniques. Lifting loose bark; turning logs and stones; raking conifer needles; pitfall trapping. References. Lindroth, 1961 (natural history); Kavanaugh, 1992 (natural history). Olisthopus brevicornis Casey, 1913 Natural History. Unknown. Olisthopus filicornis Casey, 1913 Ecology. Macrohabitat unknown. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: May. Dispersal power. Wing-condition unknown. Attracted to artificial lights at night. Collecting technique. Raking the leaf litter. Olisthopus innuens Casey, 1913 Natural History. Unknown. Olisthopus iterans Casey, 1913 Natural History. Unknown. Olisthopus micans LeConte, 1848 Ecology. Lowlands. Low deciduous forests (e.g., Acer) and flood-plain forests: along ponds, marshes, lakes, rivers, brooks, and rills. Wet, muddy or clayish soil covered with thick leaf litter and humus, and sparse vegetation. Nocturnal; sheltering during the day in leaf litter and under sticks. Biology. Seasonality: January-August, November-December. Adults found overwintering in woods; in leaf litter. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Raking the leaf litter. References. Lindroth, 1966 (natural history); Kirk, 1969 (dispersal power); Erwin, 1981 (natural history); Will and Liebherr, 1997 (ecology). Olisthopus parmatus (Say, 1823) Ecology. Lowlands and mountains. Deciduous forests (mostly; e.g., Acer, Quercus), mixed forests (often along brooks), forest clearings, roads, and edges; hedges and swampforests. Shaded ground; moderately moist, often sandy or gravelly soil covered with thick leaf litter. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: January-December. Copulating pairs: early June (QU). Gravids: mid-May (QU). Tenerals: late July (IL) to mid-August (KY, ON). Adults found overwintering in woods and at the edge of woods; in leaf litter (mostly), in standing rotten tree-stumps, and under

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pieces of wood. Adult food, in captivity: dead carabids and tenebrionids, earthworms, and meat. Predators: toads. Parasites: Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult escapes by running and digging into the leaf litter. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Frequent in seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Raking the leaf litter; pitfall trapping; turning drift material. References. Lindroth, 1966 (natural history); Larochelle, 1974a (ecology); Erwin, 1981 (natural history). Olisthopus pusio Casey, 1913 Natural History. Unknown. Omophron americanum Dejean, 1831 Ecology. Lowlands and mountains. Beaches of rivers, brooks, lakes, ponds, pools, marshes, and estuaries; sand pits near bodies of water; salt meadows. Open ground; wet soil consisting of sand or sand mixed with clay, sometimes saline, bare or almost so. Mostly nocturnal; sheltering during the day in burrows dug in the soil (adult and larva). Gregarious. Biology. Seasonality: March-November. Copulating pairs: April-May (QU). Gravids: May-June (ON). Tenerals: July-September. Adults found overwintering on higher and drier ground, in the upper zone of river banks and sand pits, at the edge of woods and in woods; in the soil (mostly), under dead leaves, stones, logs, and bark. Adult food, in captivity: mealworms and meat. Larval food, in captivity: mealworms. Predators: frogs, toads, bluebirds, and starlings. Parfasites: Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult stridulates or escapes by running and burrowing; body colorpattern matching the soil. Dispersal power. Macropterous. Occasional flier (on humid, sunny days; to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Fast runner. Strong burrower. Collecting techniques. Pouring water over the ground; night collecting with a headlamp. References. Lindroth, 1955a, 1961, and 1963a (natural history); Rivard, 1964c (biology); Larochelle, 1972j (biology, dispersal power); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Omophron dentatum LeConte, 1852 Ecology. Lowlands and mountains. Beaches of rivers, brooks, and ponds. Open ground; wet, sandy or gravelly soil. Nocturnal; sheltering during the day in burrows dug in the soil (adult and larva). Biology. Seasonality: February-August. Defense-mechanism: when alarmed, the adult escapes by running and burrowing; body color-pattern matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Strong burrower. Collecting technique. Pouring water over the ground. Omophron gilae LeConte, 1852 Ecology. Uplands. Beaches of water bodies. Nocturnal; sheltering during the day in burrows dug in the soil. Gregarious. Biology. Seasonality: April-August. Defense-mech-

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anism: body color-pattern matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Strong burrower. Collecting technique. Pouring water over the ground. Omophron grossum Casey, 1909 Ecology. Lowlands. River beaches. Open ground; wet soil, sandy or mixed with mud. Nocturnal; sheltering during the day in burrows dug in the ground or in mole cricket burrows. Gregarious. Biology. Seasonality: June-August. Defense-mechanism: body color-pattern matching the soil. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Lago and Zucarro, 1984 (ecology, dispersal power). Omophron labiatum (Fabricius, 1801) Ecology. Coastal lowland. Beaches of ponds, lakes, and rivers; margins of ditches; sea beaches. Cultivated fields (e.g., corn). Open ground; wet, sandy, sometimes saline soil, bare or almost so. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in burrows dug in the soil. Gregarious. Biology. Seasonality: January-December. Larval food, in the field: grains and young shoots of corn. Economic importance: sometimes a pest of young corn. Predators: frogs and bluebirds. Defensemechanism: when alarmed, the adult escapes by running, burrowing or sheltering under debris; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Fast runner. Strong burrower. Collecting techniques. Pouring water over the ground; light trapping. References. Ulke, 1903 (dispersal power); Blatchley, 1910 (ecology); Leng, 1915 (ecology, dispersal power); Kirk, 1969 (ecology, dispersal power). Omophron nitidum LeConte, 1848 Ecology. Beaches of lakes and rivers. Open ground; wet, muddy soil, bare or covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil (mostly) and in leaf litter. Gregarious. Biology. Seasonality: April-October. Predators: bluebirds. Defense-mechanism: when alarmed, the adult escapes by running; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Fast runner. Strong burrower. Collecting techniques. Pouring water over the ground; light trapping. Omophron obliteratum G.H. Horn, 1870 Ecology. Uplands. River beaches. Open ground. Nocturnal; sheltering during the day in burrows dug in the soil. Gregarious. Biology. Seasonality: April-November. Defensemechanism: when alarmed, the adult escapes by running; body color-pattern matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Strong burrower. Collecting technique. Pouring water over the ground.

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Omophron ovale G.H. Horn, 1870 Ecology. Lowlands and mountains. Beaches of rivers (mostly), brooks, lakes, ponds, and reservoirs. Open ground; wet sandy, bare soil, often mixed with clay. Nocturnal; sheltering during the day in burrows dug in the soil (mostly), under stones and vegetal debris. Gregarious. Biology. Seasonality: April-October. Copulating pairs: early July (WY). Tenerals: first half of July (MT, WA, WY), October (BC). Defense-mechanism: when alarmed, the adult escapes by running; body color-pattern matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Lindroth, 1961 (natural history). Omophron robustum G.H. Horn, 1870 Ecology. Beaches of lakes, rivers, and brooks. Open ground; wet, sandy, sometimes alkaline soil. Nocturnal; sheltering during the day in burrows dug in the soil (mostly) and under debris. Gregarious. Biology. Seasonality: May-September. Tenerals: August (AB). Adult and larval food, in captivity: mealworms. Defense-mechanism: when alarmed, the adult escapes by running; body color-pattern matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Strong burrower. Collecting technique. Pouring water over the ground. References. Schwarz, 1876 (ecology); Lindroth, 1961 (natural history). Omophron solidum Casey, 1897 Ecology. Beaches of water bodies. Nocturnal; sheltering during the day in burrows dug in the soil. Gregarious. Biology. Seasonality: April-November. Defense-mechanism: when alarmed, the adult escapes by running; body color-pattern matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Strong burrower. Collecting technique. Pouring water over the ground. Omophron tessellatum Say, 1823 Ecology. Lowlands and mountains. Beaches of lakes, ponds, estuaries, slow brooks, and rivers; sea beaches; margins of pools (e.g., in sand pits) and ditches. Open ground; wet, sandy or clayish, sometimes saline soil, bare or almost so. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in burrows dug in the soil (adult and larva). Gregarious. Biology. Seasonality: April-November. Copulating pairs: June-July (QU). Tenerals: August-October. Adult food, in captivity: mealworms and dead carabids. Larval food, in captivity: mealworms. Predators: frogs and toads. Defensemechanism: when alarmed, the adult stridulates or escapes by running; body colorpattern matching the soil. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Strong burrower. Collecting techniques. Pouring water over the ground; night collecting with a headlamp. References. Benschoter and Cook, 1956 (biology); Lindroth, 1961 (natural history); Larochelle, 1972j (biology); Landry and Bousquet, 1984 (biology).

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Onota floridana G.H. Horn, 1881 Ecology. Lowlands. Forests (e.g., Quercus, Sabal). Shaded ground. Diurnal; active in the sunshine on the foliage of trees. Biology. Seasonality: February-March, November. Dispersal power. Wing-condition and running type unknown. Frequent climber (on trees). Collecting technique. Sweeping or beating trees. References. Schwarz, 1884 (ecology); Grossbeck, 1913 (ecology). Oodes amaroides Dejean, 1831 Ecology. Lowlands and uplands. Swamps; forested borders of marshes (small and large), lakes, and ponds; flood-plain forests. Shaded ground; soil inundated or, when not inundated, soft, very wet, muddy or silty, rich in organic debris. Diurnal and nocturnal; during the day, active under water along plant stems or among floating debris, or, sheltering in the soil, under dead leaves, fallen branches, and logs; also active on trees. Biology. Seasonality: January-October. Tenerals: March (rarely; FL), mid-June to mid-August (FL, IN, ME, ON, VA). Adults found overwintering (SC). Defense-mechanism: when alarmed, the adult escapes by diving into the water, crawling under floating debris, and disappearing from view. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (along plant stems, under the water surface; on trees). Excellent swimmer. Long-distance disperser; found at sea 30 km away from the mainland. Collecting techniques. Treading substrate with the feet; light trapping; using aquatic nets; using tanglefoot screens; using Japanese beetle traps; pyrethrum spraying. References. Ulke, 1903 (ecology); Leng, 1915 (ecology); Kirk, 1969 (ecology, dispersal power); Lindroth, 1969a (natural history); Lee, 1978 (ecology, dispersal power); Erwin, 1981 (natural history); Krinsky and Oliver, 1988 (ecology); Bousquet, 1996a (taxonomy, ecology). Oodes americanus Dejean, 1826 Ecology. Lowlands. Borders of cypress-swamps. Shaded ground; soft, wet, muddy soil covered with dead leaves and some vegetation. Nocturnal; sheltering during the day. Biology. Seasonality: April-May. Phoretic mites found on adult body. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Treading the ground with the feet; pouring water on the ground. Reference. Bousquet, 1996a (taxonomy, ecology). Oodes americanus americanus Dejean, 1826, see Oodes americanus Dejean, 1826. Reference. Bousquet, 1996a (taxonomy). Oodes americanus fluvialis LeConte, 1863, see Oodes fluvialis LeConte, 1863. Reference. Bousquet, 1996a (taxonomy).

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Oodes brevis Lindroth, 1957 Ecology. Lowlands and uplands. Swamps; flood-plain forests (e.g., Liriodendron, Liquidambar); forested borders of eutrophic marshes, lakes, ponds, pools, slow rivers, and brooks. Shaded ground; soil flooded, covered with dense tall vegetation (e.g., Typha) or, when not flooded, soft, very wet, muddy, rich in organic debris, covered with thick leaf litter and similar vegetation. Diurnal and nocturnal; during the day, active under water along plant stems or sheltering under dead leaves and logs. Biology. Seasonality: February-July, September. Tenerals: late May (DC), mid-July (VA), September (DC). Defense-mechanism: when alarmed, the adult escapes by diving into the water. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Frequent climber (on plants). Excellent swimmer. Collecting techniques. Treading the Typha-vegetation into the water; night collecting with a headlamp; pitfall trapping. References. Lindroth, 1969a (natural history); Anderson et al. 1995 (ecology); Bousquet, 1996a (taxonomy). Oodes fluvialis LeConte, 1863 Ecology. Lowlands and uplands. Borders of eutrophic marshes, ponds, and lakes; swamps. Shaded ground; soil, when flooded, covered with dense tall vegetation (e.g., Typha, Carex, Scirpus) or, when not flooded, soft, very wet, muddy or clayish, rich in organic debris, covered with similar vegetation. Diurnal and nocturnal; during the day, active under water along plant stems or sheltering under vegetal debris (e.g., Typhastems, pieces of wood), in moss, in clumps of grass, and under stones. Biology. Seasonality: February-December. Tenerals: late June (IN) to September (OH). Adults found overwintering on higher and drier ground, in woods, at the edge of woods, and in swamps; under and in logs, under stones, and in the soil. Adult food, in captivity: mealworms. Predators: ducks. Defense-mechanism: when alarmed, the adult emits a strong odor from the pygidial glands or escapes by swimming, diving into the water, staying submerged for a while before reappearing for oxygen. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Frequent climber (on plants). Excellent swimmer. Collecting techniques. Treading the Typha-vegetation into the water; light trapping. References. Lindroth, 1969a (natural history; as americanus Dejean); Attygalle et al. 1991 (biology, dispersal power; as as americanus Dejean); Will et al. 1995 (dispersal power); Bousquet, 1996a (taxonomy, ecology). [Oodes schaumi Chaudoir, 1882 References. Bousquet and Larochelle, 1993 (distribution); Bousquet, 1996a (taxonomy; absent from North America).] Oodinus alutaceus (Bates, 1882) Ecology. Borders of swamps and marshes. Shaded ground; soft, wet, muddy or silty soil covered with some vegetation (e.g., Sagittaria, Typha). Nocturnal; sheltering during the

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day. Biology. Seasonality: April, August, October, December. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Bousquet, 1996a (taxonomy, ecology). Oodinus piceus Motschulsky, 1865, see Oodinus pseudopiceus Bousquet, 1996. Reference. Bousquet, 1996a (taxonomy). Oodinus pseudopiceus Bousquet, 1996 Ecology. Lowlands. Swamp-forests and and borders of marshes. Shaded ground; very wet, muddy or silty soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day under pieces of wood. Biology. Seasonality: February-July, October, December. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Turning pieces of wood. Reference. Bousquet, 1996a (taxonomy). Ophonus puncticeps Stephens, 1828 Ecology. Lowlands and mountains. Vacant lots, old fields, meadows, cultivated fields, roadsides, gravel pits, sand pits, orchards (e.g., apple), and open forests. Open ground; dry soil consisting of clay, loam or gravel-mixed loam, covered with sparse tall vegetation (e.g., umbellifers). Mostly nocturnal; sometimes active during the day and often at night on umbellifers (Daucus carota Linné); usually sheltering during the day under stones, dead leaves, clumps of grass, pieces of wood, plant rosettes (Verbascum), and in the soil at the base of plants. Gregarious. Biology. Seasonality: June-October. Tenerals: early June (CT, ON, VT). Adult and larval food, in the field (Europe): mostly umbellifer seeds (Daucus) and pollen; also snails. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Frequent climber (on plants, e.g., Daucus). Strong burrower. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Turning stones; sweeping plants; pitfall trapping; light trapping. References. Larsson, 1939 (biology). Lindroth, 1945 and 1968 (natural history); Brandmayr Zetto and Brandmayr, 1975 (biology); Steffens and Davidson, 1979 (dispersal power); Brandmayr et al. 1980 (taxonomy); Lindroth, 1986 (natural history); Krinsky and Oliver, 1988 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology); Will et al. 1995 (dispersal power); Kataev, 1996 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Ophonus rufibarbis (Fabricius, 1792) Ecology. Lowlands. Vacant lots, meadows, and lawns. Europe: meadows, parks, gardens, cultivated fields, and open woods. Open ground; moderately dry soil, usually clayish, humus-mixed, covered with some vegetation (e.g., grass, herbs). Nocturnal; sheltering during the day under stones and bricks. Gregarious (Europe). Biology. Seasonality: May-August. Adult food, in the field: umbellifer seeds (Anthriscus; Europe).

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Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Regular climber (on plants; Europe). Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Turning stones; sweeping plants; pitfall trapping. References. Lindroth, 1945 and 1968 (natural history); Brandmayr et al. 1980 (taxonomy); Lindroth, 1986 (natural history); Koch, 1989 (ecology); Luff. 1993 (biology); Kataev, 1996 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Ophryogaster flohri Bates, 1882 Ecology. Mountains. Riparian forests. Moist or wet soil. Nocturnal; sheltering during the day in leaf litter and under fallen fruits (e.g., palm fruit). Biology. Unknown. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Collecting technique. Raking the leaf litter. References. Bousquet, 1999 (dispersal power); Ball and Bousquet, 2001 (ecology). Opisthius richardsoni Kirby, 1837 Ecology. Lowlands and mountains. Banks of cold, swift rivers and large brooks. Close to water. Open ground; well-drained, soft, wet, bare sandy soil, somewhat mixed with clay. Diurnal, active in the sunshine; sheltering on cloudy or cool days under stones. Gregarious. Associated with staphylinids (Bledius). Larval habitat: burrows dug in the soil. Biology. Seasonality: June-September. Tenerals: early July to August. Adult and larval food, in captivity: mealworms. Defense-mechanism: when alarmed, the adult escapes by flying or running; when disturbed, the larva performs sudden jumps. Dispersal power. Macropterous. Frequent flier (in the warm sunshine). Fast runner. Collecting techniques. Turning stones; pouring water over the ground. References. Lindroth, 1960 and 1961 (natural history). Oxycrepis (Stolonis) intercepta (Chaudoir, 1873) Ecology. Lowlands and mountains. Mexico: tropical forests (gallery; evergreen seasonal; tropical deciduous scrub; palm), along dry stream beds. Shaded ground; moist soil. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: February, JuneOctober (USA-Mexico). Defense-mechanism: the white spots of the antennae contrasting with the dark colored body may distract predators. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Raking the leaf litter. References. Allen and Ball, 1980 (ecology); Ball and Bousquet, 2001 (ecology). Oxydrepanus rufus (Putzeys, 1846) Ecology. Lowlands. Swamps (hammock forests) and pool margins. Wet soil, rich in organic debris. Nocturnal; sheltering during the day in leaf litter or in floating vegetation. Biology. Seasonality: June. Dispersal power. Macropterous, probably capable of flight. Slow runner. Collecting techniques. Raking the leaf litter; treading the floating vegetation into water. Reference. Nichols, 1988 (ecology).

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Oxypselaphus pusillus (LeConte, 1854) (Agonum puncticeps (Casey, 1920)) Ecology. Lowlands and mountains. Borders of eutrophic marshes, ponds, lakes, slow rivers and brooks; swamps, swamp-forests, flood-plain forests, wet meadows, and beaver houses. Shaded ground; moist muddy or clayish soil, rich in organic debris, covered with some vegetation. Nocturnal; sheltering during the day in leaf litter (mostly), under pieces of wood, logs, and stones. Gregarious (in winter). Biology. Seasonality: JanuaryDecember. Tenerals: April (rarely; QU), June-November. Adults found overwintering on higher and drier ground, in the upper banks of marshes, rivers, and swamps, in woods, at the edge of woods, in hawthorn hedges, beaver houses, dunes, and vacant lots; among and under dead leaves (mostly), stones, logs, the loose bark of logs, fallen trees, and standing tree-stumps, in rotten logs, tree-stumps, fallen branches, and in the soil (as deep as 15 cm). Active at low temperatures under snow in October-November. Adult food, in captivity: dipterous eggs (Aedes) and mealworms. Larval food, in captivity: mealworms. Predators: toads. Parasites: Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult may emit a pop noise accompanied by a strong Brachinus-like smoke from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Frequent climber (on trees and plants). Favored by spring floods along large rivers. Collecting techniques. Raking the leaf litter; sweeping or beating the vegetation. References. Blatchley, 1910 (biology; as Platynus pusillus LeConte); Lindroth, 1966 (natural history; as puncticeps (Casey)). Ozaena lemoulti Bänninger, 1932 Ecology. Mountains. Forests. Biology. Seasonality: June-July. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Pachyteles (Pachyteles) gyllenhalii (Dejean, 1825) Ecology. Lowlands and mountains. Forests. Dry soil. Nocturnal; sheltering during the day under the loose bark of dead tree trunks and in rotting saguaro cactus. Biology. Seasonality: January-February, July-August, October-December. Parasites: Laboulbeniales (Fungi). Dispersal power. Wing-condition unknown. Attracted to artificial lights at night. Frequent climber (on trees). Collecting technique. Lifting the loose bark of dead tree trunks. Reference. Ball and McCleve, 1990 (natural history). Pachyteles (Goniotropis) kuntzeni (Bänninger, 1927) Ecology. Mountains. Forests (Quercus). Dry soil. Nocturnal; sheltering on dead trees. Biology. Seasonality: July-August, November. Dispersal power. Wing-condition unknown. Attracted to artificial lights at night. Occasional climber (on trees). Reference. Ball and McCleve, 1990 (natural history).

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Pachyteles (Goniotropis) parca LeConte, 1884 Ecology. Mountains. Vicinity of forests (Quercus, Pinus). Biology. Seasonality: June-August. Dispersal power. Wing-condition unknown. Frequently attracted to artificial lights at night. Occasional climber (on trees). Reference. Ball and McCleve, 1990 (natural history). Panagaeus crucigerus Say, 1823 Ecology. Lowlands. Borders of saline marshes. Meadows near the sea (often close to tidal streams). Open ground; wet soil consisting of loose sand, often saline. Nocturnal; sheltering during the day in the soil, at the base of plants (e.g., Hudsonia), and under pieces of wood and stones. Biology. Seasonality: January-December. Tenerals: September (NJ). Adults found overwintering in ant nests. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strong burrower. Collecting techniques. Inspecting ant nests in winter; pitfall trapping; light trapping. References. Schwarz, 1890 (biology); Leng, 1915 (dispersal power); Löding, 1945 (ecology). Panagaeus fasciatus Say, 1823 Ecology. Lowlands. Open forests (mostly; e.g., Pinus), grassy hills, cypress-swamp edges, railway embankments, and river banks. Open ground; dry sandy soil covered with some vegetation (e.g., grass). Mostly nocturnal; sometimes active during the day on forest paths; usually sheltering in leaf litter, under stones, pieces of wood, and logs, in ant nests and clumps of broomsedges. Biology. Seasonality: January-December. Adults found overwintering in ant nests (frequently), under stones, and logs. Predators: crows. Parasites: Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult emits a Brachinus-like smoke from the pygidial glands. Dispersal power. Macropterous. Frequent flier (often swarming to artificial lights at night). Frequent in sea- and lakeshore drift material which indicates flight ability. Moderate runner. Occasional climber (on trees). Collecting techniques. Pitfall trapping; raking the leaf litter; turning stones and wood debris; light trapping; inspecting ant nests in winter; turning drift material. References. Hamilton, 1888b and 1895 (ecology, biology); Ulke, 1903 (ecology, dispersal power); Blatchley, 1910 (ecology, biology); Smith, 1910 (ecology); Leng, 1915 (ecology, dispersal power); Löding, 1945 (ecology); Kirk, 1969 (ecology, dispersal power) and 1970 (ecology). Panagaeus sallei Chaudoir, 1862 Ecology. Open woods. Open ground; dry soil. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: April, June-September. Defense-mechanism: body color-pattern offers a striking protective resemblance to mutilids. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Raking the leaf litter.

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Paranchus albipes (Fabricius, 1796) Ecology. Coastal lowlands. Steep slopes along the seashore; shores of lakes and ponds; banks of rivers and brooks. Open or slightly shaded ground; wet clay soil, bare or covered with sparse vegetation. Nocturnal; sheltering during the day under stones (mostly) and vegetal debris. Biology. Seasonality: May-September. Tenerals: early July (NS) to late August (NF). Dispersal power. Submacropterous, probably incapable of flight. Moderate runner. Collecting technique. Turning stones. References. Larsson, 1939 (biology); Lindroth, 1945, 1955a, 1963a (as ruficorne (Goeze)) and 1966 (natural history); ° Kurka, 1976 (biology; as ruficorne (Goeze)); Lindroth, 1986 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology). Paratachys aeneipennis (Motschulsky, 1862) Natural History. Unknown. Paratachys albipes (LeConte, 1863) Ecology. Lowlands. Pond margins. Moist, muddy soil. Nocturnal; sheltering during the day in leaf litter and in the soil. Biology. Seasonality: April-August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Collecting techniques. Pouring water over the ground; raking the leaf litter. Paratachys austinicus Casey, 1918 Ecology. Unknown. Biology. Seasonality: July-September. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Reference. Kirk, 1975c (dispersal power) Paratachys columbiensis (Hayward, 1900) Ecology. Lowlands. Lake shores; margins of saltwater bodies. Nocturnal; sheltering during the day in debris. Biology. Seasonality: March-December. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Fast runner. Collecting techniques. Pouring water over the ground; raking debris. Reference. Leng, 1915 (ecology). Paratachys edax (LeConte, 1852) Ecology. Margins of streams, lakes, and reservoirs. Open ground; soft, wet, clayish or muddy, bare soil. Nocturnal; sheltering during the day in cracks in the soil. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Pouring water over the ground. Reference. Hatch, 1953 (ecology). Paratachys hyalinus (Casey, 1918) Natural History. Unknown.

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Paratachys oblitus (Casey, 1918) (Tachys obliquus Casey, 1918) Ecology. Lowlands and mountains. Marshy edges of ponds, lakes, slow rivers and brooks; borders of eutrophic marshes. Open or shaded ground; wet, clayish or muddy soil, sometimes with organic debris, covered with sparse or moderate vegetation. Crepuscular and nocturnal; sheltering during the day under dead leaves, fallen dead plants, at the base of plant tufts (e.g., Carex), and in soil crevices. Biology. Seasonality: AprilNovember. Tenerals: early June to late August. Adults found overwintering on higher and drier ground, in the upper banks of marshes and rivers, and at the edge of woods; in leaf litter. Dispersal power. Macropterous. Frequent flier (before sunset; to artificial lights at night; to artifical lights in captivity). Fast runner. Favored by spring floods along large rivers. Collecting techniques. Pouring water over the ground; treading the soil with the feet; raking the leaf litter; light trapping. References. Larochelle, 1971b (ecology; as obliquus (Casey)); Erwin, 1981 (natural history; as (obliquus Casey)); Will et al. 1995 (dispersal power). Paratachys potomaca Erwin, 1981 Ecology. Lowlands. Stream and pond margins. Sandy soil mixed with clay, covered with depressed vegetation. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: March-May. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Raking the leaf litter. Reference. Erwin, 1981 (natural history). Paratachys proximus (Say, 1823) Ecology. Lowlands and mountains. Borders of slow rivers and brooks, pools, ponds, and eutrophic marshes; beaver houses. Caves (occasionally). Open or half-shaded ground; moist, clayish, muddy, or sandy soil (clay- or silt-mixed) covered with some vegetation. Mostly crepuscular and nocturnal; sometimes active in the sunshine; usually sheltering during the day under dead leaves, stones, and pieces of wood. Biology. Seasonality: February-November. Tenerals: July (MD). Adults found overwintering under pieces of wood and stones. Dispersal power. Macropterous. Frequent flier (before sunset; to artificial lights after dusk, at night; to artifical lights in captivity). Occasional in lakeshore drift material, which indicates flight ability. Fast runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet; raking the leaf litter; sifting vegetal debris; light trapping. References. Lindroth, 1966 (natural history); Kirk, 1975c (dispersal power); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Paratachys pumilus (Dejean, 1831) Ecology. Lowlands and mountains. Borders of lakes, ponds, slow rivers and brooks; roadside ditches, mud flats, wet meadows, and low forests. Open ground; wet, clayish, muddy or silty soil covered with some vegetation. Mostly crepuscular and nocturnal;

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sometimes active in the sunshine; usually sheltering during the day mostly under dead leaves and fallen plants, also in cracks in the soil and in the ground at the base of plants (e.g., grass). Biology. Seasonality: January-November. Tenerals: February (rarely; AL), July (AL, TX, WV). Adults found overwintering in mud flats; under fallen dead plants. Dispersal power. Wing-dimorphic: macropterous (capable of flight) or brachypterous (incapable of flight). Frequent flier (at sunset; to artificial lights at night). Occasional in drift material, which indicates flight ability. Fast runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet; raking and sifting the leaf litter; turning fallen plants; light trapping. References. Leng, 1915 (ecology); Erwin, 1981 (natural history; as corruscus (LeConte)). Paratachys rectangulus (Notman, 1919) Natural History. Unknown. Paratachys rhodeanus (Casey, 1918) Ecology. Lowlands. Borders of slow rivers, lake shores, and eutrophic marshes. Open ground; moist, clayish or muddy soil (sometimes sand-mixed) covered with some vegetation. Crepuscular and nocturnal; sheltering during the day under well-embedded stones or in leaf litter at the base of plants. Biology. Seasonality: April-June, August, October. Dispersal power. Macropterous. Frequent flier (at sunset; to artificial lights at night; to artifical lights in captivity). Fast runner. Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet; turning embedded stones; raking the leaf litter; light trapping. References. Lindroth, 1966 (natural history); Landry, 1975b (ecology). Paratachys sagax (Casey, 1918) Natural History. Unknown. Paratachys scitulus (LeConte, 1848) Ecology. Lowlands and mountains. Borders of rivers, brooks, ponds, lakes, and eutrophic marshes; beaver houses. Open ground; moist or wet, clayish (sometimes sand-mixed), muddy or peaty soil, bare or covered with sparse vegetation. Mostly crepuscular and nocturnal; sometimes active in the sunshine; usually sheltering during the day under dead leaves, fallen dead plants, stones, pieces of wood, in cracks in the soil, under silt cakes, and among plant roots. Biology. Seasonality: March-November. Tenerals: JuneAugust; mostly after mid-July. Adult food, in the field: lepidopterous caterpillars. Defense-mechanism: when alarmed, the adult escapes by flying. Dispersal power. Macropterous. Frequent flier (when alarmed; in the sunshine; at sunset; to artificial lights at night). Fast runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet; raking the leaf litter; light trapping. References. Lindroth, 1966 (natural history); Kirk, 1975c (dispersal power); Erwin, 1981 (natural history).

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Paratachys sequax (LeConte, 1848) Natural History. Unknown. Paratachys spadix (Casey, 1918) Natural History. Unknown. Paratachys umbripennis (Chaudoir, 1868) Natural History. Unknown. Paratachys ventricosus (LeConte, 1863) Ecology. Lowlands. Damp places. Margins of a flowing well in a dense wet hammock. Nocturnal; sheltering during the day under pieces of wood. Biology. Seasonality: March, May, August, December. Dispersal power. Wing-condition unknown. Fast runner. Paratachys vernilis (Casey, 1918) Natural History. Unknown. Paratachys vorax (LeConte, 1852) Ecology. River banks. Wet soil. Biology. Seasonality: April-May, July. Dispersal power. Wing-condition unknown. Moderate runner. Pasimachus (Pasimachus) californicus Chaudoir, 1850 Ecology. Lowlands and uplands. Grasslands, city vacant lots, vicinity of bluffs, and open forests. Open or half-shaded ground; dry, sandy (mostly), clayish or muddy soil covered with sparse vegetation (e.g., grass, yuccas). Nocturnal; sheltering during the day in burrows dug in the soil under stones, dry cattle dung, and other debris. Biology. Seasonality: January-November. Adults found overwintering under stones. Adult food, in the field: insect larvae (mostly lepidopterous caterpillars), grasshopper eggs and adults. Economic importance: may play an important role in the control of lepidopterous caterpillars. Defense-mechanism: when alarmed, the adult emits a defensive fluid from the pygidial glands or bites strongly, sometimes drawing blood. Predators: bats. Dispersal power. Brachypterous, incapable of flight. Slow runner. Strong burrower. Collecting techniques. Pitfall trapping; turning stones. Reference. McCullough, 1969a (biology). Pasimachus (Pasimachus) depressus (Fabricius, 1787) Ecology. Lowlands and uplands. Grasslands, cultivated fields (e.g., corn), hill slopes, and open forests. Open ground; dry, sandy soil covered with sparse vegetation. Noctur-

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nal; sheltering during the day in burrows dug in the soil under stones, logs, and dead leaves. Solitary or in pairs. Biology. Seasonality: January, April-October, December. Adults found overwintering (SC). Copulating pairs: June (MD). Tenerals: June (MD). Adult food, in the field: lepidopterous caterpillars (armyworms, cutworms). Economic importance: may play an important role in the control of lepidopterous caterpillars. Predators: crows, grackles, grosbeaks, and woodpeckers. Dispersal power. Brachypterous, incapable of flight. Slow runner. Strong burrower. Collecting techniques. Pitfall trapping; turning stones. References. Ulke, 1903 (ecology); Erwin, 1981 (natural history). Pasimachus (Pasimachus) duplicatus LeConte, 1853 Ecology. Lowlands and uplands. Grasslands. Open ground; dry, muddy soil covered with sparse vegetation (e.g., plants, yuccas). Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: April-November. Dispersal power. Brachypterous, incapable of flight. Slow runner. Strong burrower. Collecting technique. Pitfall trapping. Reference. McCullough, 1969a (biology). Pasimachus (Pasimachus) elongatus LeConte, 1846 Ecology. Lowlands and uplands. Grasslands, cultivated fields (e.g., wheat, alfalfa, oat, flax, corn), rangelands, meadows, pastures, roadsides, railway embankments, and open forests. Open ground; well-drained, dry, sandy soil covered with some vegetation. Often associated with the presence of Brome Grass (Bromus inermis Leysser). Nocturnal; sheltering during the day in burrows dug in loose soil, in cracks in the ground, under dry cow dung, stones, railway ties, and logs. Solitary. Associated with False wireworms (tenebrionids). Larval habitat: in burrows dug in the soil among plants or shrubs, in leaf litter and humus, and under stones; the larva waits at its burrow entrance, like a tiger beetle larva, ambushing passing insects. Biology. Seasonality: April-October. Copulating pairs: June (CO, SD), August (WY); coitus lasting 90-105 minutes. Oviposition: late June to August (SD). Adults found overwintering in burrows dug in the soil (as deep as 30 cm). Life cycle: 2 years. Adult food: lepidopterous caterpillars, false wireworm larvae (frequently), other tenebrionids, chrysomelids, carabids, acridids, and other insects (in the field); tenebrionids, mealworms, meat, boiled yolk of egg, peach, and cheese (in captivity). Larval food: weevils and other insects (in the field); mealworms and soft-bodied insects (in captivity). Economic importance: may play an important role in the control of crop pests. Predators: toads, foxes, and raccoons. Parasites: podapolipid mites. Defense-mechanism: when alarmed, the adult emits a smelling defensive fluid from the pygidial glands, spreads its mandibles in a threatening position, or bites vigorously. Dispersal power. Brachypterous, incapable of flight. Slow runner. Strong burrower; less fossorial than other Pasimachus species. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning dry cow dung and stones. References. Lindroth, 1961 (ecology, dispersal power); Cress, 1966 (biology); Bell, 1971 (ecology); Cress and Lawson, 1971 (biology); McCullough, 1972b (biology); Calkins and Kirk, 1974 (biology); Thompson and Allen, 1974 (ecology, biology); Kirk, 1975c (ecology, biology); Allen and Thompson, 1977 (ecology, biology); Lavigne, 1977 (natural history); Epstein and Kulman, 1990 (ecology, biology); McIntyre, 1995 (biology).

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Pasimachus (Pasimachus) marginatus (Fabricius, 1787) Ecology. Lowlands. Open forests (e.g., Pinus), orchards (e.g., peach, plum), grasslands, and pastures. Open or half-shaded ground; dry, sandy soil covered with sparse vegetation. Nocturnal; sheltering during the day under the loose bark of dead trees, and in burrows dug in the soil under logs and boards. Solitary. Biology. Seasonality: FebruaryDecember. Predators: crows and alligators. Dispersal power. Brachypterous, incapable of flight. Slow runner. Strong burrower. Collecting techniques. Pitfall trapping; baiting traps with malt; ground-based light trapping; lifting loose bark; turning wood debris. Reference. Leng, 1915 (ecology, biology). Pasimachus (Pasimachus) obsoletus LeConte, 1846 Ecology. Lowlands and uplands. Prairies and hills. Open ground; dry, sandy soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: February-December. Dispersal power. Brachypterous, incapable of flight. Slow runner. Strong burrower. Collecting technique. Pitfall trapping. Reference. Kirk, 1975c (ecology). Pasimachus (Pasimachus) punctulatus Haldeman, 1843 Ecology. Lowlands. Grasslands, pastures, wastelands, and open forests (deciduous, mixed). Open ground; dry, sandy soil covered with some vegetation (e.g., grass). Mostly diurnal. Solitary. Biology. Seasonality: March-December. Copulating pairs: July (OH), September (TX). Adult food, in the field: grasshopper eggs and adults. Larvae are cannibalistic in captivity. Defense-mechanism: when alarmed, the adult may emit a fine acid-tasting spray of burning water-colored fluid from the abdomen. Dispersal power. Brachypterous, incapable of flight. Slow runner. Collecting technique. Pitfall trapping. References. Fattig, 1937 (biology); Alexander, 1959 (biology); Nelson, 1969 and 1970 (ecology, biology). Pasimachus (Pasimachus) strenuus LeConte, 1874 Ecology. Lowlands. Habitats covered with sparse trees (Pinus-Quercus): sand hills, scrublands, and open forests. Open or slightly shaded ground; dry, sandy soil covered with sparse vegetation (e.g., grass, Saw palmettos). Diurnal and nocturnal; active in the sunshine or sheltering during the day under pieces of wood. Biology. Seasonality: JanuaryDecember. Copulating pairs: spring and autumn (FL). Adult food, in the field: scarabaeids. Parasites: tachinid flies (Sitophaga). Dispersal power. Brachypterous, incapable of flight. Slow runner. Collecting techniques. Pitfall trapping; baiting traps with malt; turning pieces of wood. References. Leng, 1915 (ecology); McCoy and Witz, 1994 (biology). Pasimachus (Pasimachus) sublaevis (Palisot de Beauvois, 1811) Ecology. Lowlands. Open forests (e.g., Pinus), pastures, dunes, and beaches. Shaded or open ground; dry, sandy soil. Nocturnal; sheltering during the day in burrows dug in the

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soil under logs, stones, boards, and dry cow and horse dung. Solitary. Biology. Seasonality: January-November. Adults found overwintering in burrows dug in the soil (as deep as 15 cm) and under logs. Adult food, in the field: lepidopterous caterpillars and other insect larvae. Dispersal power. Brachypterous, incapable of flight. Slow runner. Strong burrower. Collecting techniques. Pitfall trapping; baiting traps with syrup; ground-based light trapping; turning logs and stones. References. Blatchley, 1910 (ecology); Leng, 1915 (ecology, biology); Kirk, 1969 (dispersal power); Miliotis, 1974 (ecology). Pasimachus (Pasimachus) subsulcatus Say, 1823 Ecology. Lowlands. Sandhills covered with sparse trees (Pinus-Quercus), scrublands, and open forests (e.g., Quercus). Open or slightly shaded ground; dry, sandy soil covered with sparse vegetation (e.g., grass, Saw palmettos). Mostly nocturnal; sometimes active in the daytime; normally sheltering during the day in burrows dug in the soil. Biology. Seasonality: January-December. Copulating pairs: spring and autumn (FL). Parasites: tachinid flies (Sitophaga) and nematodes. Defense-mechanism: when alarmed, the adult emits a burning spray from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Slow runner. Strong burrower. Collecting techniques. Pitfall trapping; baiting traps with malt. References. Kirk, 1969 (ecology); Harris and Whitcomb, 1971 (ecology, biology); McCoy, 1987 (ecology, biology); Davidson et al. 1989 (biology); Witz and Mushinsky, 1989 (biology); McCoy and Witz, 1994 (biology). Pasimachus (Pasimachus) viridans LeConte, 1858 Ecology. Uplands. Biology. Seasonality: July-September. Dispersal power. Brachypterous, incapable of flight. Patrobus (Patrobus) fossifrons (Eschscholtz, 1823) Ecology. Lowlands and mountains. Margins of sloughs, marshes, bogs, small pools, slow rivers and brooks; mud flats. Close to water. Open ground; moist, often muddy, clayish or gravelly soil, bare or covered with sparse vegetation. Nocturnal; sheltering during the day under logs, dead leaves, and fallen grass. Biology. Seasonality: JulyAugust. Tenerals: August (BC). Adult and larval food, in captivity: mealworms. Dispersal power. Wing-dimorphic: macropterous or brachypterous (incapable of flight). Moderate runner. Collecting techniques. Pitfall trapping; turning logs; raking the leaf litter. Reference. Lindroth, 1961 (natural history). Patrobus (Patrobus) foveocollis (Eschscholtz, 1823) Ecology. Lowlands, mountains, subalpine and alpine zones. Forests, forest clearings and edges, thickets and bushes (e.g., Alnus, Salix, dwarf Betula), moraines, cultivated fields, sedge meadows, alpine meadows; upper borders of marshes, swamps, lakes, ponds, and rivers. Slightly shaded ground; moderately moist soil, rich in vegetal debris (e.g., dead leaves). Nocturnal; sheltering during the day in leaf litter (mostly), under

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and in logs, under stones, and in moss (Sphagnum). Biology. Seasonality: May-November. Gravids: late June and early July (QU). Tenerals: June-September. Adult food, in captivity: mealworms and shrimps. Larval food, in captivity: mealworms. Predators: toads. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (to artificial lights at light). Moderate runner. Strong burrower. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Lindroth, 1955a, 1961, and 1963a (natural history); Carter, 1981 (biology). Patrobus (Patrobus) lecontei Chaudoir, 1871 Ecology. Lowlands and mountains. Borders of small pools, ponds, sloughs, marshes, swamps, slow rivers and brooks; thickets (e.g., Salix) and cultivated fields (e.g., corn, barley). Open or slightly shaded ground; firm, moist or wet, clay soil covered with some vegetation (e.g., Carex, Agrostis). Nocturnal; sheltering during the day under vegetal debris (mostly; e.g., logs) and stones. Biology. Seasonality: June-November. Gravids: mid-July to mid-August (AB, QU). Tenerals: June to early August. Adult food, in captivity: mealworms and shrimps. Larval food, in captivity: mealworms. Phoretic mites found on adult body. Dispersal power. Wing-dimorphic: macropterous or brachypterous (incapable of flight). Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; turning logs and stones. References. Lindroth, 1955a, 1961, and 1963a (natural history); Kirk, 1975c (dispersal power); Carter, 1981 (ecology, biology). Patrobus (Neopatrobus) longicornis (Say, 1823) Ecology. Lowlands and mountains. Borders of rivers, brooks, lakes, ponds, pools, marshes, irrigation canals, and roadside ditches; mud flats, swamps, swamp-forests, low woods; orchards (e.g., apple, peach), meadows, pastures, cultivated fields (e.g., corn, tobacco, carrot), vacant lots, gravel pits, and sand pits. Caves (occasionally). Often close to water. Open or shaded ground; moist or wet, usually clayish soil covered with dense vegetation. Nocturnal; sheltering during the day under dead leaves, dead fallen plants, logs, pieces of wood, branches, and other debris. Biology. Seasonality: January, MarchDecember. Gravids: early July to mid-October (QU). Tenerals: May-September. Adults found overwintering on higher and drier ground, in the upper banks of marshes and rivers, at the edge of woods, in woods, on hills, and in meadows; in burrows dug in the soil (as deep as 25 cm), under stones, logs, and the loose bark of fallen trees. Adult food, in the field: 2/3 vegetal matter, 1/3 animal matter (e.g., lepidopterous caterpillars, aphids). Adult and larval food, in captivity: mealworms. Predators: frogs, toads, starlings, crows, and shrews. Parasites: parantennulid mites. Phoretic mites found on adult body. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Frequent flier (to artificial lights at night). Moderate runner. Moderately favored by human activities. Collecting techniques. Raking the leaf litter; treading the soil with the feet; pitfall trapping; baiting traps with meat; night collecting with a headlamp. References. Blatchley, 1910 (ecology); Darlington, 1936 (dispersal power); Lindroth, 1955a, 1961, and 1963a (natural history); Larochelle, 1973d (dispersal power);

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Kirk, 1975c (ecology, dispersal power); Erwin, 1981 (natural history); Levesque and Levesque, 1987 (biology); Will et al. 1995 (dispersal power). Patrobus (Patrobus) septentrionis Dejean, 1828 Ecology. Lowlands, mountains, subalpine and alpine zones. Borders of lakes, ponds, pools, marshes, cold bogs, and slow rivers. Meadows. Open country, away from water (Aleutian Islands). Open ground; moist, somewhat clayish soil covered with some vegetation (e.g., grass, Carex, sedges, moss). Nocturnal; sheltering during the day under logs and dead leaves. Biology. Seasonality: June-October. Tenerals: June-July. Adult food, in the field (Europe): insects lying on snow. Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Occasional flier (to artificial lights). Moderate runner. Collecting techniques. Treading the soil with the feet; turning logs; raking the leaf litter; pitfall trapping. References. Lindroth, 1945, 1955a, 1961, and 1963a (natural history); Carter, 1981 (biology); Bakken, 1985 (biology); Lindroth, 1985 (natural history); Dearborne and Donahue, 1993 (dispersal power); Luff, 1993 (biology). Patrobus (Patrobus) stygicus Chaudoir, 1871 Ecology. Lowlands and mountains. Borders of marshes, bogs, and quagmires; swamps; marshy borders of ponds, lakes, and slow rivers. Open ground; soft, wet, often peaty soil covered with tall dense vegetation (e.g., Carex, Comarum, Menyanthes, moss). Nocturnal; sheltering during the day under vegetal debris (e.g., dead leaves) and in moss. Biology. Seasonality: June-September. Gravids: mid-June to mid-July (QU). Tenerals: June to early August. Adult and larval food, in captivity: mealworms and shrimps. Dispersal power. Macropterous. Occasional flier (to artificial lights at night; to artifical lights in captivity). Moderate runner. Occasional climber (on plants). Good swimmer. Collecting techniques. Treading the vegetation into the water; raking the leaf litter; pitfall trapping; sweeping plants. References. Lindroth, 1955a, 1961, and 1963a (natural history); Carter, 1981 (ecology, biology). Pelmatellus (Pelmatellus) obtusus Bates, 1882 Ecology. Mountains. Cultivated fields, meadows, canyons, and forest openings (e.g., Quercus, Pinus). Open ground. Nocturnal; sheltering during the day under stones and other debris. Biology. Seasonality: March-September. Dispersal power. Wing-condition unknown. Moderate runner. Collecting technique. Turning stones. Pelmatellus (Pelmatellus) stenolophoides parallelus Goulet, 1974 Ecology. Mountains. Biology. Seasonality: June. Dispersal power. Unknown. Pelmatellus (Pelmatellus) stenolophoides stenolophoides Bates, 1882 Ecology. Cultivated fields, meadows, roadsides, and open forests (e.g., Quercus, Pinus). Biology. Unknown. Dispersal power. Macropterous. Occasional flier. Moderate runner.

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Pelophila borealis (Paykull, 1790) Ecology. Lowlands, mountains, subalpine and alpine zones. Borders of slow rivers, lakes, eutrophic ponds, pools, and marshes; swamps. Close to water. Open ground; wet, soft, muddy soil, rich in organic debris, or gravelly soil often covered with dense vegetation (e.g., Juncus, Carex, Eriophorum), sometimes with moss carpets. Nocturnal; sheltering during the day under vegetal debris, stones, and in moss carpets. Able to withstand freezing temperatures (as low as –5ºC). Larval habitat: under stones. Biology. Seasonality: June-August. Gravids: late June and early July (QU). Eggs are laid in the ground. Tenerals: early July (QU). Adults found overwintering in cavities in the soil (Europe). Adult and larval food, in the field (Europe): worms, insect larvae, and carrion. Adult food, in captivity: mealworms. Larval food, in captivity: mealworms and meat. Defense-mechanism: when alarmed, the adult emits a strong smell; when disturbed, the larva runs about rapidly. Dispersal power. Macropterous. Flight records (Europe). Fast runner. Excellent swimmer. Collecting techniques. Treading the vegetation into the water; turning vegetal debris and stones. References. Johnson and Carpenter, 1898 (natural history); Lindroth, 1945, 1955a, 1961, and 1963a (natural history); Östbye and Sömme, 1972 (biology); Conradi-Larsen and Sömme, 1973a and 1973b (biology); Sömme, 1974 (biology); Luff, 1993 (biology). Pelophila rudis (LeConte, 1863) Ecology. Lowlands. Margins of eutrophic marshes; marshy borders of slow streams; swamps and lakeside forests. Open ground; soft, muddy soil covered with dense vegetation (e.g., Carex, Juncus, Eleocharis, moss). Biology. Seasonality: May-June. Tenerals: September (AB). Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous, probably capable of flight. Fast runner. Collecting technique. Treading the vegetation into the water. References. Lindroth, 1961 and 1963a (natural history). Pentagonica bicolor (LeConte, 1863) Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Pentagonica felix Bell, 1987 Ecology. Mountains. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Pentagonica flavipes flavipes (LeConte, 1853) Ecology. Lowlands. Damp places, marshy areas, and lake shores. Somewhat open ground; moist or wet soil. Mostly diurnal; active in the sunshine on plants and dead epiphytes growing on trees; usually sheltering on cloudy or cool days on the ground among dead leaves. Biology. Seasonality: January-August, November-December. Adult found over-

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wintering in weed debris. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Fast runner. Frequent climber (on plants and trees). Collecting techniques. Sweeping or beating the vegetation; light trapping; raking the leaf litter; sifting wood debris. Reference. Kirk, 1970 (dispersal power). Pentagonica marshalli Mateu, 1994 Ecology. Lowlands. Biology. Seasonality: January, March. Dispersal power. Macropterous, probably capable of flight. Reference. Mateu, 1994 (taxonomy). Pentagonica nigricornis Darlington, 1934 Ecology. Lowlands. Biology. Seasonality: June. Dispersal power. Macropterous, probably capable of flight. Pentagonica picticornis Bates, 1883 Ecology. Lowlands and mountains. Open forests and shrublands. Open or slightly shaded ground; dry mossy soil. Mostly diurnal; active in the sunshine on flowering trees, shrubs (e.g., Prunus), and plants; sheltering on cloudy or cool days under moss, lichens, among dead leaves, and other debris. Gregarious (in winter). Biology. Seasonality: February, May-October. Adults found overwintering in woods; among dead leaves. Dispersal power. Macropterous. Frequent flier (in the sunshine; to artificial lights at night). Fast runner. Frequent climber (on trees, shrubs, and plants). Collecting techniques. Beating or sweeping the vegetation; light trapping; turning moss carpets. Reference. Kiteley, 1977 (ecology). Pericompsus (Pericompsus) ephippiatus (Say, 1830) Ecology. Lowlands. Banks of rivers and brooks. At a some distance from water. Open ground; moderately moist soil consisting of very fine sand, often with silty or muddy coating, bare or covered with sparse vegetation (e.g., Carex, Equisetum). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in burrows dug in the ground or in soil crevices. Gregarious. Biology. Seasonality: February-October. Tenerals: July-August (LA, MD, TX). Adults found overwintering in leaf litter. Defense-mechanism: body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (in the sunshine; to artificial lights at night). Fast runner. Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping; sifting leaf litter in early spring. References. Ulke, 1903 (ecology); Blatchley, 1910 (ecology); Kirk, 1969 and 1975c (dispersal power); Erwin, 1974 and 1981 (natural history). Pericompsus (Pericompsus) laetulus LeConte, 1852 Ecology. Mountains. Biology. Seasonality: January, June-September. Tenerals: August-September (AZ, NM). Dispersal power. Macropterous. Frequent flier (to artifi-

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cial lights at night). Collecting technique. Light trapping. Reference. Erwin, 1974 (natural history). Pericompsus sellatus LeConte, 1852 Ecology. River banks. Sandy soil. Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: January, March, July-August. Dispersal power. Wing-condition unknown. Fast runner. Collecting technique. Raking the leaf litter. Perigona nigriceps (Dejean, 1831) Ecology. Lowlands. Gardens. Borders of ponds, lakes, and rivers. Open ground; moist or wet soil, rich in organic debris. Crepuscular and nocturnal; sheltering during the day in compost or grass heaps, and under dead leaves. Tropical regions: occurring in decaying vegetation or in the soil near plant roots (e.g., peanut). Gregarious. Biology. Seasonality: April-November. Tenerals: late July (QU), mid-August (VT). Dispersal power. Macropterous. Frequent flier (at dusk; to artificial lights at night). Moderate runner. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Sifting or lifting compost heaps; light trapping. References. Ulke, 1903 (ecology); Jeannel, 1942 (ecology); Lindroth, 1986 (ecology, dispersal power); Koch, 1989 (ecology); Nelson, 1991 (ecology); Will et al. 1995 (dispersal power). Perigona pallipennis (LeConte, 1853) Ecology. Lowlands. Pond borders, open woodlands, sawmills, and piles of sawdust. Open ground; moist or wet soil, rich in vegetal debris. Nocturnal; sheltering during the day in leaf litter, in piles of fermenting sawdust, among decaying wood chips, and under other debris. Gregarious. Biology. Seasonality: March-September, November. Adults found overwintering in woods; under the loose bark of dead trees (e.g., Carya) and at their base. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; sifting (e.g., with Berlese funnels) piles of sawdust or decaying wood chips. References. Ulke, 1903 (ecology); Nelson, 1991 (ecology). Philorhizus atriceps (LeConte, 1880) Ecology. Lowlands. Wetlands. Open ground; moist soil covered with clumps of broomsedges (Andropogon). Nocturnal; sheltering during the day among grass stems. Biology. Seasonality: February-August, December. Adults found overwintering in the stems and roots of broomsedges (Andropogon) and among debris. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Frequent climber (on plants). Collecting techniques. Inspecting and sweeping broomsedges; sifting debris surrounding the roots of broomsedges; light trapping. References. Schwarz, 1884 (ecology); Blatchley, 1928 (ecology, biology); Kirk, 1969 (ecology, biology).

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Phloeoxena (Oenaphelox) signata (Dejean, 1825) Ecology. Lowlands. Deciduous forests (mostly; e.g., Quercus), coniferous forests (e.g., Pinus), and shrublands (e.g., Castanea). Also tropical gallery forests and scrub forests, with evergreens and palms (Mexico). Shaded ground. Nocturnal and diurnal; sheltering during the day mostly under the loose bark of trees (e.g., Quercus); sometimes active on logs where fungi grow or foraging on Castanea-blooms. Biology. Seasonality: January-June, August-December. Adults found overwintering under the loose bark of trees (eg., Quercus, Pinus). Dispersal power. Macropterous. Frequent climber (on shrubs and trees). Collecting techniques Lifting the loose bark of trees; sweeping or beating Castanea-blooms. References. Leng, 1915 (ecology, biology); Löding, 1945 (ecology, biology); Kirk, 1969 (ecology); Ball, 1975 (ecology); Shpeley and Ball, 2000 (ecology); Ball and Bousquet, 2001 (taxonomy). Phrypeus rickseckeri (Hayward, 1897) Ecology. Banks of running waters, mostly large rivers; sometimes on lake shores. At considerable distance from water. Open ground; rather dry, sandy-gravelly soil. Nocturnal; sheltering during the day among gravel. Biology. Seasonality: May-July. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Strong burrower. Collecting techniques. Pouring water over the ground; raking the loose gravel. Reference. Lindroth, 1963b (natural history). Physea hirta LeConte, 1853 Ecology. Lowlands. Forests (Mexico). In and around ant nests. Biology. Seasonality: August. Dispersal power. Unknown. Collecting technique. Inspecting ant nests. Reference. Ball and McCleve, 1990 (natural history). Piesmus submarginatus (Say, 1823) Ecology. Forests (e.g., Pinus, Quercus, Liquidambar). Shaded ground; moderately moist soil. Nocturnal; sheltering (adult and larva) during the day under the loose bark of logs and fallen trees (Pinus). Biology. Seasonality: January-May, July-December. Adults found overwintering in tree-stumps and under loose bark. Dispersal power. Macropterous. Moderate runner. Frequent climber (on logs, fallen trees, and tree-stumps). Collecting techniques. Lifting the loose bark of logs and fallen trees; beating trees; pitfall trapping; collecting at night with a headlamp. References. Leng, 1915 (ecology); Löding, 1945 (ecology); Davidson, 1995 (ecology); Purrington and Stanton, 1996 (ecology). Piosoma setosum LeConte, 1848 Ecology. Uplands. Grasslands, prairies, rangelands (short- and mid-grass), pastures, and hills. Open ground; sandy (mostly) or clayish soil covered with some vegetation (e.g., grass). Mostly nocturnal; sometimes active in the sunshine on plants; usually shel-

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tering during the day in burrows dug in the soil under pieces of wood, dry cattle dung, and stones. Biology. Seasonality: May-August. Tenerals: early June (BC). Adult food, in the field: plant seeds. Dispersal power. Wing-dimorphic: macropterous or brachypterous (incapable of flight). Frequent flier (to artificial lights at night). Occasional climber (on plants). Strong burrower. Collecting techniques. Pitfall trapping; turning debris; sweeping plants. References. Lindroth, 1968 (natural history); Kirk, 1975c (dispersal power); Lavigne, 1977 (ecology). Platidiolus vandykei Kurnakov, 1960 Ecology. Mountains. Margins of cold, swift rivers. Close to water. Open ground; moist, gravelly-stony, bare soil. Nocturnal; sheltering during the day among gravel and large stones. Biology. Seasonality: May-September. Tenerals: June (AK), September (AB). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Strong burrower. Collecting technique. Raking the soil. Reference. Lindroth, 1961 (natural history). Platynus (Microplatynus) agilis LeConte, 1863 Ecology. Mountains. Biology. Seasonality: April-June, August-September. Dispersal power. Brachypterous, incapable of flight. Platynus angustatus Dejean, 1828, see Batenus (Platynidius) angustatus angustatus (Dejean, 1828). Reference. Morvan, 2000 (taxonomy). Platynus (Platynus) brunneomarginatus (Mannerheim, 1843) Ecology. Lowlands. Margins of lakes and streams. Shaded ground; wet stony soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: January-February, April-May, July, September, December. Tenerals: late June (ID), August (WA). Predators: robins and Scaphinotus punctatus. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Turning stones. References. Hatch, 1953 (ecology); Lindroth, 1966 (natural history). Platynus cazieri Liebherr and Will, 1996 Ecology. Mountains. Biology. Seasonality: July, September. Dispersal power. Macropterous, probably capable of flight. Reference. Liebherr and Will, 1996 (taxonomy). Platynus cincticollis (Say, 1823), see Batenus (Batenus) cincticollis (Say, 1823) Reference. Morvan, 2000 (taxonomy).

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Platynus cohni Liebherr and Will, 1996 Ecology. Mountains. Deciduous forests (e.g., Quercus). Nocturnal; sheltering during the day under loose bark. Biology. Seasonality: June-August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Lifting loose bark. Reference. Liebherr and Will, 1996 (taxonomy). Platynus (Platynus) decentis (Say, 1823) Ecology. Lowlands, mountains, and subalpine zone. Low forests (deciduous, mixed, coniferous) bordering rivers, brooks, lakes, and ponds; swamp-forests, swamps, borders of eutrophic marshes, and subalpine forest clearings. Usually close to water. Shaded ground; moist soil, rich in organic debris. Mostly nocturnal; sometimes active during the day on shrubs (e.g., Cornus, Salix, Corylus); usually sheltering under the loose bark of trees (deciduous, coniferous) or under dead leaves, pieces of wood, logs, and stones. Adults show an early season preference for running on the ground, while in late June and early July a preference for tree climbing (NH). Gregarious (in winter). Biology. Seasonality: January-December. Copulating pairs: late June (QU). Gravids: mid-May to late June (IL, QU); late June to mid-July (AB). Tenerals: April-December; mostly from mid-July on. Adults found overwintering in woods and at the edge of woods; under well-embedded stones (as deep as 15 cm) and in leaf litter (mostly), also in rotten logs, branches, and tree-stumps. Adult food: lepidopterous immature stages (in the field); diplopods, mealworms, shrimps, and a piece of earthworm (in captivity). Larval food, in captivity: mealworms. Economic importance: may play an important role in the control of the Spruce Budworm (Choristoneura fumiferana (Clemens)). Predators: frogs and toads. Parasites: nematodes, mites, and Laboulbeniales (Fungi). Dispersal power. Submacropterous, probably incapable of flight. Moderate runner. Strong burrower. Frequent climber (on trees, tree-stumps, shrubs, and logs). Favored by spring floods along rivers. Collecting techniques. Lifting the loose bark of trees; pitfall trapping; using tree bands; beating the vegetation; sugaring tree-trunks; light trapping. References. Lindroth, 1955a (as sinuatum Dejean), 1963a (as decens (Say)), and 1966 (natural history); Gilbert, 1957 (biology); Bousquet and Pilon, 1977 (biology); Levesque et al. 1979 (biology); Erwin, 1981 (natural history); Reeves et al. 1983 (ecology, biology); Epstein and Kulman, 1990 (ecology, biology); Holliday, 1991 (ecology, biology); Niemelä et al. 1992 (ecology, biology). Platynus falli (Darlington, 1936) Ecology. Mountains. Biology. Seasonality: April. Dispersal power. Brachypterous, incapable of flight. Platynus hypolithos (Say, 1823), see Batenus (Platynidius) angustatus hypolithos (Say, 1823) Reference. Morvan, 2000 (taxonomy).

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Platynus indecentis Liebherr and Will, 1996 Ecology. Lowlands and mountains. Sphagnum-bogs. Open ground; soft, wet, mossy soil. Nocturnal; sheltering during the day under the loose bark of dead trees (e.g., Pinus, Larix, Populus) and in the rotten wood of standing dead trees. Biology. Seasonality: MaySeptember. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Regular climber (on trees). Collecting techniques. Lifting the loose bark of dead trees; beating trees; collecting at night with a headlamp. Reference. Liebherr and Will, 1996 (taxonomy, ecology). Platynus (Platynus) lyratus (Chaudoir, 1879) Ecology. Lowlands and mountains. Stream banks (Mexico). Sandy soil. Nocturnal; sheltering during the day under stones. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning stones. Platynus mannerheimii (Dejean, 1828), see Batenus (Batenus) mannerheimii (Dejean, 1828 Reference. Morvan, 2000 (taxonomy). Platynus megalops (Bates, 1882) Ecology. A riparian forest. Biology. Seasonality: August-September. Dispersal power. Macropterous. Frequent flier (often swarmings to artificial lights at night). Collecting technique. Light trapping. Platynus (Platynus) opaculus LeConte, 1863 Ecology. Lowlands. Flood-plain forests (e.g., Acer saccharinum, Quercus bicolor): along large rivers and lakes. Shaded ground; inundated (mostly) or wet, muddy or clay soil, rich in organic debris (e.g., dead leaves, branches, logs, fallen trees). Nocturnal; sheltering during the day mostly under the loose bark of tree-stumps, trees, and logs protruding above the water surface, also in logs lying on or near the shore. Gregarious (in winter). Biology. Seasonality: January-November. Tenerals: August (NY). Adults found overwintering on higher and drier ground, at a good distance (15-100 m) from water, on wooded hills and at the edge of woods; under stones. Adult and larval food, in captivity: mealworms. Phoretic mites found on adult body. Defense-mechanism: when alarmed, the adult escapes by swimming and diving into water (as deep as 25 cm). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in seashore drift material and on mountain tops, which indicates flight ability. Frequent climber (on logs, tree-stumps, and trees). Good swimmer; able to remain submerged for 4 minutes. Collecting techniques. Lifting the loose bark of tree-stumps, trees, and logs protruding above the water surface; night collecting with a headlamp. Reference. Davidson and Bell, 1977 (ecology).

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Platynus (Glyptolenopsis) ovatulus (Bates, 1884) Ecology. Mountains. Forests (e.g., Quercus, Pinus). Shaded ground; moderately moist soil. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: April-August, November. Dispersal power. Submacropterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. Platynus (Platynus) ovipennis (Mannerheim, 1843) Ecology. Lowlands and mountains. Coniferous (e.g., Pseudotsuga) and deciduous forests (e.g., Betula). Shaded ground; moist soil. Nocturnal; sheltering during the day under stones, pieces of wood, and in rotten wood. Biology. Seasonality: March-May, July-December. Tenerals: late July (OR). Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones or pieces of wood. Reference. Gray and Hatch, 1941 (ecology). Platynus (Platynus) parmarginatus Hamilton, 1893 Ecology. Lowlands and mountains. Deciduous (mostly; e.g., Acer, Quercus) and coniferous forests: along brooks and ravines. Shaded ground; wet soil. Mostly nocturnal; usually sheltering during the day under the loose bark of trees and under stones; sometimes active on cloudy or cool days on trees. Biology. Seasonality: February-September, December. Tenerals: late July (KY), August (MD). Adults found overwintering under the loose bark of trees. Adult food, in the field: lepidopterous immature stages. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Frequent climber (on trees). Collecting techniques. Pitfall trapping; lifting the loose bark of trees; beating trees; sugaring trees; using tree bands; using malaise traps; pyrethrum spraying; collecting at night with a headlamp; turning stones. References. Kirk, 1970 (dispersal power); Erwin, 1981 (natural history); Krinsky and Oliver, 1988 (ecology); Krinsky, 1989 (ecology). Platynus (Microplatynus) pecki Barr, 1982 Ecology. Mountains. A coniferous forest (Picea): on cool, moist soil, under boulders (NM). Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Platynus prognathus Van Dyke, 1926 Ecology. Lowlands. Biology. Seasonality: April-May, July. Dispersal power. Brachypterous, incapable of flight. Reference. Liebherr, 1989 (dispersal power). Comments. The supraspecific status of this species was not dealt with by Morvan (2000).

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Platynus rufiventris (Van Dyke, 1926) Ecology. Mountains. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Platynus (Platynus) tenuicollis (LeConte, 1848) Ecology. Deciduous and mixed forests: along turbulent brooks (mostly) and rivers. Caves (frequently). Shaded ground; wet, gravelly or gravelly-stony, bare soil. Nocturnal; sheltering during the day under stones (mostly) and dead leaves. Biology. Seasonality: March-December. Tenerals: mid-June to September. Adults found overwintering in woods; under stones. Predators: toads. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on trees). Collecting techniques. Turning stones; light trapping. References. Blatchley, 1910 (ecology; as reflexus LeConte); Lindroth, 1966 (natural history); Larochelle, 1974a (ecology); Kirk, 1975c (ecology, dispersal power); Erwin, 1981 (natural history); Liebherr, 1986a (dispersal power) and 1988b (ecology, dispersal power). Platynus (Platynus) trifoveolatus Beutenmüller, 1903 Ecology. Mountains. Coniferous forests (e.g., Picea, Abies): along streams, springs, seepages, and ravines. Shaded ground; often steep, moist or wet soil. Nocturnal; sheltering during the day under stones (mostly), the loose bark of logs and dead trees, in leaf litter, in and under moss. Biology. Seasonality: April-September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning stones; lifting the loose bark of logs and trees. Reference. Barr, 1969 (ecology). Platypatrobus lacustris Darlington, 1938 Ecology. Lowlands and mountains. Beaver houses: along rivers, brooks, lakes, and ponds. Open ground; moist or wet, muddy soil, bare or almost so. Nocturnal; sheltering during the day in the outer layers of beaver house walls consisting of mud, sticks, and branches. Gregarious. Associated with Pterostichus castor Goulet and Bousquet and Atranus pubescens Dejean. Biology. Seasonality: June-October. Copulating pairs: July (QU), August-October (AB). Oviposition: August-October (AB, QU). Tenerals: mid-May to early July. Adult food, in captivity: mealworms and dipterous larvae. Larval food, in captivity: mealworms. Phoretic mites found on adult body. Defense-mechanism: when alarmed, the adult escapes by burrowing into the substrate or diving into the water. Dispersal power. Macropterous. Frequent flier (to artificial lights at night; in captivity). Moderate runner. Strong burrower. Good swimmer. Favored by laws protecting the beaver habitat. Collecting techniques. Inspecting the walls of beaver houses; light trapping. References. Goulet, 1965b (ecology, biology) and 1977b (natural history); Landry, 1980b (biology); Dearborne and Donahue, 1993 (dispersal power); Bousquet and Grebennikov, 1999 (biology). Comments. The beetle was considered a rarity until its habitat was discovered.

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Plochionus (Menidius) amandus amandus Newman, 1840 Ecology. Lowlands. Marshes and swamps. Open or slightly shaded ground; moist soil. Sheltering during the day among the dead leaves of Cabbage palmettos and in Spanish moss bunches. Biology. Seasonality: February. Dispersal power. Macropterous, probably capable of flight. Frequent climber (on trees). Collecting techniques. Raking dead palmetto leaves; beating trees (e.g., with Spanish moss). Plochionus (Menidius) amandus discoideus LeConte, 1880 Ecology. Lowlands. Marshes. Open or slightly shaded ground; moist soil. Sheltering during the day among the dead leaves of Cabbage palmettos and on herbage. Biology. Seasonality: February, April-June. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Frequent climber (on trees and plants). Collecting techniques. Raking palmetto dead leaves; sweeping plants. Reference. Leng, 1915 (ecology). Plochionus (Menidius) amandus vittatus LeConte, 1844 Ecology. Lowlands. In leafroller nests on the American olive (AL). Biology. Seasonality: February, December. Adult food, in the field: lepidopterous caterpillars. Adults found overwintering in leafroller nests. Dispersal power. Macropterous, probably capable of flight. Frequent climber (on trees). Collecting technique. Inspecting leafroller nests on trees. References. Leng, 1915 (ecology); Löding, 1945 (biology). Plochionus (Menidius) bicolor Notman, 1919 Ecology. Lowlands. Biology and Dispersal power. Unknown. Plochionus (Plochionus) pallens (Fabricius, 1775) Ecology. Coastal lowlands. Parks and their vicinity (Europe). Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Plochionus (Menidius) timidus Haldeman, 1843 Ecology. Lowlands. Deciduous forests (e.g., Quercus, Acer, Carya, Morus, Salix), cypressswamps, and rows of deciduous trees. Shaded ground; moderately moist soil. Nocturnal and diurnal; sheltering during the day under the loose bark of standing and fallen trees, and on trees ( e.g., Quercus, Acer, Carya), in the nests of the Fall Webworm (Hyphantria cunea Drury). Larval habitat: same as adult; also found on persimmon trees (Diospyros). Biology. Seasonality: January-December. Gravids: May-August (MO). Eggs are laid on tree leaves. Tenerals: June-September. Adults found overwintering under the loose bark or bark scales of trees (Quercus, Taxodium, Salix). Adult and larval food, in the field: lepidopterous caterpillars (Fall Webworm). Adult food, in captivity: aphids and other soft-bodied insects. Larval food, in captivity: lepidopterous caterpillars. Eco-

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nomic importance: may play an important role in the control of webworms. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Frequent climber (adult and larva; on trees). Collecting techniques. Lifting the loose bark of trees; using tanglefoot screens; sticky boards; beating trees; sugaring trees; light trapping; collecting at night with a headlamp; inspecting the nests of the Fall Webworm. References. Duffey, 1891 (natural history); Murtfeldt, 1892 (ecology, biology); Ulke, 1903 (ecology); Blatchley, 1910 (ecology); Emden, 1942 (biology); Löding, 1945 (ecology, biology); Erwin, 1981 (natural history); Braun et al. 1990 (biology); Zhou and Goyer, 1993 (ecology, biology). Poecilus (Poecilus) chalcites (Say, 1823) Ecology. Lowlands and mountains. Cultivate fields (corn (mostly), soybean, peanut, cotton, oat, hay, barley, wheat, tobacco, cabbage, asparagus, rye, alfalfa), rangelands, pastures, meadows, lawns, vacant lots, gravel pits, roadsides, orchards (e.g., apple), open forests, and upper banks of bodies of water. Open ground; moderately moist soil consisting often of clay loam, covered with moderate or dense vegetation. Mostly nocturnal; usually sheltering during the day in cracks in the soil, under clods of earth, dead leaves and dead fallen plants, pieces of wood, logs, and stones. Biology. Seasonality: January-December. Copulating pairs: May (QU); coitus lasting 6-17 minutes (13 minutes on average). Oviposition: late May to mid-August, mostly mid-June (SD). Eggs are laid in masses of 2-5 in hard cells in cracks in the soil or under clods of earth. Tenerals: mostly early August (SD). Adults found overwintering in cultivated fields, vacant lots, meadows, on hills, and at the edge of woods; into the soil (as deep as 15 cm) and under well-embedded stones. Able to survive winter inundations. Adult food: lepidopterous caterpillars (frequently), flies, hymenopterans, other insects, fungi and other vegetal tissue (in the field); lepidopterous caterpillars, chrysomelid eggs, mealworms, earthworms, slugs, meat, corn kernels, and raisin (in captivity). Larval food, in captivity: mealworms. Adults and larvae are cannibalistic in captivity. Economic importance: may play an important role in the control of lepidopterous caterpillars. Predators: catbirds, crows, kingbirds, starlings, thrushes, and woodpeckers. Parasites: ascid mites, halolaelaphid mites, tapeworms, and Laboulbeniales (Fungi). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Frequent in lakeshore drift material, which indicates flight ability. Moderate runner. Strong burrower. Strongly favored by human activities. Collecting techniques. Pitfall trapping; turning debris; using tanglefoot screens; light trapping; night collecting with a headlamp; turning drift material. References. Lindroth, 1966 (natural history); Kirk, 1975b (biology) and 1975c (ecology, dispersal power); Hsin et al. 1979 (biology); Best et al. 1981 (biology, dispersal); Erwin, 1981 (natural history); Lesiewicz et al. 1983 (ecology, biology); Los and Allen, 1983 (biology); Ferguson and McPherson, 1985 (biology); Will et al. 1995 (dispersal power); Chen and Willson, 1996 (biology). Poecilus (Poecilus) coloradensis (Csiki, 1930) Ecology. Uplands. Biology and Dispersal power. Unknown.

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Poecilus (Poecilus) corvus (LeConte, 1873) Ecology. Lowlands and uplands. Cultivated fields (e.g., wheat, barley) and tall-grass prairies. Open ground; rather dry, often clay soil, sometimes alkaline. Nocturnal; sheltering during the day. Biology. Seasonality: May-September. Adult food: dipterous larvae (in the field); grasshopper eggs (mostly), mealworms, and cat food (in captivity). Larval food, in captivity: mealworms. Dispersal power. Macropterous. Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting technique. Pitfall trapping. References. Lindroth, 1966 and 1971a (natural history); Frank, 1971b (biology). Poecilus (Poecilus) cursitor LeConte, 1852 Ecology. Uplands. Orchards (e.g., apple). Biology. Seasonality: April, June-August. Adult food, in captivity: lepidopterous caterpillars. Economic importance: the adult may play an important role in the control of the Codling Moth. Dispersal power. Macropterous, probably capable of flight. References. Riddick and Mills, 1995 and 1996 (ecology, biology). Poecilus (Poecilus) cyanicolor Chaudoir, 1876 Ecology. Uplands. Biology and Dispersal power. Unknown. Poecilus (Poecilus) diplophryus Chaudoir, 1876 Ecology. Uplands. Biology and Dispersal power. Unknown. Poecilus (Poecilus) laetulus (LeConte, 1863) Ecology. Unknown. Biology. Seasonality: April. Dispersal power. Macropterous, probably capable of flight. Poecilus (Poecilus) lucublandus acomanus Casey, 1924 Ecology and Biology. Unknown. Dispersal power. Submacropterous, probably incapable of flight. Poecilus (Poecilus) lucublandus louisinus Casey, 1924 Ecology and Biology. Unknown. Dispersal power. Submacropterous, probably incapable of flight. Poecilus (Poecilus) lucublandus lucublandus (Say, 1823) Ecology. Lowlands, mountains, subalpine and alpine zones. Cultivated fields (corn (mostly), hay, alfalfa, asparagus, cabbage, soybean, hop, rye, oat, wheat, barley, flax, fescue, tobacco, potato, carrot, raspberry), tilled fields, pastures, meadows, relict prairies, grass-

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lands, hills, gardens, lawns, vacant lots, fencerows, roadsides, gravel pits, sand pits, hedges, orchards (e.g., apple), thickets, forest edges and clearings, light and open woods; upper zone of lake shores, brook and river banks; ravines. Greenhouses (occasionally). Open (mostly) or slightly shaded ground; moderately dry soil consisting generally of clay loam or clay (gravel- or sand-mixed), covered with some vegetation (e.g., grass). Mostly nocturnal; sometimes active in the sunshine on plants; usually sheltering during the day in the soil at the base of plants, under stones, logs, branches, pieces of wood, dead fallen plants, dead leaves, heaps of hay, silt cakes, and reindeer moss (Cladonia). Active at low temperatures under snow in October-November. Larval habitat: in the soil and under dry cow dung. Biology. Seasonality: January-December. Copulating pairs: April-July (QU); coitus lasting 20-121 minutes (81 minutes on average). Oviposition: April to early September, mostly May-June (NY, ON, QU, SD). Tenerals: April-May (rarely; QU), July-November (throughout the distribution range). Adults found overwintering in cultivated fields, vacant lots, pastures, on hills, in the upper zone of sand pits, at the edge of woods, and in woods; in the soil (as deep as 25 cm), under dead leaves, plant rosettes (Verbascum), crop debris, stones, logs, under the lose bark of logs and tree-stumps. Adult able to survive extreme low temperatures and winter inundations. Adult food: lepidopterous caterpillars; dipterous larvae, pupae, and adults; hymenopterous larvae, chrysomelids, scarabaeid larvae, elaterids, dead Pterostichus chalcites, other insects, fungi, and vegetal tissue (in the field); lepidopterous caterpillars; dipterous eggs, pupae and adults; weevil immatures stages, larval and adult elaterids, carabids, a dead Pterostichus chalcites, larval chrysomelids, earthworms, slugs, meat, egg yolk and white, seeds, corn kernels, and raisin (in captivity). Larval food: elaterid and scarabaeid larvae (in the field); mealworms (in captivity). Adults are cannibalistic in captivity. Economic importance: may play an important role in the control of lepidopterous caterpillars. Predators: frogs, toads, snakes, bluebirds, crows, grouses, starlings, swallows, thrushes, and spiders. Parasites: ascid mites, podapolipid mites, gordiid worms (Nematomorpha), and tachinid flies. Defense-mechanism: when alarmed, the adult escapes by running around erratically; females unwilling to mate may discharge a defensive secretion against males. Dispersal power. Submacropterous. Occasional flier (in the spring sunshine; to artificial lights at night; to artifical lights in captivity). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Strong burrower. Favored by human activities (strongly) and spring floods along large rivers. Collecting techniques. Pitfall trapping; turning stones; baiting traps with molasses and meat; turning drift material; sweeping plants. References. Schaupp, 1881a (biology); Blatchley, 1910 (biology); Rivard, 1964c (biology); Lindroth, 1966 (natural history); Barlow, 1970 (biology); Frank, 1971b (biology); Kirk, 1971b (natural history); Kirk and Dupraz, 1972 (biology); Kirk, 1975c (ecology); Larochelle, 1975l (biology); Randolph et al. 1976 (biology); Doane, 1981 (ecology, biology); Erwin, 1981 (natural history); Levesque et al. 1979 (biology); Bousquet, 1986b (biology); Levesque and Levesque, 1987 and 1994 (biology); Epstein and Kulman, 1990 (ecology, biology); Holliday, 1991 (biology); Will et al. 1995 (dispersal power). Poecilus (Poecilus) mexicanus Chaudoir, 1876 Natural History. Unknown.

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Poecilus (Derus) nearcticus (Lindroth, 1966) Ecology. Lowlands. An open sandy place, among grass, under driftwood. Biology. Seasonality: June-August. Dispersal power. Brachypterous, incapable of flight. Reference. Lindroth, 1966 (natural history). Poecilus (Poecilus) occidentalis (Dejean, 1828) Ecology. Unknown. Biology. Seasonality: June. Adult food, in captivity: grasshopper eggs. Dispersal power. Unknown. Poecilus (Poecilus) scitulus LeConte, 1848 Ecology. Lowlands and uplands. Grasslands, rangelands, cultivated fields (e.g., corn, wheat), sod fields; vicinity of sloughs, alkaline ponds, lakes, reservoirs, and streams. Open ground; moist clay soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day. Biology. Seasonality: April-September. Predators: swallows. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; light trapping. References. Lindroth, 1966 (natural history); Hsin et al. 1979 (biology); Chen and Willson, 1996 (biology). Poecilus (Poecilus) texanus (LeConte, 1863) Ecology. Unknown. Biology. Seasonality: September. Dispersal power. Unknown. Pogonistes planatus (G.H. Horn, 1876), see Diplochaetus planatus (G.H. Horn, 1876). Reference. Bousquet and Laplante, 1997 (taxonomy). Pogonodaptus mexicanus (Bates, 1878) Ecology. Lowlands. Borders of marshes; swamps; marshy borders of ponds, lakes, and slow rivers. Open ground; soft, wet soil consisting of mud mixed with dead leaves. Nocturnal; sheltering during the day in burrows dug in the soil. Defense-mechanism: when alarmed, the adult may escape by flying. Biology. Seasonality: July-September. Tenerals: late July (TX). Dispersal power. Macropterous. Occasional flier (when alarmed; to artificial lights at night). Moderate runner. Collecting techniques. Treading the soil with the feet; pouring water over the ground. Pogonus parallelus LeConte, 1874, see Diplochaetus rutilus (Chevrolat, 1863). Reference. Bousquet and Laplante, 1997 (taxonomy).

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Pogonus texanus Chaudoir, 1868 Ecology. Coastal lowlands. Borders of salt marshes; muddy salt flats. Open ground; moist, clayey or muddy, saline soil covered with sparse vegetation. Nocturnal; sheltering during the day in clay cracks (mostly) and under vegetal debris (e.g., logs). Gregarious. Biology. Seasonality: June, November. Defense-mechanism: when alarmed, the adult may escape by running. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. References. Lindroth, 1985 (natural history); Bousquet and Laplante, 1997 (taxonomy). Polpochila (Phymatocephalus) capitata (Chaudoir, 1852) Ecology. Mountains. Mexico: Borders of standing waters in deserts and semi-deserts. Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Unknown. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; light trapping. Polpochila (Phymatocephalus) erro (LeConte, 1854) Ecology. Mountains. Borders of a small semi-permanent pond, covered with dense vegetation (e.g., Juncus). Biology. Seasonality: August. Dispersal power. Macropterous, probably capable of flight. Polpochila (Polpochila) rotundicollis Bates, 1882 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Polyderis diaphana (Casey, 1918) Natural History. Unknown. Polyderis laevis (Say, 1823) Ecology. Lowlands. Pastures, meadows, hills, vacant lots, gravel pits; upper banks of lakes, marshes and brooks; woodchuck burrows and sawmills. Open ground; moderately dry or moist, often sandy soil. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in ant nests, under well-embedded stones, in the soil at the base of plants (e.g., Solidago), in piles of sawdust, among wood chips, and in leaf litter. Gregarious (in winter). Biology. Seasonality: January-November. Tenerals: September-October. (QU). Adults found overwintering in pastures, meadows, vacant lots, on hills, in gravel pits, in hawthorn hedges, at the edge of woods, in clearings, and in woods; against the undersurface of wellembedded stones, in leaf litter, moss, and clumps of grass, under wood chips and logs. Adult food: an acridid (in the field); meat (in captivity). Dispersal power. Macropterous. Frequent flier (during the day; to artificial lights at night). Moderate runner. Strong burrow-

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er. Strongly favored by human activities. Collecting techniques. Inspecting ant nests; turning embedded stones; sifting (e.g., with Berlese funnels) piles of decaying sawdust or wood chips; light trapping. References. Lindroth, 1966 (natural history); Kirk, 1969 (dispersal power) and 1970 (biology); Erwin, 1981 (natural history); Nelson, 1991 (ecology). Polyderis rufotestacea (Hayward, 1900) Ecology. Unknown. Biology. Seasonality: June-July. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Reference. Kirk, 1975c (dispersal power). Porotachys bisulcatus (Nicolai, 1822) Ecology. Lowlands. Mostly forests (coniferous), sawmills, and beaver houses. Also river banks and vacant lots. Open ground; wet soil, mostly rich in fermenting vegetal debris. Crepuscular and nocturnal; sheltering during the day mostly in wet piles of fermenting sawdust (e.g., Picea), bark, and wood chips, also under the loose bark of logs, fallen tree-stumps, and under stones; could also occur under the loose fermenting bark of fallen or standing, dead or dying coniferous trees (e.g., Picea, Pinus, Abies, Larix). Europe: also in ants nests, garbage deposits, and inundation debris. Biology. Seasonality: April-September. Tenerals: mid- to late June (ON), August (ME). Parasites: Laboulbeniales (Fungi; Europe). Dispersal power. Frequent flier (at sunset; to artificial lights at night). Moderate runner. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Sifting fermenting sawdust, pieces of bark, and wood chips; light trapping. References. Lindroth, 1945, 1966, and 1985 (natural history); Liard and Garneau, 1977 (ecology); Koch, 1989 (ecology). Promecognathus crassus LeConte, 1868 Ecology. Lowlands and mountains. Forests: often along valleys and canyons. More or less shaded ground; moist or rather dry soil, rich in humus. Nocturnal; sheltering during the day in burrows dug in the soil and under stones. Associated with millipedes. Biology. Seasonality: January-December. Adult food, in the field: millipedes. Dispersal power. Brachypterous, incapable of flight. Slow runner. Strong burrower. Collecting techniques. Pitfall trapping; turning stones. References. Van Dyke, 1925 (ecology); Lindroth, 1961 (natural history); Chen and Willson, 1996 (biology). Promecognathus laevissimus (Dejean, 1829) Ecology. Lowlands and mountains. Coniferous forests (e.g., Pseudotsuga), their edges, and ravines. More open ground and drier soil than P. crassus; soil rich in humus. Nocturnal; sheltering during the day in burrows dug in the soil and under stones. Associated with millipedes (Xystocheir francisca Chamberlin). Biology. Seasonality: January-July, OctoberDecember. Gravids: mid-March (CA). Tenerals: December-January (CA). Adult food: millipedes (in the field); millipedes, tipulid larvae, mealworms, and meat (in captivity). Dispersal power. Brachypterous, incapable of flight. Slow runner. Strong burrower. Collect-

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ing techniques. Pitfall trapping; baiting traps with meat; turning stones. References. Garnett, 1920 (ecology); Van Dyke, 1925 (ecology; as debilis Casey); MacSwain and Garner, 1956 (biology); LaBonte, 1983 (biology); Weber and Kavanaugh, 1992 (ecology). Pseudamara arenaria (LeConte, 1848) Ecology. Lowlands and mountains. Deciduous forests (e.g., Acer, Ostrya), their clearings and edges. Shaded ground; preferably sloping, moderately moist, often gravelly soil covered with a thick layer of dead leaves and humus. Nocturnal; sheltering during the day in leaf litter. Gregarious (in winter and spring). Biology. Seasonality: April-November. Tenerals: early May and October (QU). Adults found overwintering on higher and drier ground; in leaf litter (mostly), humus, under logs, stones, and in rotten tree-stumps. Adult food, in captivity: meat. Predators: toads. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (to artificial lights at night). Frequent in sea- and lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Collecting techniques. Raking the leaf litter; turning drift material; sweeping plants. References. Lindroth, 1968 (natural history); Larochelle, 1974a (ecology); Bousquet, 1987a (dispersal power). Pseudanophthalmus acherontis Barr, 1959 Ecology. Caves: along stream banks. Wet, silty soil. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus alabamae Valentine, 1932 Ecology. Caves: along stream banks. Wet, muddy soil. Occurring on wet rotting pieces of wood. Biology. Seasonality: July, October. Dispersal power. Brachypterous, incapable of flight. Fast runner. Climber (on pieces of wood). Pseudanophthalmus assimilis Barr, 1981 Ecology. Caves. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus audax (G.H. Horn, 1883) Ecology. In deep soil (mostly). Caves (occasionally). Occurring on wet rotting wood. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Occasional climber (on pieces of wood). Pseudanophthalmus avernus Valentine, 1945 Ecology. Caves. Biology. Seasonality: June-July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting techniques. Pitfall trapping; baiting traps.

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Pseudanophthalmus barberi Jeannel, 1928 Ecology. Caves. Biology. Seasonality: August, October. Adult food, in the field: tubificid worms. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus barri Krekeler, 1973 Ecology. Caves. Biology. Seasonality: January, August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus beaklei Valentine, 1937 Ecology. Caves: along stream banks. Wet soil. Sheltering in leaf debris. Biology. Seasonality: April, July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Inspecting leaf debris. Pseudanophthalmus bendermani Barr, 1959 Ecology. Caves: at the back of entrance rooms (mostly) and along stream banks. Moist or wet, gravelly soil. Sheltering among gravel. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Raking the gravel. Pseudanophthalmus brevis Valentine, 1932 Ecology. Caves: upper stalactitic rooms and along river banks. Occurring on moist rotting accumulations of wood. Biology. Seasonality: March-April. Dispersal power. Brachypterous, incapable of flight. Fast runner. Excellent climber (on cave walls and pieces of wood). Pseudanophthalmus calcareus Barr, 1981 Ecology. Caves. Moist, silty soil, rich in organic debris. Sheltering under stones. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones. Pseudanophthalmus catherinae Barr, 1959 Ecology. Caves: along stream banks. Wet soil. Sheltering among loose gravel. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Raking the gravel. Pseudanophthalmus catoryctos Krekeler, 1973 Ecology. Caves. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Pseudanophthalmus cerberus cerberus Barr, 1985 Ecology. Caves. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus cerberus completus Barr, 1985 Ecology. Caves. Biology. Seasonality: February, April. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus chthonius Krekeler, 1973 Ecology. Caves: along river banks. Moist, muddy or gravelly soil. Sheltering under stones. Biology. Seasonality: July-August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones. Pseudanophthalmus ciliaris ciliaris Valentine, 1937 Ecology. Caves: along river banks, in passageways, and stalactitic chambers. Moist soil. Occurring on mud and rotting pieces of wood. Biology. Seasonality: April-August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Excellent climber (on cave walls and pieces of wood). Pseudanophthalmus ciliaris colemanensis Barr, 1959 Ecology. Caves: at the base of rapid drips of water falling from ceilings. Wet soil. Sheltering among small stones. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus ciliaris orlindae Barr, 1959 Ecology. Caves. Wet soil. Biology. Seasonality: April, June. Adult food, in the field: small invertebrates. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus cnephosus Krekeler, 1973 Ecology. Caves. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus conditus Krekeler, 1973 Ecology. Caves. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Pseudanophthalmus cordicollis Barr, 1981 Ecology. Caves. Biology. Seasonality: November. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus cumberlandus Valentine, 1937 Ecology. Caves: along the banks of streams and pools, and on bat guano. Muddy soil. Biology. Seasonality: April, August. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus darlingtoni darlingtoni Barr, 1985 Ecology. Caves. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus darlingtoni persimilis Barr, 1985 Ecology. Caves. Biology. Seasonality: September. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus deceptivus Barr, 1981 Ecology. Caves: base of steep talus slopes. Sheltering under stones. Biology. Seasonality: July. Tenerals: September. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones. Pseudanophthalmus delicatus Valentine, 1932 Ecology. Caves: along pool and stream banks. Wet, muddy soil covered with rotting wood. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus desertus desertus Krekeler, 1973 Ecology. Caves. Biology. Seasonality: January, June-July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus desertus major Krekeler, 1973 Ecology. Caves. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Pseudanophthalmus digitus Valentine, 1932 Ecology. Caves: along stream banks. Wet soil. Sheltering among rotten pieces of wood and bark. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus egberti Barr, 1965 Ecology. Caves: along steep stream banks. Wet soil. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus elevatus Valentine, 1932 Ecology. Caves: along stream banks. Moist, rocky and muddy soil. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus elongatus Krekeler, 1973 Ecology. Caves. Biology. Seasonality: July-August. Tenerals: July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus emersoni Krekeler, 1958 Ecology. Caves. Biology. Seasonality: August-September. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus engelhardti (Barber, 1928) Ecology. Caves: in stairways and walkways. Moist soil. Sheltering among wet rotting pieces of wood. Biology. Seasonality: April-May, July-September. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus eremita (G.H. Horn, 1871) Ecology. Caves. Biology. Seasonality: July-August, October. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus exiguus exiguus Krekeler, 1973 Ecology. Caves. Biology. Seasonality: February, May-August, and December. Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Pseudanophthalmus exiguus furtivus Krekeler, 1973 Ecology. Caves. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus exoticus Krekeler, 1973 Ecology. Caves. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus farrelli Barr, 1959 Ecology. Caves. Wet soil. Sheltering under stones or rotten wood. Biology. Seasonality: June, December. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus fastigatus Barr, 1981 Ecology. Caves. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus fowlerae Barr, 1980 Ecology. Caves: along stream banks. Moist or wet soil. Sheltering under stones. Biology. Seasonality: February, April, August, December. Tenerals: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique.Turning stones. Pseudanophthalmus frigidus Barr, 1981 Ecology. Caves: along temporary pools. Sheltering among wet stalactites and under stones. Biology. Seasonality: August, October. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus fulleri Valentine, 1932 Ecology. Caves: on terraced floors. Moist or wet soil. Sheltering among pebbles. Biology. Seasonality: April, July-September. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning pebbles. Pseudanophthalmus fuscus constrictus Valentine, 1932 Ecology. Caves. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Pseudanophthalmus fuscus fuscus Valentine, 1931 Ecology. Caves: along stream banks. Wet soil. Sheltering in debris. Biology. Seasonality: February-March, August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus georgiae Barr, 1981 Ecology. Caves. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus globiceps Barr, 1985 Ecology. Caves. Biology. Seasonality: July-August, December. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus gracilis Valentine, 1931 Ecology. Caves: along stream banks. Wet, muddy soil covered with rotting wood. Biology. Seasonality: January, March, August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus grandis Valentine, 1931 Ecology. Caves: along stream banks. Wet soil. Biology. Seasonality: February-March, August. Parasites: Laboulbeniales (Fungi). Defense-mechanism: the adult may emit a repugnant smell from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus hadenoecus Barr, 1965 Ecology. Caves: along stream banks. Wet, silty soil. Sheltering under small stones. Biology. Seasonality: April, August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones. Pseudanophthalmus henroti Jeannel, 1949 Ecology. Caves. Moist soil. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus hesperus Barr, 1959 Ecology. Caves: along stream banks. Moist soil. Sheltering under logs and stones. Biology. Seasonality: February, June. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning logs and stones.

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Pseudanophthalmus higginbothami Valentine, 1931 Ecology. Caves: along stream banks and in higher chambers. Moist or wet, muddy soil. Biology. Seasonality: February-March, August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Excellent climber (on cave walls). Pseudanophthalmus hirsutus Valentine, 1931 Ecology. Caves: along pool margins. Occurring on wet rotting pieces of wood. Biology. Seasonality: July-August. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. Climber (on pieces of wood). Pseudanophthalmus hoffmani Barr, 1965 Ecology. Caves. Biology. Seasonality: January, July-August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus holsingeri Barr, 1965 Ecology. Caves. Moist, silty soil. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus horni abditus Krekeler, 1973 Ecology. Caves. Biology. Seasonality: April, July-September. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus horni caecus Krekeler, 1973 Ecology. Caves. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus horni horni (Garman, 1892) Ecology. Caves. Moist, gravelly soil. Occurring among wood debris and on gravel. Biology. Seasonality: July-November. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus hortulanus Barr, 1965 Ecology. Caves. Sheltering in rotting plant debris. Biology. Seasonality: September. Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Pseudanophthalmus hubbardi (Barber, 1928) Ecology. Caves. Sheltering among stones in wet, muddy wall pockets. Biology. Seasonality: January, May, August, October-December. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Baiting traps. Pseudanophthalmus hubrichti Valentine, 1948 Ecology. Caves. Sheltering among loose concretions in limestone pockets and loose cobbles above standing water. Biology. Seasonality: April, July, September. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus humeralis Valentine, 1931 Ecology. Caves: along stream banks. Wet soil. Occurring on rotting wood or sheltering between stones. Biology. Seasonality: March, June, August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Climber (on pieces of wood). Pseudanophthalmus hypertrichosis Valentine, 1932 Ecology. Caves: along stream banks. Wet, muddy soil. Sheltering among stones. Biology. Seasonality: unknown. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus hypolithos Barr, 1981 Ecology. Caves. Sheltering under stones. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus illinoiensis Barr and Peck, 1966 Ecology. Caves: along stream banks. Wet, gravelly or muddy soil. Sheltering among gravel or occurring on mud. Biology. Seasonality: June-July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus inexpectatus inexpectatus Barr, 1959 Ecology. Caves. Wet soil. Sheltering under wet decaying wood. Biology. Seasonality: January-April, June. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus inexpectatus orientalis Krekeler, 1973 Ecology. Caves. Biology. Seasonality: June, August. Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Pseudanophthalmus inquisitor Barr, 1980 Ecology. Caves: along stream banks. Biology. Seasonality: April, August, December. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus insularis Barr, 1959 Ecology. Caves: along stream banks. Moist or wet, gravelly soil. Sheltering under rotten pieces of wood, or occurring on stones. Biology. Seasonality: August-December. Dispersal power. Brachypterous, incapable of flight. Fast runner. Climber (on stones). Pseudanophthalmus intermedius (Valentine, 1931) Ecology. Caves: along stream banks and stalactic passages. Wet soil. Standing on damp decaying pieces of wood, on the ground, or sheltering in crevices. Biology. Seasonality: March, November. Dispersal power. Brachypterous, incapable of flight. Fast runner. Excellent climber (on cave walls and pieces of wood). Pseudanophthalmus intersectus Barr, 1965 Ecology. Caves. Wet soil. Sheltering under stones. Biology. Seasonality: August, October. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones. Pseudanophthalmus jonesi Valentine, 1945 Ecology. Caves. Moist soil. Occurring on the ground or climbing on walls. Biology. Seasonality: June, October. Dispersal power. Brachypterous, incapable of flight. Fast runner. Occasional climber (on cave walls). Pseudanophthalmus krameri Krekeler, 1973 Ecology. Caves. Biology. Seasonality: March, August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus krekeleri Barr, 1965 Ecology. Caves. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus lallemanti Jeannel, 1949 Ecology. Caves. Biology. Seasonality: August. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Pseudanophthalmus leonae Barr, 1960 Ecology. Caves: along stream banks. Wet soil. Sheltering among small stones and rootlets. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus limicola Jeannel, 1931 Ecology. Caves. Biology. Seasonality: February-May, August, October. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Pitfall trapping. Pseudanophthalmus loedingi aladdini Valentine, 1945 Ecology. Caves. Biology. Seasonality: June-July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus loedingi aquaticus Valentine, 1945 Ecology. Caves. Biology. Seasonality: March, August-September, December. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus loedingi distinguens Valentine, 1948 Ecology. Caves. Biology. Seasonality: September. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus loedingi fluviatilis Valentine, 1948 Ecology. Caves. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus loedingi loedingi Valentine, 1931 Ecology. Caves: along stream banks. Wet soil. Sheltering among wet rotting pieces of wood. Biology. Seasonality: January, March, August-September. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus loedingi meridionalis Valentine, 1945 Ecology. Caves. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus loedingi profundus Valentine, 1945 Ecology. Caves. Biology. Seasonality: November. Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Pseudanophthalmus loganensis Barr, 1959 Ecology. Caves: along stream banks. Wet, cobbled or gravelly soil. Sheltering in mud cracks, among cobbles and gravel, or standing on rotten pieces of wood. Biology. Seasonality: January, April, June-July. Adult food, in the field and in captivity: small invertebrates. Dispersal power. Brachypterous, incapable of flight. Fast runner. Climber (on pieces of wood). Pseudanophthalmus longiceps Barr, 1981 Ecology. Caves. Sheltering under stones and rotting wood. Biology. Seasonality: March, September, November. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones and rotting wood. Pseudanophthalmus macradei Valentine, 1948 Ecology. Caves. Wet, silty soil. Sheltering under stones or climbing on walls. Biology. Seasonality: January, August. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. Occasional climber (on cave walls). Collecting technique. Turning stones. Pseudanophthalmus menetriesii campestris Barr, 1985 Ecology. Caves. Biology. Seasonality: March. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus menetriesii menetriesii (Motschulsky, 1862) Ecology. Caves. Moderately moist soil. Sheltering among dead leaves, sticks, and piles of rotting wood. Biology. Seasonality: January-December. Gravids: same period. Tenerals: mid-summer to fall. Adult food, in the field: springtails, mites, and worms. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. Climber (on pieces of wood). Collecting technique. Turning vegetal debris. References. McKinney, 1974 (ecology) and 1975 (biology); Kane and Ryan, 1983 (biology). Pseudanophthalmus montanus Barr, 1965 Ecology. Caves. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus nelsoni Barr, 1965 Ecology. Caves. Biology. Seasonality: April, June-July. Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Pseudanophthalmus nickajackensis Barr, 1981 Ecology. Caves. Biology. Seasonality: July, November. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus nortoni Barr, 1981 Ecology. Caves. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus occidentalis Barr, 1959 Ecology. Caves. Wet, silty soil. Sheltering under stones. Biology. Seasonality: JanuaryFebruary, June. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones. Pseudanophthalmus ohioensis Krekeler, 1973 Ecology. Caves. Biology. Seasonality: March, June. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus orthosulcatus Valentine, 1932 Ecology. Caves: along stream banks. Moist, muddy soil. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus packardi Barr, 1959 Ecology. Caves: along stream banks. Moist or wet soil. Sheltering in organic debris. Biology. Seasonality: May. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus pallidus Barr, 1981 Ecology. Caves: along stream banks. Wet, silty soil. Sheltering under stones. Biology. Seasonality: March, July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones. Pseudanophthalmus paradoxus Barr, 1981 Ecology. Caves. Sheltering under stones and rotten wood. Biology. Seasonality: MayJuly. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones and rotten wood.

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Pseudanophthalmus parvicollis Jeannel, 1931 Ecology. Caves. Biology. Seasonality: September-October. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus parvus Krekeler, 1973 Ecology. Caves. Biology. Seasonality: August-October. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus paulus Barr, 1981 Ecology. Caves: along stream banks. Wet, muddy soil. Biology. Seasonality: January. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus paynei Barr, 1981 Ecology. Caves. Wet soil covered with rotting wood. Sheltering under logs and stones. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning logs and stones. Pseudanophthalmus petrunkevitchi Valentine, 1945 Ecology. Caves: along stream banks. Wet soil. Sheltering under stones and pieces of wood. Biology. Seasonality: May-July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones and pieces of wood. Pseudanophthalmus pholeter Krekeler, 1973 Ecology. Caves. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus pilosus Barr, 1985 Ecology. Caves. Biology. Seasonality: December. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus pontis Barr, 1965 Ecology. Caves: along stream banks. Wet, silty soil. Biology. Seasonality: January, August. Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Pseudanophthalmus potomacus potomacus Valentine, 1932 Ecology. Caves: along stream banks. Wet, muddy soil. Occurring on wet, rotten wood. Biology. Seasonality: May, July-August. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus potomacus senecae Valentine, 1932 Ecology. Caves: along pool margins. Wet soil. Sheltering among pebbles. Biology. Seasonality: May-July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus praetermissus Barr, 1981 Ecology. Caves. Biology. Seasonality: July, October. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus princeps Barr, 1979 Ecology. Caves: along stream banks. Wet soil. Biology. Seasonality: July. Adult food, in captivity: dead cave crickets and raw beef. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus productus Barr, 1980 Ecology. Caves: sometimes along stream banks. Moist or wet soil. Biology. Seasonality: March, October. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus pubescens intrepidus Barr, 1985 Ecology. Caves: along stream banks. Wet soil. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus pubescens pubescens (G.H. Horn, 1868) Ecology. Caves: along stream banks. Wet silty soil. Sheltering in organic debris. Biology. Seasonality: August-November. Tenerals: mostly spring. Adult food, in the field: springtails, mites, and tubificid worms. Dispersal power. Brachypterous, incapable of flight. Fast runner. References. McKinney, 1974 (ecology) and 1975 (biology). Pseudanophthalmus punctatus Valentine, 1931 Ecology. Caves: along stream banks. Wet, muddy soil. Occurring on wet rotting wood or sheltering in crevices. Biology. Seasonality: July-August. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. Climber (on pieces of wood).

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Pseudanophthalmus pusillus Barr, 1981 Ecology. Caves. Biology. Seasonality: January, November. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus pusio (G.H. Horn, 1868) Ecology. Caves. Moist or wet, muddy soil. Sheltering under stones and straw, or occurring on wet decaying pieces of wood. Biology. Seasonality: July-August. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. Climber (on pieces of wood) Pseudanophthalmus puteanus Krekeler, 1973 Ecology. Caves. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus quadratus Barr, 1965 Ecology. Caves. Biology. Seasonality: August-September. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus rittmani Krekeler, 1973 Ecology. Caves: at some distance from water. Moist or wet, muddy soil. Sheltering under stones or occurring on the ground. Biology. Seasonality: June-August. Adult food, in the field: cave-cricket eggs. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus robustus Valentine, 1931 Ecology. Caves: along stream banks. Wet soil. Sheltering under stones, pieces of wood, and in rotting wood. Biology. Seasonality: January, March, May, July, December. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. Climber (on cave walls). Collecting technique. Turning stones and pieces of wood. Pseudanophthalmus rogersae Barr, 1981 Ecology. Caves. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus rotundatus Valentine, 1932 Ecology. Caves: along stream banks. Wet soil. Sheltering under stones, sticks, and dead leaves. Biology. Seasonality: March-May, July-August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones and vegetal debris.

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Pseudanophthalmus sanctipauli Barr, 1981 Ecology. Caves. Biology. Seasonality: April. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus scholasticus Barr, 1981 Ecology. Caves: along stream banks. Wet soil. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus scutilus Barr, 1981 Ecology. Caves: along stream banks. Wet soil. Sheltering in wet, rotten wood. Biology. Seasonality: August-September, November. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus seclusus Barr, 1981 Ecology. Caves: along stream banks. Wet, muddy soil. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus sequoyah Barr, 1981 Ecology. Caves. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus sericus Barr, 1981 Ecology. Caves. Biology. Seasonality: October. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus shilohensis boonensis Krekeler, 1958 Ecology. Caves. Biology. Seasonality: July, November. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus shilohensis mayfieldensis Krekeler, 1958 Ecology. Caves: along stream banks. Wet soil. Sheltering among cobbles. Biology. Seasonality: November. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus shilohensis shilohensis Krekeler, 1958 Ecology. Caves. Biology. Seasonality: January-June, December. Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Pseudanophthalmus sidus Barr, 1965 Ecology. Caves. Occurring on wet, rotting wood. Biology. Seasonality: July, September. Dispersal power. Brachypterous, incapable of flight. Fast runner. Climber (on pieces of wood) Pseudanophthalmus simplex Barr, 1981 Ecology. Caves: along stream banks. Wet, silty soil. Sheltering under stones or occurring on the ground in the open. Biology. Seasonality: May-June, October. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones. Pseudanophthalmus simulans Barr, 1985 Ecology. Caves. Biology. Seasonality: November-December. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus solivagus Krekeler, 1973 Ecology. Caves. Biology. Seasonality: July, October-November. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus steevesi Barr, 1981 Ecology. Caves. Biology. Seasonality: March-April, November. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus striatus (Motschulsky, 1862) Ecology. Caves: along stream banks (mostly) and in dome pits. Wet, silty soil. Biology. Seasonality: August. Adult food, in the field: tubificid worms. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus sylvaticus Barr, 1967 Ecology. Mountains. Coniferous forests (e.g., Picea): along stream banks. Wet, sandygravelly soil. Nocturnal; sheltering under stones. Biology. Seasonality: April. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones. Reference. Barr, 1969 (ecology). Pseudanophthalmus templetoni Valentine, 1948 Ecology. Caves. Dry or moist soil. Biology. Seasonality: April, June, August. Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Pseudanophthalmus tenebrosus Krekeler, 1973 Ecology. Caves. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus tenesensis Valentine, 1937 Ecology. Caves: along stream banks. Moist or wet, silty soil. Occurring on wet rotting pieces of wood or sheltering under stones. Biology. Seasonality: March, May, July-August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Climber (on pieces of wood). Pseudanophthalmus tenuis blatchleyi Barr, 1960 Ecology. Caves. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus tenuis jeanneli Krekeler, 1958 Ecology. Caves. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus tenuis morrisoni Jeannel, 1931 Ecology. Caves. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus tenuis stricticollis Jeannel, 1931 Ecology. Caves. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus tenuis tenuis (G.H. Horn, 1871) Ecology. Caves. Moist soil composed of mud sand or stones. Occurring on the ground, walls, and ceilings, or sheltering under stones. Biology. Seasonality: April-December. Tenerals: July-September. Adult food, in the field: springtails, worms (oligochaetes), diplurans, and diplopods. Predators: salamanders and spiders. Parasites: Laboulbeniales (Fungi). Defensemechanism: when alarmed, the adult escapes into cracks or crevices. Dispersal power. Brachypterous, incapable of flight. Fast runner. Good climber (on cave walls and ceilings). Pseudanophthalmus thomasi Barr, 1981 Ecology. Caves: along stream banks. Wet, muddy soil. Biology. Seasonality: June, November. Adult food, in the field: mites. Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Pseudanophthalmus tiresias Barr, 1959 Ecology. Caves. Moderately moist, muddy soil, rich in organic debris. Occurring on wet rotting wood or on the ground. Biology. Seasonality: December. Dispersal power. Brachypterous, incapable of flight. Fast runner. Climber (on pieces of wood). Pseudanophthalmus transfluvialis Barr, 1985 Ecology. Caves. Biology. Seasonality: September. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus troglodytes Krekeler, 1973 Ecology. Caves. Biology. Seasonality: October. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus tullahoma Barr, 1959 Ecology. Caves: along stream banks. Wet, muddy soil. Sheltering under stones or occurring on rotting wood. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Climber (on pieces of wood). Pseudanophthalmus umbratilis Krekeler, 1973 Ecology. Caves. Biology. Seasonality: January, June-August, December. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus unionis Barr, 1981 Ecology. Caves. Biology. Seasonality: January, August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus valentinei Jeannel, 1949 Ecology. Caves. Biology. Seasonality: August. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus vanburenensis Barr, 1959 Ecology. Caves: along pool margins. Wet soil. Sheltering under stones. Biology. Seasonality: June. Adult food, in the field: cave-cricket eggs. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones.

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Pseudanophthalmus ventus Barr, 1981 Ecology. Caves. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus vicarius Barr, 1965 Ecology. Caves: along stream banks. Sheltering in heaps of rotting twigs, in crevices, and under stones. Biology. Seasonality: July, September. Dispersal power. Brachypterous, incapable of flight. Fast runner. Climber (on pieces of wood). Pseudanophthalmus virginicus (Barr, 1960) Ecology. Caves: along stream banks. Wet soil. Sheltering in heaps of broken stones. Biology. Seasonality: unknown. Tenerals: September. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones. Pseudanophthalmus wallacei Barr, 1981 Ecology. Caves. Biology. Seasonality: February-April. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus youngi donaldsoni Krekeler, 1958 Ecology. Caves. Biology. Seasonality: June, August-September. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudanophthalmus youngi youngi Krekeler, 1958 Ecology. Caves. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Fast runner. Pseudaptinus lecontei (Dejean, 1831) Ecology. Lowlands. Borders of salt marshes; salt flats. Open ground; wet, clayish or muddy soil, rich in organic debris, covered with some vegetation (e.g., Juncus, grass). Nocturnal; sheltering during the day in vegetal debris. Biology. Seasonality: MarchOctober. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Collecting techniques. Treading the soil with the feet; pouring water over the ground; light trapping. References. Leng, 1915 (dispersal power); Kirk, 1969 and 1970 (dispersal power); Davidson, 1995 (dispersal power).

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Pseudaptinus oviceps Van Dyke, 1926 Ecology. Borders of rivers. Moist soil. Nocturnal; sheltering during the day under stones. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Turning stones. Pseudaptinus tenuicollis (LeConte, 1851) Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Pseudomorpha alleni Van Dyke, 1953 Ecology. Unknown. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Pseudomorpha alutacea Notman, 1925 Ecology. Unknown. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Pseudomorpha angustata G.H. Horn, 1883 Ecology. Mountains. Ant nests. Biology. Seasonality: March, June-August. The female is ovoviviparous. Tenerals: July (AZ). Defense-mechanism: as quick in flight as a Cicindela; difficult to pursue. Dispersal power. Macropterous. Frequent flier (when alarmed; to artificial lights at night). Fast runner. Reference. Liebherr and Kavanaugh, 1985 (natural history). Pseudomorpha behrensi G.H. Horn, 1870 Ecology. Unknown. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Pseudomorpha castanea Casey, 1909 Ecology. Unknown. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Pseudomorpha champlaini Notman, 1925 Ecology. Unknown. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight.

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Pseudomorpha consanguinea Notman, 1925 Ecology. Unknown. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Pseudomorpha cronkhitei G.H. Horn, 1867 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Pseudomorpha cylindrica Casey, 1889 Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Pseudomorpha excrucians Kirby, 1825 Ecology. Macrohabitat unknown. On sand (SC). Biology. Seasonality: May-July. Dispersal power. Macropterous, probably capable of flight. Pseudomorpha falli Notman, 1925 Ecology. Unknown. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Pseudomorpha hubbardi Notman, 1925 Ecology. Mountains. Ant nests. Biology. Seasonality: June-September. The female is ovoviviparous. Dispersal power. Macropterous, probably capable of flight. Fast runner. Reference. Liebherr and Kavanaugh, 1985 (biology). Pseudomorpha parallela Van Dyke, 1943 Ecology. Sand dunes. Open ground; dry soil, bare or covered with sparse vegetation. Biology. Seasonality: spring and autumn. Dispersal power. Macropterous, probably capable of flight. Fast runner. Pseudomorpha ruficollis Casey, 1924 Ecology. Unknown. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Pseudomorpha schwarzi Notman, 1925 Ecology. Unknown. Biology. Seasonality: June. Dispersal power. Macropterous, probably capable of flight.

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Pseudomorpha tenebroides Notman, 1925 Ecology. Unknown. Biology. Seasonality: June. Dispersal power. Macropterous, probably capable of flight. Pseudomorpha vandykei Notman, 1925 Ecology. Unknown. Biology. Seasonality: August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Pseudomorpha vicina Notman, 1925 Ecology. Unknown. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Pseudomorpha vindicata Notman, 1925 Ecology. Unknown. Biology. Seasonality: August. Dispersal power. Macropterous, probably capable of flight. Psydrus piceus LeConte, 1846 Ecology. Lowlands and mountains. Coniferous forests (e.g., Pinus, Tsuga, Pseudotsuga, Larix, Abies). Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under the loose, thick (1 cm or more), moist, slightly dehiscent bark of fallen trees (commonly 80 cm in diameter) and logs. Gregarious. Larval habitat: under the loose bark of trees. Biology. Seasonality: February-December. Defense-mechanism: when alarmed, the adult may emit from the pygidial glands a slight Brachinus-like explosion with a strong malodorous smell (less offensive than in Nomius). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Lifting the loose bark of fallen trees or logs. References. Schwarz, 1884 (ecology); Hatch, 1935b (ecology); Lindroth, 1961 (natural history); Noonan, 1967a (ecology); McCleve, 1975 (ecology); Goulet, 1977a (ecology); Dajoz, 1997 (ecology). Pterostichus (Cylindrocharis) acutipes acutipes Barr, 1971 Ecology. Lowlands and mountains. Deciduous or mixed forests: along ravines, rivers, and brooks. Shaded ground; moist soil. Nocturnal; sheltering during the day under large stones. Biology. Seasonality: April-September. Defense-mechanism: when alarmed, the adult escapes by burrowing deep into the soil. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on trees). Strong burrower. Collecting techniques. Pitfall trapping; turning large stones.

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Pterostichus (Cylindrocharis) acutipes kentuckensis Barr, 1971 Ecology. Mountains. Forests. Shaded ground; moist soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: April, June-November. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting techniques. Pitfall trapping; turning stones. Reference. Hoffman, 1998 (ecology, biology). Pterostichus (Hypherpes) adoxus (Say, 1823) Ecology. Lowlands and mountains. Forests (mostly deciduous, e.g., Acer; also mixed, coniferous): along slopes, ravines, brooks, seepages, and roadsides. Shaded ground; moist or wet soil. Nocturnal; sheltering during the day under the loose bark of fallen rotten trees and logs (mostly), in rotten logs, under fallen trees, logs, and stones. Biology. Seasonality: April-November. Tenerals: July (mostly)-August. Adult food, in the field: lepidopterous immature stages. Adult and larval food, in captivity: mealworms. Parasites: nemathelminth worms, ascid mites, and Laboulbeniales (Fungi). Phoretic mites found on adult body. Defense-mechanism: when alarmed, the adult emits a strong smell, similar to that of burnt chocolate. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Frequent climber (on trees and logs). Collecting techniques. Lifting the loose bark of fallen trees or logs; pitfall trapping. References. Blatchley, 1910 (ecology); Levesque et al. 1979 (biology); Reeves et al. 1983 (biology); Bousquet, 1986b (biology); Levesque and Levesque, 1986 (biology). Pterostichus (Bothriopterus) adstrictus Eschscholtz, 1823 Ecology. Lowlands, mountains, subalpine and alpine zones. Forests (mostly coniferous; also mixed, and, more rarely, deciduous), their edges and openings; lakeside forests. Thickets, bushes, and hedges (e.g., Alnus, Betula, Populus). Grasslands, cultivated fields (e.g., wheat, barley, fava bean), vacant lots, gardens, Coastal toundra, moraines, alpine meadows, roadsides, gravel pits, sand pits; upper zone of river and brook banks; sprucebogs, marsh edges, supra-tidal meadows, and seashore. Vicinity of human habitations (rural and urban sites). Greenhouses (occasionally). Shaded or open ground; moderately moist or rather dry soil consisting of gravel (often clay- or humus-mixed), sometimes composed of sand or stones, covered with some vegetation (e.g., grass, weeds), stones or dead leaves. Mostly nocturnal; usually sheltering during the day in leaf litter, in and under logs, under fallen branches, sticks, and boards. Gregarious (in winter). Larval habitat: in rotten logs and leaf litter. Biology. Seasonality: April-November. Copulating pairs: May-June (QU); autumn (occasionally; AB); coitus lasting 12 hours on average. Gravids: May-October; mostly before mid-July (AB, ON, QU). Eggs are laid in rotten logs. Tenerals: late June to September. Adults found overwintering on higher and drier ground, in woods; in logs and rotten tree-stumps, under the loose bark of tree-stumps, under pieces of wood, stones, and in leaf litter. Adult food: lepidopterous eggs and larvae, sawfly cocoons, dipterous larvae (in the field); lepidopterous and sawfly cocoons, elaterids, mealworms, and crushed snails (in captivity). Larval food, in captivity: mealworms. Larvae are cannibalistic in captivity. Predators: toads (in the field) and

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shrews (in captivity). Parasites: mites, nematodes, and fungi. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Frequent in sea- and lakeshore drift material, which indicates flight ability. Occasional climber (on bushes). Strongly suffering from competition for resources with Pterostichus melanarius (western Canada). Collecting techniques. Pitfall trapping; baiting traps with meat; raking the leaf litter; turning drift material. References. Lindroth, 1945, 1955a, 1963a, and 1966 (natural history); Barlow, 1970 (biology); Frank, 1971b (biology); Goulet, 1974 (biology); Kirk, 1975c (ecology, dispersal power); Bousquet and Pilon, 1977 (biology); Levesque et al. 1979 (biology); Bousquet, 1986b (biology); Lindroth, 1986 (ecology); Holliday, 1991 (biology); Niemelä and Spence, 1991 (ecology); Niemelä et al. 1992 (ecology, biology); Dearborne and Donahue, 1993 (dispersal power); Luff, 1993 (biology); Currie and Digweed, 1996 (biology). Pterostichus (Lenapterus) agonus G.H. Horn, 1880 Ecology. Tundra. Open ground; somewhat moist, muddy soil, rich in organic debris, covered with some vegetation (e.g., Eriophorum, Carex). Nocturnal; sheltering during the day. Biology. Seasonality: May-August. Tenerals: August (AK). Dispersal power. Brachypterous, incapable of flight. Moderate runner. References. Lindroth, 1966 (natural history); Bousquet, 1999 (taxonomy). Pterostichus (Hypherpes) alamedae (Casey, 1918) Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) algidus LeConte, 1852 Ecology. Lowlands, mountains, and subalpine zone. Mostly forests (deciduous, mixed, and coniferous): along streams and above sea beaches. Also open places with dense vegetation. Greenhouses (occasionally). Shaded ground; moist soil covered with logs. Nocturnal; sheltering during the day under logs and pieces of wood. Biology. Seasonality: May-November. Tenerals: April-June (WA). Adult food, in captivity: dipterous eggs and tree seeds. Predators: spiders (Black Widow). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning logs. References. Exline and Hatch, 1934 (biology); Johnson et al. 1966 (biology); Lindroth, 1966 (natural history); Kavanaugh, 1992 (natural history). Pterostichus (Hypherpes) amethystinus Mannerheim, 1843 Ecology. Lowlands, mountains, and subalpine zone. Coniferous forests: along streams or above sea beaches. Shaded ground; moist soil covered with logs. Nocturnal; sheltering during the day under and in old logs, and under the loose bark of fallen trees. Biology. Seasonality: February-October. Copulating pairs: February (BC). Tenerals: February, July, November. Adult food, in captivity: dipterous eggs, mealworms, and tree seeds. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collect-

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ing techniques. Pitfall trapping; turning and dismembering old logs. References. Lindroth, 1966 (natural history); Kavanaugh, 1992 (natural history). Pterostichus (Hypherpes) angustus (Dejean, 1828) Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) annosus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) arcanus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Cryobius) arcticola (Chaudoir, 1868) Ecology. Lowlands, mountains, subalpine and alpine zones. Coastal tundra, moraines, and alpine meadows. Open ground; soil composed of stones, gravel, sand or clay, covered with some vegetation.Nocturnal; sheltering during the day under stones and in moss. Biology. Seasonality: May-October. Gravids: July (NH). Tenerals: late July to September (NH, QU). Phoretic mites found on adult body. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning stones; pitfall trapping. Reference. Lindroth, 1966 (ecology, dispersal power); Bousquet, 1999 (biology). Pterostichus (Abacidus) atratus (Newman, 1838) Ecology. Lowlands and mountains. Mostly forests (mixed or deciduous); also neighboring cultivated fields (e.g., tobacco, alfalfa). Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under logs and in leaf litter. Biology. Seasonality: January, April-December. Tenerals: late June (ON), August and October (PA). Adult food, in the field: fresh hickory nut meat from nut scraps left behind by squirrels. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning logs; night collecting with a headlamp. References. Lindroth, 1966 (natural history); Morrill, 1992 (biology); Will et al. 1995 (ecology, dispersal power). Pterostichus (Cryobius) auriga Ball, 1962 Ecology. Mountains. Biology. Seasonality: June-July. Dispersal power. Brachypterous, incapable of flight.

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Pterostichus (Hypherpes) baldwini (Casey, 1924) Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) barbarinus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Cryobius) barryorum Ball, 1962 Ecology. Tundra. Biology. Seasonality: May-June. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) beyeri Van Dyke, 1925 Ecology. Mountains. Forests. Shaded ground; moderately moist soil covered with vegetal debris. Nocturnal; sheltering during the day under well-embedded debris. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting techniques. Pitfall trapping; turning embedded debris. Pterostichus (Cryobius) biocryus Ball, 1962, see Pterostichus (Cryobius) bryanti biocryus Ball, 1962. Reference. Ball and Currie, 1997 (taxonomy). Pterostichus (Cryobius) brevicornis (Kirby, 1837) Ecology. Lowlands, mountains, subalpine and alpine zones. Alpine heaths and meadows, coastal moraines, thickets (Alnus), forest clearings, light or dense forests (coniferous, mixed, deciduous; e.g., Alnus, Betula, Abies, Picea ). Ground open or shaded by bushes (e.g., Alnus) and shrubs; usually rather dry, sometimes moist (in forests) soil covered with grass or dead leaves. Nocturnal; sheltering during the day under stones, dead leaves, moss, pieces of rotten wood, and the loose bark of fallen trees. Gregarious (autumn and winter). Biology. Seasonality: January-December. Gravids: June-July (QU). Copulating pairs: late June (QU). Oviposition: May-July (AK). Tenerals: August (AK, NH), September (QU). Adults found overwintering in partially decayed tree-stumps. Adult food, in captivity: mealworms, small insects, and other arthropods. Larval food, in captivity: mealworms, other insects, dead carabids, meat, vegetables, fruit, and bread. Predators: trouts. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones; raking the leaf litter. References. Lindroth, 1963a and 1966 (natural history); Miller, 1969 (biology); Baust and Miller, 1970 (biology); Kaufmann, 1971 (biology); Baust, 1972 (biology); Bousquet, 1999 (biology).

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Pterostichus (Hypherpes) breviusculus breviusculus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) breviusculus mimus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Cryobius) bryanti biocryus Ball, 1962. Ecology. Mountains. Tundra. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Reference. Ball and Currie, 1997 (taxonomy). Pterostichus (Cryobius) bryanti bryanti (Van Dyke, 1951) Ecology. Mountains. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Reference. Ball and Currie, 1997 (taxonomy). Pterostichus (Cryobius) bryanti bryantoides Ball, 1962 Ecology. Unknown. Biology. Seasonality: probably June (label data “6.9.1920” are ambiguous). Dispersal power. Brachypterous, incapable of flight. Reference. Ball and Currie, 1997 (taxonomy). Pterostichus (Cryobius) bryanti cacumenis Ball, 1966 Ecology. Mountains. A meadow with rather dry soil covered with dense vegetation (e.g., Betula, Dryas, Epilobium), under stones. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. References. Ball, 1966 (ecology); Lindroth, 1966 (natural history); Ball and Currie, 1997 (taxonomy). Pterostichus (Cryobius) bryanti stantonensis Ball, 1966 Ecology. A river bank. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Reference. Ball and Currie, 1997 (taxonomy). Pterostichus (Cryobius) bryanti tiliaceoradix Ball, 1962 Ecology. Mountains. Moraine slopes: along brooks. Open ground; rather dry, sandy soil covered with sparse vegetation (e.g., Epilobium, Artemisia). Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. References. Ball, 1966 (ecology); Lindroth, 1966 (ecology, dispersal power); Ball and Currrie, 1997 (taxonomy).

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Pterostichus (Hypherpes) bucolicus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Cryobius) cacumenis Ball, 1966, see Pterostichus (Cryobius) bryanti cacumenis Ball, 1966. Reference. Ball and Currie, 1997 (taxonomy). Pterostichus (Hypherpes) californicus (Dejean, 1828) Ecology. Mountains. Forests. Damp soil. Nocturnal; sheltering during the day under stones and logs. Biology. Seasonality: April-May, July, September. Adult food, in the field: lepidopterous caterpillars. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning stones and logs. Reference. Riddick and Mills, 1996 (biology). Pterostichus (Hypherpes) caligans G.H. Horn, 1891 Ecology. Mountains. Forests. Damp soil. Nocturnal; sheltering during the day under well-embedded logs and other debris. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting technique. Turning embedded logs. Pterostichus (Pseudoferonina) campbelli Bousquet, 1985 Ecology. Deciduous forests and marshy areas: along streams. Shaded ground; moist soil. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: May-October. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. Reference. Bousquet, 1999 (natural history). Pterostichus (Hypherpes) canallatus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Cryobius) caribou Ball, 1962 Ecology. Lowlands. Tundra: along river banks. Wet or dry soil. Biology. Seasonality: May-September. Dispersal power. Brachypterous, incapable of flight. References. Ball, 1966 (ecology); Lindroth, 1966 (ecology, dispersal power). Pterostichus (Monoferonia) carolinus carolinus Darlington, 1931 Ecology. Mountains. Forests: along stream banks and gorges. Shaded ground; wet soil. Nocturnal; sheltering during the day in leaf litter (mostly), under moss, logs, and stones.

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Biology. Seasonality: July-September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; pitfall trapping. Reference. Barr, 1969 (ecology). Pterostichus (Monoferonia) carolinus fumorum Darlington, 1931 Ecology. Mountains. Forests (Picea). Shaded ground; moderately moist soil. Nocturnal; sheltering during the day in rotten logs or under stones and moss. Biology. Seasonality: July-September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; dismembering rotten logs. Pterostichus (Hypherpes) castaneus (Dejean, 1828) Ecology. Lowlands, mountains, and subalpine zone. Coniferous forests: along stream banks and above sea beaches. Shaded ground; moist or rather dry soil covered with logs. Nocturnal; sheltering during the day under and in rotten logs, under fallen tree trunks, under loose bark and mats of moss growing on fallen tree trunks. Biology. Seasonality: February, May-August. Tenerals: mid-July to late August (WA). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Dismembering rotten logs; pitfall trapping. References. Lindroth, 1966 (natural history); Kavanaugh, 1992 (natural history). Pterostichus (Hypherpes) castanipes (Ménétriés, 1843) Ecology. Mountains. Biology. Seasonality: May-June, November. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Melanius) castor Goulet and Bousquet, 1983 Ecology. Lowlands and mountains. Beaver houses: along ponds and slow streams. Open ground; wet, muddy soil, bare or almost so. Nocturnal; sheltering during the day in the outer layers and under the well-embedded branches of beaver house walls. Gregarious. Associated with Platypatrobus lacustris Darlington and Atranus pubescens Dejean. Biology. Seasonality: May-November. Gravids: May (QU). Tenerals: late June to September (NH, QU). Adult and larval food, in captivity: mealworms. Phoretic mites found on adult body. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Strong burrower. Good swimmer. Favored by laws protecting the beaver. Collecting technique. Inspecting the walls of beaver houses. References. Goulet and Bousquet, 1983 (natural history); Bousquet, 1999 (biology). Pterostichus (Lamenius) caudicalis (Say, 1823) Ecology. Lowlands and mountains. Flood-plain forests (e.g. Acer, Ulmus, Fraxinus); swampforests (e.g., Acer, Salix) bordering lakes and ponds; borders of eutrophic pools and ponds; small brook margins surrounded by shrubs and bushes (e.g., Rhododendron). Shaded

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ground; soft, wet, clayish or muddy soil, rich in organic debris. Nocturnal; sheltering during the day under logs, pieces of wood, dead leaves, stones, and other debris. Gregarious (in winter). Biology. Seasonality: January-December. Copulating pairs: early July (QU). Gravids: late May to early July (QU). Tenerals: August and October. Adults found overwintering under stones, in leaf litter, under the loose bark of fallen trees, logs, and pieces of wood. Active at low temperatures under snow in October-November. Adult and larval food, in captivity: mealworms. Predators: toads. Parasites: Laboulbeniales (Fungi). Phoretic mites found on adult body. Dispersal power. Macropterous. Moderate runner. Favored by spring floods along large rivers. Collecting techniques. Turning logs and pieces of wood; pitfall trapping. References. Blatchley, 1910 (biology); Lindroth, 1955a, 1963a, and 1966 (natural history); Kirk, 1975c (ecology); Erwin, 1981 (natural history); Epstein and Kulman, 1990 (ecology, biology); Bousquet, 1999 (taxonomy, natural history). Pterostichus (Cryobius) chipewyan Ball, 1962 Ecology. Forests (Salix, Alnus). Ground shaded by bushes. Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: May-September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. References. Ball, 1966 (ecology); Lindroth, 1966 (ecology, dispersal power). Pterostichus (Eosteropus) circulosus Lindroth, 1966 Ecology. Borders of bogs and small marshes. Open ground; soft, wet, muddy soil covered with vegetation (mostly Carex; moss). Nocturnal; sheltering during the day in moss and under debris. Biology. Seasonality: June-July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Treading the soil with the feet; turning moss carpets. References. Lindroth, 1966 (natural history); Bousquet, 1999 (taxonomy). Pterostichus (Hypherpes) cochlearis Hacker, 1968 Ecology. Mountains. Biology. Seasonality: September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Pterostichus (Argutor) commutabilis (Motschulsky, 1866) (Pterostichus leconteianus (Lutshnik, 1922)) Ecology. Lowlands and mountains. Borders of ponds, lakes, pools, rivers, and marshes; swamps, meadows, pastures, relict prairies, ski fields, sand pits, gravel pits, and roadsides ditches. Open ground; moderately dry, sand-mixed soil covered with some vegetation (e.g., Equisetum, Juncus, grass). Mostly nocturnal; usually sheltering during the day under dead leaves, dead fallen plants (e.g., Juncus), heaps of hay, stones, logs, pieces of wood, and other debris. Gregarious (in winter). Biology. Seasonality: March-December. Gravids: May (QU). Tenerals: April (rarely), July-August, October. Adults found overwintering on higher and drier ground, in the upper zone of sand pits, gravel pits, and roadsides, on hills, in pastures and meadows, in tree hedges, at the edge of woods, and in

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woods; in burrows dug in the soil (as deep as 5 cm), under stones, in leaf litter, under pieces of wood, in rotten logs, and in moss. Adult and larval food, in captivity: mealworms. Predators: frogs and toads. Also eaten by insectivorous plants (Drosera). Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Occasional flier (in the sunshine). Frequent in lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Moderately favored by human activities. Collecting techniques Raking the leaf litter; turning dead fallen plants; pitfall trapping; turning drift material. References. Blatchley, 1910 (ecology; as erythropus (Dejean)); Lindroth, 1966 (natural history; as leconteianus (Lutshnik)); Bousquet, 1986b (biology; as leconteianus (Lutshnik)); Bousquet, 1999 (taxonomy, natural history); Ball and Bousquet, 2001 (taxonomy). Pterostichus (Hypherpes) congestus (Ménétriés, 1843) Ecology. Lowlands and mountains. Forest edges. Biology. Seasonality: February-March, May. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Pterostichus (Euferonia) coracinus (Newman, 1838) Ecology. Lowlands, mountains, subalpine and alpine zones. Mostly forests (deciduous, mixed, coniferous); forest clearings, edges and roadsides; tree plantations (e.g., Pinus) and orchards. Also wastelands, cultivated fields (e.g., wheat, tobacco, corn), and meadows. Shaded or open ground; usually moderately moist soil (wet in coastal areas) covered with thick leaf litter (in forests) or some vegetation (in open country). Nocturnal; sheltering during the day under dead leaves, stones, and in rotten logs. Biology. Seasonality: January, March-December. Copulating pairs: throughout most of the activity period; coitus lasting 50-53 minutes. Gravids: May-October (QU). Tenerals: early June to mid-August. Adults found overwintering in woods and at their edges; in the soil, under stones, in leaf litter, and in rotten tree-stumps. Adult food: lepidopterous immature stages (in the field); lepidopterous and sawfly cocoons, carabids, mealworms, earthworms, meat, and a corn kernel (in captivity). Larval food, in captivity: mealworms. Predators: toads and crows (in the field); shrews (in captivity). Parasites: braconid larvae, ascid mites, nemathelminth worms, and fungi (Laboulbeniales). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on plants). Moderately favored by human activities. Collecting techniques. Pitfall trapping; baiting traps with meat; raking the leaf litter; turning stones; sweeping plants. References. Blatchley, 1910 (ecology); Lindroth, 1955a, 1963a, and 1966 (natural history); Rivard, 1964a (biology); Barr, 1969 (ecology); Barlow, 1970 (ecology, biology); Randolph et al. 1976 (biology); Bousquet and Pilon, 1977 (biology); Levesque et al. 1979 (biology); Erwin, 1981 (natural history); Reeves et al. 1983 (biology); Levesque and Levesque, 1986 (biology); Bousquet, 1986b (biology) and 1999 (taxonomy). Pterostichus (Phonias) corrusculus LeConte, 1873 Ecology. Lowlands. Swamp-forests, swamps, and low forests: often along lakes and rivers. Also meadows. Shaded ground; moist or wet soil covered with dead leaves or

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moss. Nocturnal; sheltering during the day under stones, logs, moss, among dead grass and other debris. Biology. Seasonality: February-June, September-October, December. Adults found overwintering at the edge of woods; in leaf litter. Parasites: mites. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning stones and logs; pitfall trapping. References. Darlington, 1936 (dispersal power); Lindroth, 1966 (natural history); Bousquet, 1999 (taxonomy). Pterostichus (Melanius) corvinus (Dejean, 1828) Ecology. Lowlands and uplands. Borders of eutrophic marshes; marshy borders of often small ponds and lakes, slow rivers and brooks; swamps, swamp-forests, flood-plain forests; beaver houses. Close to water. Open or shaded ground; soft, wet, clayish or muddy soil, rich in organic debris, covered with dense vegetation (e.g., Typha, Salix, Carex, grass). Mostly nocturnal; usually sheltering during the day under fallen plants, dead leaves, pieces of wood, logs, stones, other debris, and in mud layers. Gregarious (in winter). Biology. Seasonality: March-December. Copulating pairs: May-June (QU). Gravids: early May to late June (NY, QU, VT). Tenerals: late June to October; mostly August-September. Adults found overwintering on higher and drier ground, in forests, on hills, in vacant lots, in the upper zone of roadsides, gravel pits, and sand pits; in the soil (mostly; as deep as 25 cm), in leaf litter and under the loose bark of fallen trees. Adult food, in captivity: mealworms, slugs, and meat. Larval food, in captivity: mealworms. Predators: frogs and toads. Parasites: ascid and halolaelapid mites, and Laboulbeniales (Fungi). Phoretic mites found on adult body. Dispersal power. Macropterous. Occasional flier (in the sunshine). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Treading the soil with the feet; turning dead fallen plants; raking the leaf litter. References. Lindroth, 1966 (natural history); Erwin, 1981 (natural history); Bousquet, 1986b (biology). Pterostichus (Lenapterus) costatus (Ménétriés, 1851) Ecology. Tundra: along margins of dry pools. Open ground; very moist peaty or muddy soil covered with bare spots surrounded by some vegetation (e.g., Eriophorum, Carex). Biology. Seasonality: June-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. References. Lindroth, 1966 (ecology, dispersal power); Bousquet, 1999 (taxonomy). Pterostichus (Hypherpes) craterensis (Hatch, 1949) Ecology. Mountains. Forests. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Pterostichus (Hypherpes) crenicollis LeConte, 1873 Ecology. Lowlands and mountains. Mostly deciduous or coniferous forests (often above sea beaches). Also open sites modified by human activities (e.g., gardens). Shaded

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ground; moderately moist soil. Nocturnal; sheltering during the day under embedded stones and logs, the loose bark of fallen trees, and in leaf litter. Biology. Seasonality: April-May, July-October. Tenerals: mid-July to late August (WA). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Moderately favored by human activities. Collecting techniques. Turning embedded stones and logs; pitfall trapping. References. Lindroth, 1966 (natural history); Kavanaugh, 1992 (natural history). Pterostichus (Hypherpes) cuneatulus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) diabolus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Monoferonia) diligendus (Chaudoir, 1868) Ecology. Lowlands and mountains. Deciduous (mostly; e.g., Acer, Quercus), mixed (e.g., Acer, Quercus, Tsuga), and coniferous forests (rarely; e.g., Picea; WV). Swamp-forests: along brooks, rills, rivulets, springs, and seepages. Associated with ravines and slopes. Cool, shaded ground; well-drained, very wet, often mudddy soil. Nocturnal; sheltering during the day under stones and logs (mostly), under dead leaves, moss, fallen trees and their loose bark, and in rotten logs. Associated with the Two-lined Salamander (Eurycea bislineata bislineta Green) (QU) and Pterostichus superciliosus (WV). Larval habitat: similar to adult, but closer to water. Biology. Seasonality: April-October. Gravids: May-July (QU). Eggs are laid in late July under moss lying on emergent roots and logs (QU). Tenerals: late July to October. Adult food, in the field: lepidopterous immature stages. Adult and larval food, in captivity: mealworms. Parasites: mermithid worms (Nematoda) and fungi (Laboulbeniales). Phoretic mites found on adult body. Defensemechanism: brood care (eggs and larvae) exhibited by females. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on trees). Collecting techniques. Turning stones and logs; pitfall trapping. References. Lindroth, 1966 (natural history); Barr, 1969 (ecology); Bousquet, 1983 and 1986b (biology). Pterostichus (Melanius) ebeninus (Dejean, 1828) Ecology. Lowlands. Swamps, swamp-forests, flood-plain forests, and borders of marshes. Shaded ground; wet, clayish or muddy soil. Nocturnal; sheltering during the day under logs, pieces of wood and other debris. Biology. Seasonality: January-July, October-November. Adults found overwintering in rotten logs. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Turning logs and pieces of wood; night collecting with a headlamp; pitfall trap-

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ping. References. Blatchley, 1910 (ecology, biology); Leng, 1915 (ecology); Löding, 1945 (ecology); Kirk, 1969 (ecology, dispersal power); Goulet and Bousquet, 1983 (biology); Bousquet, 1999 (biology). Pterostichus (Hypherpes) ecarinatus Hatch, 1936 Ecology. Mountains. Forests. Shaded ground; somewhat moist soil. Nocturnal; sheltering during the day in or under old logs (mostly), and under the loose bark of fallen trees. Biology. Seasonality: June-August. Tenerals: early July to late August (BC). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Inspecting old logs. Reference. Lindroth, 1966 (natural history). Pterostichus (Cryobius) empetricola (Dejean, 1828) Ecology. Lowlands and mountains. Vicinity of brooks and rivers, in the shade of bushes (Alnus, Salix), on moist soil. Meadows and heaths, along small streams (Aleutian Islands). Nocturnal; sheltering during the day under dead leaves and stones. Biology. Seasonality: May-July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; turning stones. References. Ball, 1966 (ecology); Lindroth, 1966 (ecology, dispersal power). Pterostichus (Gastrosticta) enodis Bousquet, 1992 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Pterostichus (Hypherpes) esuriens Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Pterostichus (Abacidus) fallax (Dejean, 1828) Ecology. Lowlands. Open deciduous forests (e.g., Quercus, Carya). Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under boards and wood chips. Biology. Seasonality: January-August, October-December. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning pieces of wood. Pterostichus (Hypherpes) falli Van Dyke, 1925 Ecology. Lowlands. Wooded canyons. Nocturnal; sheltering during the day under wellembedded debris. Biology. Seasonality: January. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting techniques. Turning embedded debris; pitfall trapping.

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Pterostichus (Phonias) femoralis (Kirby, 1837) Ecology. Lowlands, mountains, and subalpine zone. Grasslands, prairies, meadows, pastures, cultivated fields (e.g., corn, wheat, barley, oat, flax) and their borders, gardens, vacant lots, roadsides, sand pits, gravel pits, hills; upper zone of river banks and lake shores; hedges, thickets, forest edges, and light forests (e.g., Acer, Ulmus) neighboring fields. Not associated with water. Open or shaded ground; slightly moist or dry, sandy or clayish soil covered with moderate or dense vegetation (e.g., grass and weeds). Nocturnal; sheltering during the day under stones (mostly), logs, pieces of wood, and reindeer moss (Cladonia). Biology. Seasonality: January-December. Gravids: April (QU). Tenerals: early April-May (rarely; CT, QU), late July to November. Adults found overwintering on hills, in pastures, in the upper zone of sand pits and lake shores, in hedges, at the edge of woods, and in woods; under stones (mostly), in the soil, and under dead leaves. Active at low temperatures under snow in October-November. Adult food: dipterous larvae (in the field); grasshopper eggs (mostly), mealworms, and meat (e.g., cat food) (in captivity). Larval food, in captivity: mealworms. Predators: bluebirds. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (to artificial lights at night). Frequent in lake shore drift material, which indicates flight ability. Moderate runner. Strongly favored by human activities. Collecting techniques. Turning stones; pitfall trapping; turning drift material. References. Blatchley, 1910 (biology); Lindroth, 1966 (natural history); Larochelle, 1972k (dispersal power); Kirk, 1975c (ecology, dispersal power); Bousquet, 1986b (biology); Bousquet, 1999 (taxonomy, biology). Pterostichus (Hypherpes) fenyesi fenderi Hacker, 1968 Ecology. Lowlands and mountains. Damp woods. Biology. Seasonality: January. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) fenyesi fenyesi Csiki, 1930 Ecology. Lowlands and mountains. Damp woods. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) fuchsi Schaeffer, 1910 Ecology. Lowlands. Forests. Shaded ground; moderately moist soil. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Pterostichus (Cryobius) gerstlensis Ball, 1962 Ecology. Forests (e.g., Alnus). Shaded ground; soil usually composed of sand-mixed clay. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. References. Ball, 1966 (ecology); Lindroth, 1966 (ecology, dispersal power).

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Pterostichus (Hypherpes) gliscans Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Pterostichus (Hypherpes) gracilior LeConte, 1873 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) gregalis Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Cryobius) haftorni Lindroth, 1969, see Pterostichus (Cryobius) bryanti acumenis Ball, 1966. Reference. Ball and Currie, 1997 (taxonomy). Pterostichus (Abacidus) hamiltoni G.H. Horn, 1880 Ecology. Lowlands and mountains. Open deciduous forests, clearings, roadsides, thickets; hills covered with trees and herbage. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under flat stones and dead leaves. Gregarious (2-4 individuals under each stone). Biology. Seasonality: May-September. Tenerals: mid-June (PA). Defense-mechanism: when alarmed, the adult remains motionless. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones. Pterostichus (Hypherpes) hatchi Hacker, 1968 Ecology. Mountains. Biology. Seasonality: September. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) herculaneus Mannerheim, 1843 Ecology. Lowlands. Coniferous or mixed forests. Greenhouses (occasionally). Shaded ground; rather dry soil. Nocturnal; sheltering during the day under logs (mostly) and the loose bark of trees and tree-stumps. Gregarious (in spring; 2-3 individuals under each piece of loose bark). Biology. Seasonality: June-October. Tenerals: late May to late September. Adults found overwintering in powdery dry tree-stumps, in dry decaying logs, at the base of moss-covered tree-stumps, and under mossy logs. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning logs; lifting loose bark; pitfall trapping. References. Hippisley, 1922 (ecology, biology); Lindroth, 1966 (natural history).

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Pterostichus (Hypherpes) hornii LeConte, 1873 Ecology. Mountains. Forests. Shaded ground; moderately moist soil. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Pterostichus (Cryobius) hudsonicus LeConte, 1863 Ecology. Wooded areas (Alnus, Salix): along rivers. Shaded ground; moist, soil composed of sand-mixed clay. Nocturnal; sheltering during the day among dead leaves. Biology. Seasonality: May-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. References. Ball, 1966 (ecology); Lindroth, 1966 (ecology, dispersal power). Pterostichus (Hypherpes) humboldti Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Pterostichus (Pseudoferonina) humidulus (Van Dyke, 1943) Ecology. Lowlands and mountains. Moist places. Biology. Seasonality: May. Tenerals: September (WA). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Reference. Bousquet, 1999 (natural history). Pterostichus (Hypherpes) humilis Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Cylindrocharis) hypogeus Barr, 1971 Ecology. Mountains. Forests (deciduous): often along ravines. Cool, shaded ground; moderately moist soil. Nocturnal; sheltering during the day under well-embedded stones. Biology. Seasonality: April-August. Tenerals: mid-August. Defense-mechanism: when alarmed, the adult stays motionless for several minutes. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting techniques. Turning embedded stones; pitfall trapping. Reference. Barr, 1971a (ecology, biology). Pterostichus (Hypherpes) idahoae Csiki, 1930 Ecology. Mountains. Forests. Shaded ground; moderately moist soil covered with vegetal debris. Nocturnal; sheltering during the day in leaf litter or in rotten logs. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; dismembering rotten logs; pitfall trapping.

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Pterostichus (Hypherpes) illustris LeConte, 1851 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Pterostichus (Hypherpes) inanis G.H. Horn, 1891 Ecology. Lowlands and mountains. Coniferous forests. Shaded ground; moderately moist soil covered with vegetal debris. Nocturnal; sheltering during the day under pieces of wood. Biology. Seasonality: June-July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning pieces of wood; pitfall trapping. Reference. Papp, 1978 (ecology, biology). Pterostichus (Hypherpes) inermis Fall, 1901 Ecology. Mountains. Biology. Seasonality: November. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) infernalis Hatch, 1936 Ecology. Lowlands and mountains. Forests. Shaded ground; moderately moist soil covered with vegetal debris. Nocturnal; sheltering during the day in leaf litter or in rotten logs. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; dismembering rotten logs; pitfall trapping. Pterostichus (Hypherpes) inopinus (Casey, 1918) Ecology. Lowlands and mountains. Forests. Shaded ground; moderately moist soil covered with vegetal debris. Nocturnal; sheltering during the day in leaf litter or in rotten logs. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; dismembering rotten logs; pitfall trapping. Pterostichus (Hypherpes) intectus (Casey, 1918) Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Euferonia) iripennis (Nicolay and Weiss, 1934) Ecology. Unknown. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Reference. Bousquet, 1999 (taxonomy). Pterostichus (Hypherpes) isabellae LeConte, 1851 Ecology. Mountains. Biology. Seasonality: February-March, August, October. Dispersal power. Brachypterous, incapable of flight.

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Pterostichus (Hypherpes) jacobinus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Orsonjohnsonus) johnsoni Ulke, 1889 Ecology. Mountains. Forests: at the foot of waterfalls (in water spray); on brook margins sprinkled with water. Shaded or open ground; sloping, wet, soil composed of gravel or boulders, sometimes covered with moss. Nocturnal; sheltering during the day under stones, fallen tree bark, and moss. Biology. Seasonality: July. Tenerals: August (OR). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning stones. References. Piper, 1911 (ecology); Hatch, 1933 (ecology). Pterostichus (Hypherpes) kansanus (Casey, 1918) Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Cryobius) kotzebuei Ball, 1962 Ecology. Tundra. Ground open or shaded by Salix-shrubs; peaty or gravelly soil. Nocturnal; sheltering during the day among dead leaves or in dense moss. Biology. Seasonality: June-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. References. Ball, 1966 (ecology); Lindroth, 1966 (ecology, dispersal power). Pterostichus (Hypherpes) laborans Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) lacertus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Euferonia) lachrymosus (Newman, 1838) Ecology. Lowlands and mountains. Deciduous forests (mostly; e.g., Acer, Quercus); mixed forests; coniferous forest (e.g., Picea, Abies; high elevations in NC, WV); forest clearings and roadsides. Shaded ground; moderately moist, often gravelly or stony soil. Nocturnal; sheltering during the day under stones (mostly), logs, dead leaves, fallen trees and their loose bark, and in rotten wood. Biology. Seasonality: April-November. Copulating pairs: May-June (QU); coitus lasting up to 46 minutes. Gravids: June (QU). Tenerals: mid-July (VA). Adult food: lepidopterous immature stages (in the field); lepidopterous caterpillars (frequently), chrysomelids, an ant larva, and

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mealworms, worms, meat, and fruit (in captivity). Larval food, in captivity: mealworms, worms, and meat. Predators: toads. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; baiting traps with meat or butyric acid; turning stones. References. Lindroth, 1966 (natural history); Barr, 1969 (ecology); Levesque et al. 1979 (biology); Erwin, 1981 (natural history); Bousquet, 1986b (biology) and 1999 (taxonomy). Pterostichus (Hypherpes) lama (Ménétriés, 1843) Ecology. Lowlands and mountains. Coniferous forests (e.g., Pseudotsuga, Pinus). Shaded ground; moderately moist soil covered with conifer needles or moss. Nocturnal; sheltering during the day mostly in damp decaying logs, tree-stumps and standing trees, also under logs and under the loose bark of fallen trees. Biology. Seasonality: MarchApril, June-August, October. Tenerals: August (BC), October (WA). Adult food: termites (mostly), pentatomid bugs, cerambycid larvae, lucanids, and rhipicerids (in the field); beetles and Pseudotsuga-seeds (in captivity). Predators: owls and skunks. Defense-mechanism: when alarmed, the adult emits a strong malodorous smell, more powerful than that of the large associated tenebrionids (Coelocnemis and Eleodes). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Dismembering rotten logs and tree-stumps; pitfall trapping. References. Linsley and Michener, 1943 (biology; as Pterostichus ater LeConte); La Rivers, 1946 (ecology, biology; as Feronia ater LeConte); Lindroth, 1966 (natural history); Kavanaugh, 1992 (natural history); Weber and Kavanaugh, 1992 (ecology). Pterostichus (Hypherpes) lanei (Hatch, 1935) Ecology. Lowlands and mountains. Coniferous rainforests. Shaded ground; wet soil, rich in humus. Nocturnal; sheltering during the day in the tunnels of small rodents and under large well-embedded logs. Biology. Seasonality: June, September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting techniques. Pitfall trapping; inspecting the tunnels of small rodents; turning large embedded logs. Reference. Hatch, 1935a (ecology). Pterostichus (Pseudoferonina) lanei Van Dyke, 1925 Ecology. Coastal mountains. Coniferous rainforests: often along river banks. Shaded ground; wet soil covered with thick leaf litter, humus, and moss. Nocturnal; sheltering during the day under well-embedded logs, stones, and fallen tree-bark. Biology. Seasonality: May, July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting techniques. Pitfall trapping; turning embedded logs. Pterostichus (Hypherpes) lassulus (Casey, 1920) Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner.

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Pterostichus (Hypherpes) lobatus Hacker, 1968 Ecology. Lowlands and mountains. Biology. Seasonality: March. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Paraferonia) lubricus LeConte, 1852 Ecology. Mountains. Coniferous forests (e.g., Tsuga) covered with ravines and gorges: along very narrow brooks (less than 1 m wide), springs, and seepages. Shaded ground; wet soil. Nocturnal; sheltering during the day mostly in small mats of dead leaves piled up by periods of high water, also under stones and under moss. Biology. Seasonality: March-July, September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; pitfall trapping. References. Barr, 1969 (ecology); Hoffman, 1998 (ecology, biology). Pterostichus (Pseudomaseus) luctuosus (Dejean, 1828) Ecology. Lowlands and mountains. Margins of eutrophic marshes, ponds, lakes, and slow rivers and brooks; swamps, swamp-forests, and flood-plain forests; beaver houses. Shaded ground; often soft, wet, clayish or muddy soil, rich in organic debris, covered with some vegetation (e.g., Typha, Salix). Nocturnal; sheltering during the day under dead leaves and fallen plants (mostly), pieces of wood, and stones. Gregarious. Biology. Seasonality: March-November. Copulating pairs: April-July (QU). Gravids: June (QU). Tenerals: April (rarely; QU), August-November. Adults found overwintering on higher and drier ground, in woods, at the edge of woods, on hills and hillocks, and in vacant lots; mostly in the soil (as deep as 50 cm) and under well-embedded stones, also under dead leaves, pieces of wood, and in rotten standing tree-stumps. Adult and larval food, in captivity: diplopods and mealworms. Predators: toads. Parasites: nemathelminth worms, halolaelapid and ascid mites, and Laboulbeniales (Fungi). Phoretic mites found on adult body. Dispersal power. Macropterous. Occasional flier (to artificial lights). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber. Strong burrower. Favored by spring floods along rivers. Collecting techniques. Treading the vegetation into the water; raking the leaf litter; turning fallen dead plants. References. Snider, 1984 (biology); Bousquet, 1986b (biology); Davidson 1988b (ecology). Pterostichus (Bothriopterus) lustrans LeConte, 1851 Ecology. Lowlands and mountains. Brook borders, orchards (e.g., apple), and pinewoods. Greenhouses (occasionally). Open ground; rather dry soil. Nocturnal; sheltering during the day under logs and stones. Biology. Seasonality: January-March, May-September. Tenerals: September (WA). Adult food, in captivity: lepidopterous caterpillars. Economic importance: may play an important role in the control of the Codling Moth. Predators: thrushes. Dispersal power. Macropterous. Moderate runner. Collecting techniques. Pitfall trapping; turning logs or stones. References. Lindroth, 1966 (natural history); Riddick and Mills, 1994 (ecology, biology).

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Pterostichus (Hypherpes) malkini (Hatch, 1953) Ecology. Lowlands. Coniferous rainforests. Shaded ground; wet soil, rich in dead leaves and humus. Nocturnal; sheltering during the day under well-embedded logs. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting techniques. Turning embedded logs; pitfall trapping. Pterostichus (Monoferonia) mancus (LeConte, 1852) Ecology. Mountains. Forests: along gorges. Shaded ground; moist soil. Nocturnal; sheltering during the day under dead leaves, moss, logs, and stones. Biology. Seasonality: June-July, September. Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; pitfall trapping. References. Barr, 1969 (ecology); Bousquet, 1999 (taxonomy). Pterostichus (Monoferonia) mancus plethorus Darlington, 1931, see Pterostichus (Monoferonia) mancus (LeConte, 1852). Reference. Bousquet, 1999 (taxonomy) Pterostichus (Cryobius) mandibularoides Ball, 1966 Ecology. Wooded areas: often near rivers. Ground shaded by shrubs (e.g., Alnus, Salix); sandy soil covered with leaf litter. Nocturnal; sheltering during the day among dead leaves. Biology. Seasonality: May. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. References. Ball, 1966 (ecology); Lindroth, 1966 (ecology, dispersal power). Pterostichus (Hypherpes) marinensis Hacker, 1968 Ecology. Lowlands and mountains. Biology. Seasonality: February, July. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) mattolensis Hacker, 1968 Ecology. Lowlands and mountains. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Morphnosoma) melanarius (Illiger, 1798) Ecology. Lowlands, mountains, subalpine and alpine zones. Gardens, cultivated fields (e.g., corn, asparagus, alfalfa, cabbage, barley, fava bean, dry bean, fescue, patato, carrot, clover, sainfoin, raspberry), vacant lots, golf courses; rangelands, meadows, grasslands, sand pits, gravel pits, and roadsides; hedges, orchards (e.g., apple, peach), thickets (e.g., Alnus, Populus), forest edges and clearings, tree plantations (e.g., Pinus),

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light forests (mostly deciduous; mixed, coniferous); upper zone of river and brook banks, lake shores, and reservoirs. Vicinity of human habitations (urban and rural sites). Open or slightly shaded ground; moderately moist, often sandy, sometimes clayish soil covered with moderate or dense vegetation. Active during June and July in the first part of the night, but more active during the day toward the end of August; quite active on rainy or cloudy days. Nocturnal; sheltering during the day under stones, pieces of wood, dead leaves, heaps of hay, in moss, under fallen trees and other debris. Biology. Seasonality: April-November. Copulating pairs: July-October (QU); coitus lasting 50-150 minutes. Gravids: May-October, mostly mid-July to September (ON, QU). Tenerals: April-August; mostly the first half of the summer. Adults found overwintering in cultivated fields, in the upper zone of roadsides and sand pits; mostly in the soil (as deep as 20 cm) and under stones, also under logs, fallen pieces of bark and plant rosettes (Verbascum), and in leaf litter. Adult food: lepidopterous caterpillars, dipterous larvae and pupae, earwigs, aphids, gryllids, and chrysomelid eggs (in the field); lepidopterous caterpillars, dipterous pupae and adults, chrysomelid eggs and larvae, weevil immature stages and adults, scarabaeid larvae, carabid larvae and dead adults, mealworms, earthworms, young salamanders, small dead invertebrates, meat, plant and tree seeds, grass tissue (in captivity). Adults are voracious feeders, killing prey even when satiate. Larval food, in captivity: mealworms. Economic importance: play an important role in the control of earwigs and, possibly also, of lepidopterous caterpillars; sometimes a pest of strawberry fruits and Douglas fir seeds. Predators: toads (frequently), frogs, and foxes. Parasites: mites, tachinids, braconids, nematodes, and nematomorphs. Defense-mechanism: when alarmed, the adult may emit a malodorous spray from the pygidial glands. Dispersal power. Wing-dimorphic: macropterous (mostly) or brachypterous (incapable of flight). Mostly brachypterous (Europe). The frequency of macropterous forms in North America is a sign of recent arrival and subsequent dispersion of this species; in stable, well-established populations, brachypterous forms are frequent. Occasional flier (at sunset; to artificial lights at night). Frequent in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on trees). Clearly effective as a colonist. Strongly reducing populations of Pterostichus adstrictus in urban and agricultural areas (western Canada) through competition for resources. Strongly favored by human activities. Collecting techniques. Turning stones and pieces of wood; pitfall trapping; turning drift material; using nearly empty soft drink bottles as traps. References. Lindroth, 1945 and 1963a (natural history); Rivard, 1964c (biology), 1965 (dispersal power), and 1966 (biology); Briggs, 1965 (biology); Lindroth, 1966 (natural history); Barlow, 1970 (biology); Krehan, 1970 (biology); Cornic, ° 1975 (biology); Tomlin, 1975a and 1975b (biology); Randolph et 1973 (biology); Hurka, al. 1976 (biology); Holliday and Hagley, 1978 (biology); Larochelle, 1978c (dispersal power); Ericson, 1979 (ecology); Levesque et al. 1979 (biology); Jørum, 1980 (biology); Reeves et al. 1983 (biology); Desender et al. 1985 (ecology); Wallin, 1985 (biology, dispersal power); Bousquet, 1986b (biology); Lindroth, 1986 (natural history); Levesque and Levesque, 1987 (biology); Ovaska and Smith, 1988 (biology); Wallin and Ekblom, 1988 (dispersal power); Wheater, 1988 (biology); Koch, 1989 (ecology); Wallin, 1989 (biology, dispersal power); Niemelä and Spence, 1991 (natural history); Luff, 1993 (biology); Basedow, 1994 (biology); Boivin and Hance, 1994 (biology); Dülge, 1994 (biology);

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Levesque and Levesque, 1994 (biology); Wallin and Ekblom, 1994 (biology, dispersal power); Will et al. 1995 (dispersal power). Pterostichus (Hypherpes) menetriesii LeConte, 1873 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) mercedianus (Casey, 1918) Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) miscellus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Eosteropus) moestus (Say, 1823) Ecology. Mountains. Deciduous (mostly; e.g., Quercus, Acer), mixed, and coniferous forests (e.g., Abies, Picea, Pinus): along slopes, ravines, gorges, and streams. Shaded ground; wet soil. Nocturnal; sheltering during the day in rotten logs and fallen trees (mostly), under logs and stones, in standing trees, under the loose bark of standing tree-stumps, and in leaf litter. Biology. Seasonality: April-October. Copulating pairs: late July (KY). Tenerals: early July to mid-August (KY, PA). Adult food, in the field: lepidopterous immature stages. Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Dismembering rotten logs and fallen trees; pitfall trapping. References. Hamilton, 1884 (ecology); Lindroth, 1966 (natural history); Barr, 1969 (ecology); Erwin, 1981 (natural history); Bousquet, 1986b (biology) and 1999 (taxonomy, biology). Pterostichus (Hypherpes) morionides (Chaudoir, 1868) Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Gastrosticta) mutoides Bousquet, 1992 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Bothriopterus) mutus (Say, 1823) Ecology. Lowlands and mountains. Light forests (mostly deciduous; e.g., Acer, Quercus; also mixed and coniferous) and tree plantations (e.g., Pinus). Forest edges, roads, clearings, openings, and roadsides. Vicinity of forests: hedges, thickets (e.g., Juniper), wastelands, cultivated fields, sand pits, and gravel pits. Open or slightly shaded ground; rather dry soil consisting of gravel, sand or clay-loam, covered with grass (in open areas) or thick leaf litter (in forests). Nocturnal; sheltering during the day under dead

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leaves and stones (mostly), pieces of wood, logs, fallen trees, fallen pieces of bark, and reindeer moss (Cladonia). Gregarious (in winter). Larval habitat: under dry cow dung. Biology. Seasonality: January, March-December. Copulating pairs: April (QU). Gravids: mid-April to early July (IL, QU), October (once, QU). Tenerals: March-September; mostly August (QU). Adults found overwintering on higher and drier ground, at the edge of woods, in woods, and in vacant lots; mostly in the soil and under well-embedded stones, also under the loose bark of fallen trees and in leaf litter. Adult food: lepidopterous caterpillars (in the field); lepidopterous caterpillars, diplopods, mealworms, and meat (in captivity). Larval food, in captivity: mealworms. Predators: frogs and toads. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strong burrower. Moderately favored by human activities. Collecting techniques. Raking the leaf litter; turning stones; pitfall trapping. References. Schaupp, 1881a (biology); Blatchley, 1910 (biology); Gilbert, 1957 (biology); Lindroth, 1963a and 1966 (natural history); Barlow, 1970 (biology); Liebherr and Mahar, 1979 (biology); Bousquet, 1986b (biology); Levesque and Levesque, 1986 (biology). Pterostichus (Hypherpes) neobrunneus Lindroth, 1966 Ecology. Mountains. Forests (mostly) and open places. Mostly shaded ground; often rather dry soil. Biology. Seasonality: May-September. Tenerals: July, September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Reference. Lindroth, 1966 (natural history). Pterostichus (Hypherpes) nigrocaeruleus Van Dyke, 1925 Ecology. Lowlands and mountains. Forests: along cold streams. Shaded ground; moist, gravelly soil. Nocturnal; sheltering during the day under well-embedded stones. Biology. Seasonality: May, August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting techniques. Turning embedded stones; pitfall trapping. Reference. Van Dyke, 1925 (ecology). Pterostichus (Cryobius) nivalis (R.F. Sahlberg, 1844) Ecology. Lowlands and mountains. Tundra. Open ground; rather dry, loamy soil, often covered with thick moss. Biology. Seasonality: June-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. References. Ball, 1966 (ecology); Lindroth, 1966 (ecology, dispersal power). Pterostichus (Euferonia) novus Straneo, 1944 Ecology. Lowlands. Deciduous forests (e.g., Quercus, Acer, Fagus), mesic cottonwood forests (Populus deltoides Marshall), drier upland forests (Quercus), alvar shrublands, alvar pavements, alvar savanna grasslands, alvar grasslands, and mesic old fields. Shaded or open ground; moderately moist or dry soil covered with dead leaves or with some

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vegetation (e.g., grass). Nocturnal; sheltering during the day in leaf litter, under logs, and under stones. Biology. Seasonality: April-September. Tenerals: June-July. Adult food, in captivity: diplopods. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking the leaf litter; turning logs and stones. References. Lindroth, 1966 (natural history); Kirk, 1975c (ecology); Erwin, 1981 (natural history); Epstein and Kulman, 1990 (ecology, biology); Bouchard et al. 1998 (ecology, biology); Bousquet, 1999 (taxonomy). Pterostichus (Gastrosticta) obesulus LeConte, 1873 Ecology. Lowlands. Biology. Seasonality: January, April. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) obsidianus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) occultus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Gastrosticta) ophryoderus (Chaudoir, 1878) Ecology. Lowlands. Cypress-swamps: along river banks. Wet soil. Biology. Seasonality: February, June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Pterostichus (Hypherpes) ordinarius Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Bothriopterus) oregonus LeConte, 1861 Ecology. Mountains. Open forests. Dry soil. Biology. Seasonality: June, August. Tenerals: August (WA). Dispersal power. Macropterous. Moderate runner. References. Lindroth, 1966 (natural history); Bousquet, 1999 (dispersal power). Pterostichus (Hypherpes) ovalipennis Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Feronina) palmi Schaeffer, 1910 Ecology. Mountains. Coniferous forests (e.g., Picea, Abies): along streams. Shaded ground; wet, well drained soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: June-September. Tenerals: September. Dispersal power. Brachypterous, inca-

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pable of flight. Moderate runner. Collecting techniques. Turning stones; pitfall trapping. Reference. Barr, 1969 (ecology). Pterostichus (Hypherpes) panticulatus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Pterostichus (Cryobius) parasimilis Ball, 1962 Ecology. Lowlands and mountains. Tundra. Dry, peaty soil covered with some shrubs and bushes (e.g., Betula, Salix, Vaccinium, Lupinus, Rubus). Biology. Seasonality: JuneAugust. Dispersal power. Brachypterous, incapable of flight. Moderate runner. References. Ball, 1966 (ecology); Lindroth, 1966 (ecology, dispersal power). Pterostichus (Hypherpes) parens Casey, 1913, see Pterostichus (Hypherpes) ybousqueti Berlov, 1999. Reference. Berlov, 1999 (taxonomy). Pterostichus (Phonias) patruelis (Dejean, 1831) Ecology. Lowlands, mountains, subalpine and alpine zones. Swamps, swamp-forests, flood-plain forests, Sphagnum-bogs; borders of eutrophic marshes (e.g., Carex), ponds, lakes, slow rivers and brooks; wet meadows, alpine meadows; beaver houses. Open or slightly shaded ground; soft, wet, clayish, muddy or peaty soil covered with moss (mostly) or vegetal debris. Mostly nocturnal; usually sheltering during the day in Sphagnum-carpets (mostly), under vegetal debris (e.g., grass, Typha-stems, dead leaves), among the roots of grass bunches, under logs, pieces of wood, stones, and in rotten logs. Gregarious (in winter). Biology. Seasonality: January-December. Gravids: June (QU). Tenerals: April (rarely; QU), late June to November; mostly August. Adults found overwintering on higher and drier ground, in woods, at the edge of woods, in hawthorn hedges, in the upper zone of lake shores, and in fields; mostly under stones and in leaf litter, also under pieces of wood and in standing rotten tree-stumps; sometimes in large aggregations (up to 30 individuals under a single stone). Active at low temperatures under snow in October-November. Adult and larval food, in captivity: mealworms. Predators: frogs, crows, starlings, and thrushes. Parasites: halolaelapid mites. Phoretic mites found on adult body. Defense-mechanism: when alarmed, the adult may dive into the water and remain submerged for one minute. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (during the day; to artificial lights at night; to artificial lights in captivity.) Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants and tree-stumps). Good swimmer and diver. Collecting technique. Treading moss carpets into the water. References. Darlington, 1936 (dispersal power); Lindroth, 1955a, 1963a, and 1966 (natural history); Larochelle, 1972k

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(dispersal power); Bousquet, 1986b (biology); Levesque and Levesque, 1987 (biology); Bousquet, 1999 (taxonomy, biology). Pterostichus (Bothriopterus) pensylvanicus LeConte, 1873 Ecology. Lowlands and mountains. Deciduous (mostly), mixed, and coniferous forests, their edges, roads, clearings, and small openings; tree plantations, thickets (e.g., Alnus, Cornus, Acer); upper banks of rivers. Vicinity of human habitations (urban and rural sites). Shaded ground; moderately moist, usually gravelly soil covered which thick leaf litter. Nocturnal; sheltering during the day mostly under dead leaves (particularly in spring and in autumn), fallen branches, logs and fallen trees (especially in summer), also under stones and among moss, rarely under the loose bark of fallen trees. Gregarious (in winter). Larval habitat: under dead leaves and in the soil. Biology. Seasonality: FebruaryNovember. Copulating pairs: April-June; October (AB; young adults). Gravids: mid-April to mid-July (AB, QU). Tenerals: April-June, August-October; mostly August. Adults found overwintering on higher and drier ground, in woods and at the edge of woods; mostly under stones, in leaf litter and in the soil (as deep as 10 cm), sometimes in rotten branches, logs and tree-stumps, under the loose bark of tree trunks and fallen trees; sometimes in aggregations of 20 individuals or more. Active at low temperatures under snow in October-November. Adult food: lepidopterous caterpillars (in the field); sawfly cocoons, dead carabids and tenebrionids, mealworms, snails, earthworms, and meat (in captivity). Larval food, in captivity: mealworms. Economic importance: plays an important role in the control of the Spruce Budworm. Predators: toads and frogs. Parasites: ascid mites, tachinid larvae, nematodes, and fungi. Adult body infested by phoretic mites. Defense-mechanism: when alarmed, the adult escapes by burrowing in leaf litter or by feigning death. Dispersal power. Macropterous. Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on logs, tree-stumps, trees). Moderately favored by human activities. Collecting techniques. Pitfall trapping; raking the leaf litter. References. Lindroth, 1955a, 1963a, and 1966 (natural history); Rivard, 1964c (biology); Barlow, 1970 (biology); Freitag and Poulter, 1970 (biology); Frank, 1971b (biology); Goulet, 1974 (biology); Randolph et al. 1976 (biology); Levesque et al. 1979 (biology); Liebherr and Mahar, 1979 (biology); Reeves et al. 1983 (biology); Bousquet, 1986b (biology); Levesque and Levesque, 1986 (biology); Epstein and Kulman, 1990 (ecology, biology); Holliday, 1991 (biology); Niemelä and Spence, 1991 (ecology); Niemelä et al. 1992 (ecology, biology); Dearborne and Donahue, 1993 (biology). Pterostichus (Hypherpes) pergracilis (Casey, 1920) Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Pterostichus (Abacidus) permundus (Say, 1830) Ecology. Lowlands. Open forests (deciduous, e.g., Quercus, Carya; mixed, e.g., Quercus, Pinus; sometimes coniferous, e.g., Pinus), groves, and thickets: often along ravines and

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streams. Cultivated fields (e.g., corn, soybean, cotton, oat, alfalfa, sweet clover), old fields, prairies, lawns, vacant lots, fencerows, and upper zone of brook banks. Shaded or open ground; moist soil consisting of sand or clay loam, covered with leaf litter (in wooded areas) or moderate to dense vegetation (in open areas). Nocturnal; sheltering during the day under logs, pieces of wood, and stones. Biology. Seasonality: JanuaryDecember. Tenerals: June-July, September. Adults found overwintering (SC). Adult food: lepidopterous caterpillars, hymenopterans, other insects, fungi, and vegetal tissue (in the field); weevils, hymenopterous larvae, mealworms, and dog food (in captivity). Larval food, in captivity: mealworms and dog food. Predators: woodpeckers (flickers). Parasites: tapeworms. Defense-mechanism: when alarmed, the adult may emit a Brachinus-like smoke from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Moderately favored by human activities. Collecting techniques. Pitfall trapping; turning logs and pieces of wood; light trapping. References. Blatchley, 1910 (ecology, biology); Kirk, 1970 (biology) and 1975c (ecology); Allen and Thompson, 1977 (ecology, biology); Barney and Armbrust, 1980 (biology). Pterostichus (Cryobius) pinguedineus (Eschscholtz, 1823) Ecology. Lowlands, mountains, subalpine and alpine zones. Forested areas: along river banks. Moraines and alpine meadows: near small trees. Shaded or open ground; moist or wet, stony or gravelly soil covered with some vegetation. Nocturnal; sheltering during the day under dead leaves or stones. Biology. Seasonality: June-October. Tenerals: late August and September (NH). Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; turning stones; pitfall trapping. References. Ball, 1966 (ecology); Lindroth, 1966 (ecology, dispersal power); Bousquet, 1999 (biology). Pterostichus (Hypherpes) placerensis (Casey, 1918) Ecology. Mountains. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) planctus LeConte, 1852 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Cryobius) planus (J.R. Sahlberg, 1887) Ecology. Mountains. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) plutonicus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight.

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Pterostichus (Monoferonia) primus Darlington, 1931 Ecology. Mountains. Coniferous forests (e.g., Picea). Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under stones, logs, and moss. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning stones and logs; pitfall trapping. References. Barr, 1969 (ecology); Bousquet, 1999 (natural history). Pterostichus (Hypherpes) protensiformis (Casey, 1924) Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) protensipennis (Casey, 1918) Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) protractus LeConte, 1860 Ecology. Mountains, subalpine and alpine zones. Coniferous (mostly; e.g., Pinus) and mixed forests (e.g., Pinus, Populus): along brooks. Sagebrush areas, subalpine and alpine meadows. Shaded or open ground; moist soil. Nocturnal; sheltering during the day under logs, stones, boards, in leaf litter (e.g., Pinus), and at the base of trees. Biology. Seasonality: April-October. Tenerals: June-September. Adult food, in captivity: snails. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning logs and stones; pitfall trapping. Reference. Lindroth, 1966 (natural history). Pterostichus (Hypherpes) pumilus pumilus Casey, 1913 Ecology. Lowlands and mountains. Coniferous (e.g., Pseudotsuga, Abies, Thuja), mixed, and deciduous forests (e.g., Acer); forest clearings. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under pieces of wood, dead leaves, stones, and in the tunnels of small rodents. Biology. Seasonality: June-September. Copulating pairs: mid-July (OR). Tenerals: late July to late August (BC, WA). Adult food, in captivity: tree seeds (Pseudotsuga). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting techniques. Turning pieces of wood; raking the leaf litter; pitfall trapping. Reference. Lindroth, 1966 (natural history). Pterostichus (Hypherpes) pumilus willamettensis Hacker, 1968 Ecology. Lowlands. Forests (e.g., Pseudotsuga, Tsuga, Quercus). Damp soil. Biology. Seasonality: January. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping.

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Pterostichus (Lenapterus) punctatissimus (Randall, 1838) Ecology. Lowlands, mountains, subalpine and alpine zones. Coniferous forests (e.g., Picea, Abies), mixed forests (e.g., Abies, Betula), thickets (e.g., Alnus, Cornus); moraines covered with dwarf Betula and ericaceous plants (coastal tundra), and alpine meadows (e.g., ericaceous plants). Ground shaded by trees, shrubs, and bushes (mostly) or open; moderately moist or dry soil covered with thick layer of dead leaves or branches. Nocturnal; sheltering during the day in leaf litter, under fallen branches, fallen pieces of bark, in moss, under and in rotten logs, also under stones and boards. Biology. Seasonality: May-October. Copulating pairs: June-August (QU); coitus lasting 3 hours. Gravids: May to late July, mostly July (NF, QU); July-August (AB). Tenerals: mid-May to September. Adult food, in captivity: sawfly cocoons, lepidopterous caterpillars, mealworms, and meat. Larval food, in captivity: mealworms. Predators: toads. Phoretic mites found on adult body. Defense-mechanism: when alarmed, the adult may emit a strong smell, similar to burnt chocolate, that clings deeeply to the hands when the beetle is seized. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (in malaise traps). Collecting techniques. Raking the leaf litter; turning pieces of wood; pitfall trapping; using malaise traps. References. Lindroth, 1955a, 1963a, and 1966 (natural history); Bousquet, 1986b (biology) and 1999 (taxonomy, biology). Pterostichus (Gastrosticta) punctiventris (Chaudoir, 1878) Ecology. Lowlands. Biology. Seasonality: January, March, May, October-December. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Pterostichus (Gastrosticta) putus Casey, 1913 Ecology and Biology. Unknown. Biology. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Pterostichus (Euferonia) relictus (Newman, 1838) Ecology. Lowlands and mountains. Deciduous and mixed forests: often along slopes, ravines, rivers, and brooks. Shaded ground; moist soil. Nocturnal; sheltering during the day under stones, logs, and fallen pieces of bark. Biology. Seasonality: April-October. Tenerals: mid-June (NC), late July (KY). Parasites: Laboulbeniales (Fungi). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning stones and logs; pitfall trapping. References. Blatchley, 1910 (ecology); Lindroth, 1966 (natural history); Kirk, 1969 (ecology); Bousquet, 1999 (taxonomy). Pterostichus (Hypherpes) restrictus (Casey, 1918) Ecology. Mountains, subalpine and alpine zones. Coniferous and mixed forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under logs.

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Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning logs; pitfall trapping. Pterostichus (Cryobius) riparius (Dejean, 1828) Ecology. Uplands, subalpine and alpine zones. Forests (e.g., Pinus, Populus): often along rivers and brooks. Scrublands and alpine meadows. Vicinity of human habitations (urban and rural sites). Shaded (mostly) or open ground; somewhat moist, sandy or gravelly soil. Nocturnal; sheltering during the day under stones, dead leaves and grass, and fallen branches. Biology. Seasonality: June-August. Tenerals: late May (BC); mostly August (Pacific Northwest). Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning stones; pitfall trapping. References. Ball, 1966 (ecology); Lindroth, 1966 (dispersal power); Niemelä and Spence, 1991 (ecology); Kavanaugh, 1992 (natural history). Pterostichus (Cylindrocharis) rostratus (Newman, 1838) Ecology. Lowlands and mountains. Deciduous forests (e.g., Acer, Quercus), mixed forests, and coniferous forests (e.g., Picea, Abies, Tsuga): often along brooks, slopes, ravines, and roadsides. Shaded ground; wet soil. Nocturnal; sheltering during the day under wellembedded stones, big logs, fallen trees, pieces of wood, fallen pieces of bark, the loose bark of fallen trees, in rotten moist logs and fallen trees, under dead leaves and moss growing on stones. Biology. Seasonality: April-November. Gravids: June-August (QU). Tenerals: mid-July to October. Adult food, in the field: lepidopterous pupae. Adult and larval food, in captivity: mealworms. Predators: toads. Parasites: braconid larvae, ascid mites, and mermithid worms (Nematoda). Phoretic mites found on adult body. Defensemechanism: when alarmed, the adult escapes by burrowing into the soil. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on tree trunks). Strong burrower. Collecting techniques. Turning stones, big logs, and fallen trees; pitfall trapping; baiting traps with molasses or butyric acid. References. Lindroth, 1966 (natural history); Barr, 1969 (ecology); Reeves et al. 1983 (biology); Bousquet, 1986b (biology); Levesque and Levesque, 1986 (biology); Dearborne and Donahue, 1993 (biology). Pterostichus (Hypherpes) rothi (Hatch, 1951) Ecology. Mountains. Coniferous rainforests. Shaded ground; wet soil, with thick leaf litter and humus. Nocturnal; sheltering during the day under large logs well-embedded in thick leaf litter. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting techniques. Turning large embedded logs. Pterostichus (Gastrosticta) sayanus Csiki, 1930 Ecology. Lowlands and mountains. Deciduous (e.g., Quercus, Carya, Fagus), mixed, and coniferous forests (rarely; e.g., Pinus). Shaded or open ground; dry soil. Nocturnal; shel-

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tering during the day under stones, dead leaves, and small pieces of wood. Biology. Seasonality: January-November. Tenerals: early July (TN) and late September (PA). Predators: raccoons. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning stones; raking the leaf litter; pitfall trapping. Pterostichus (Abacidus) sculptus LeConte, 1852 Ecology. Lowlands and mountains. Mostly deciduous and coniferous forests (e.g., Pinus); also cultivated fields (e.g., corn), grasslands, and pastures. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under stones and dead leaves. Biology. Seasonality: March-July, September-December. Adults found overwintering (SC). Adult food, in the field: lepidopterous immature stages. Defense-mechanism: when alarmed, the adult may emit a Brachinus-like smoke from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; turning stones. References. Kirk, 1970 (biology); Morrill, 1992 (ecology, biology). Pterostichus (Hypherpes) scutellaris LeConte, 1873 Ecology. Mountains. Forests. Shaded ground; wet soil. Nocturnal; sheltering during the day in or under old logs. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Inspecting old logs; pitfall trapping. Pterostichus (Hypherpes) sejungendus (Chaudoir, 1868) Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) serripes (LeConte, 1875) Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) setosus Hatch, 1951 Ecology. Mountains. Biology. July. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Pseudoferonina) shulli (Hatch, 1949) Ecology. Mountains. Biology. Seasonality: May. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) sierranus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight.

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Pterostichus (Cryobius) similis Mannerheim, 1852 Ecology. Lowlands. Tundra. Usually sandy soil. Nocturnal; sheltering during the day under dead leaves and plant rosettes (Lupinus). Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. References. Ball, 1966 (ecology); Lindroth, 1966 (ecology, dispersal power). Pterostichus (Pseudoferonina) smetanai Bousquet, 1985 Ecology. Mountains. Steep slope in a forest, along brook margins. Shaded ground; wet soil, sprinkled with water. Nocturnal; sheltering during the day under stones and fallen bark. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning stones and fallen bark. Pterostichus (Cryobius) soperi Ball, 1966 Ecology. Lowlands. Biology. Seasonality: September. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Pseudoferonina) spathifer Bousquet, 1992 Ecology. Mountains. Brook margins. Moist soil composed of alluvial cobbles partly embedded in sand and covered with moss. Nocturnal; sheltering during the day under cobbles. Biology. Seasonality: June, September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting technique. Turning cobbles. Pterostichus (Hypherpes) sphodrinus LeConte, 1863 Ecology. Mountains. Coniferous or mixed forests. Shaded ground; moderately moist soil covered with vegetal debris. Nocturnal; sheltering during the day under logs. Biology. Seasonality: May-August. Tenerals: early May (ID), late July and late August (BC). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning logs; pitfall trapping. Reference. Lindroth, 1966 (natural history). Pterostichus (Hypherpes) sponsor Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) spraguei LeConte, 1873 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight.

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Pterostichus (Cryobius) stantonensis Ball, 1966, see Pterostichus (Cryobius) bryanti stantonensis Ball, 1966. Reference. Ball and Currie, 1997 (taxonomy). Pterostichus (Hypherpes) stapedius Hacker, 1968 Ecology. Mountains. Biology. Seasonality: May. Dispersal power. Brachypterous, incapable of flight. Reference. Bousquet, 1999 (taxonomy). Pterostichus (Hypherpes) stapedius yosemitensis Hacker, 1968, see Pterostichus (Hypherpes) yosemitensis Hacker, 1968. Reference. Bousquet, 1999 (taxonomy). Pterostichus (Phonias) strenuus (Panzer, 1797) Ecology. Lowlands. Grasslands (mostly), cultivated fields, and forests. Open or shaded ground; moderately moist soil covered with some grass (in open areas), bare or covered with dead leaves (in forests). Mostly nocturnal; sheltering during the day under dead leaves, moss, and debris. Sometimes active on roads (Europe). Biology. Seasonality: June-August. Tenerals: early July (QU). Adult food, in the field (Europe): insects and vegetal tissue. Dispersal power. Brachypterous, incapable of flight (North America). Europe: brachypterous (mostly; incapable of flight) or macropterous. Moderate runner. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Raking the leaf litter; pitfall trapping. Using window traps (Europe). References. Larsson, 1939 (biology); Lindroth, 1945, 1955a, 1963a, 1966, and 1986 (natural histo° ry); Greenslade, 1965 (ecology, biology); Bousquet, 1986b (biology); Hurka, 1986b (biology); Koch, 1989 (ecology); Bousquet, 1999 (taxonomy). Pterostichus (Euferonia) stygicus (Say, 1823) Ecology. Lowlands and mountains. Mostly forests (deciduous, e.g., Acer, Quercus; mixed; rarely coniferous): often along streams. Also their edges, clearings, and roadsides. Adjoining hedges, cultivated fields (e.g., corn, soybean), grasslands (short-grass), relict prairies, and fencerows. Shaded (mostly) or open ground; wet or moist soil covered with thick leaf litter (in forests) or moderate to dense vegetation (in open areas). Nocturnal; sheltering during the day under stones and dead leaves (mostly), under the loose bark of fallen trees and logs, under logs and pieces of wood. Gregarious. Larval habitat: under the loose bark of logs. Biology. Seasonality: January-October. Gravids: June (QU). Tenerals: mid-January (rarely; MA), late June to September. Adult food: lepidopterous immature stages (in the field); diplopods and mealworms (in captivity). Larval food, in captivity: mealworms. Predators: toads and crows. Parasites: tachinid larvae. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; baiting traps with meat; turning stones; raking dead

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leaves; night collecting with a headlamp. References. Blatchley, 1910 (ecology); Gilbert, 1957 (biology); Lindroth, 1966 (ecology, dispersal power); Erwin, 1981 (natural history); Bousquet, 1986b (biology); Will et al. 1995 (dispersal power); Chen and Willson, 1996 (biology); Bousquet, 1999 (taxonomy). Pterostichus (Gastrosticta) subacutus (Casey, 1918) Ecology. Lowlands. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Metallophilus) sublaevis (J.R. Sahlberg, 1880) Ecology. Lowlands and mountains. A mountainous tundra slope with dry, sand-mixed soil (AK). Biology. Seasonality: May-August. Tenerals: mid-August (AK). Dispersal power. Brachypterous, incapable of flight. References. Lindroth, 1966 (ecology, dispersal power); Bousquet, 1999 (taxonomy). Pterostichus (Hypherpes) suffusus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Eosteropus) superciliosus (Say, 1823) Ecology. Mountains. Deciduous and coniferous rainforests (e.g., Picea). Shaded ground; wet soil covered with dead leaves or moss. Nocturnal; sheltering during the day under stones, in rotten logs and leaf litter. Biology. Seasonality: May-September. Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning stones; dismembering rotten logs; pitfall trapping. References. Bousquet, 1986b (biology) and 1999 (taxonomy, biology). Pterostichus (Cryobius) surgens LeConte, 1878 Ecology. Mountains, subalpine and alpine zones. Alpine meadows and moraines (mostly); mountain forests. Open or shaded ground; moderately moist soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: April-September. Adult food, in captivity: mealworms, chrysomelids, and snails. Larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning stones. References. Ball, 1966 (ecology); Lindroth, 1966 (ecology, dispersal power); Schmoller, 1970 (biology). Pterostichus (Hypherpes) tahoensis Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight.

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Pterostichus (Cryobius) tareumiut Ball, 1962 Ecology. Lowlands. Tundra. Open ground; rather wet, peaty soil covered with some vegetation (e.g., Dupontia, Carex, Petasites, Eriophorum). Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. References. Ball, 1963 and 1966 (ecology); Lindroth, 1966 (ecology, dispersal power). Pterostichus (Hypherpes) tarsalis LeConte, 1873 Ecology. Mountains. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Pseudomaseus) tenuis (Casey, 1924) Ecology. Lowlands and mountains. Margins of bogs, swamps, eutrophic marshes, ponds, lakes, and slow rivers. Close to water. Open or slightly shaded ground; soft, wet soil covered with moss carpets (Sphagnum). Nocturnal; sheltering during the day in moss carpets (mostly), under dead leaves, fallen dead grass, and pieces of wood. Gregarious (in winter). Biology. Seasonality: March-November. Gravids: April (QU). Tenerals: August (NH, QU). Adults found overwintering on higher and drier ground, in woods and at their edges; under stones (mostly), logs, and in leaf litter. Adult and larval food, in captivity: mealworms. Parasites: braconid larvae. Dispersal power. Macropterous. Occasional flier (to artificial lights in captivity). Moderate runner. Collecting technique. Treading moss carpets into the water. References. Bousquet and Pilon, 1977 (biology); Davidson, 1988b (ecology). Pterostichus (Hypherpes) testaceus (Van Dyke, 1926) Ecology. Mountains. Coniferous rainforests. Shaded ground; wet soil, with a thick layer of dead leaves and humus. Nocturnal; sheltering during the day under well-embedded logs, in thick leaf litter, and humus. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting techniques. Turning embedded logs; pitfall trapping. Pterostichus (Cryobius) tiliaceoradix Ball, 1962, see Pterostichus (Cryobius) bryanti tiliaceoradix Ball, 1962. Reference. Ball and Currie, 1997 (taxonomy). Pterostichus (Bothriopterus) trinarius (Casey, 1918) (Pterostichus ohionis Csiki, 1930) Ecology. Lowlands. Deciduous forests (e.g., Quercus, Carya): along rivers, brooks, ravines, and slopes. Shaded ground; dry or moist, clay soil. Nocturnal; sheltering during the day under stones, logs, dead leaves, and boards. Biology. Seasonality: March-Sep-

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tember, November. Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Moderate runner. Collecting techniques. Turning stones and logs; pitfall trapping. References. Hamilton, 1884 (ecology; as ohionis Csiki); Kirk, 1975c (ecology; as ohionus [sic] Csiki); Erwin, 1981 (natural history; as ohionis Csiki); Bousquet, 1999 (dispersal power). Pterostichus (Hypherpes) trinitensis Hacker, 1968 Ecology. Lowlands and mountains. Coniferous forests. Moderately moist soil. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Pterostichus (Hypherpes) tristis (Dejean, 1828) Ecology. Lowlands and mountains. Deciduous (mostly; e.g., Acer), mixed (e.g., Abies, Betula), and coniferous forests (e.g., Picea, Abies, Thuja). Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under the loose bark of logs and fallen trees (mostly), under logs, fallen trees and pieces of wood, in rotten logs, in standing rotten tree-stumps, under stones and dead leaves. Biology. Seasonality: March-November. Copulating pairs: August-September. Tenerals: May to early September. Adult food: lepidopterous caterpillars (in the field); mealworms, snails, meat, and apple (in captivity). Larval food, in captivity: mealworms. Economic importance: plays an important role in the control of the Spruce Budworm. Predators: toads. Parasites: nemathelminth worms, ascid mites, and fungi (Laboulbeniales). Phoretic mites found on adult body. Defense-mechanism: when alarmed, the adult emits a strong smell, similar to burnt chocolate. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Frequent climber (on fallen trees). Collecting techniques. Lifting the loose bark of logs and fallen trees; pitfall trapping; using malaise traps. References. Erwin, 1981 (natural history); Reeves et al. 1983 (biology); Bousquet, 1986b (biology); Levesque and Levesque, 1986 (biology); Will et al. 1995 (dispersal power). Pterostichus (Hypherpes) tuberculofemoratus Hatch, 1936 Ecology. Mountains. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day in or under old logs. Biology. Seasonality: March, June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Inspecting old logs. Pterostichus (Gastrosticta) tumescens LeConte, 1863 Ecology. Lowlands. Forests: along rivers and lakes. Shaded or open ground; moist or wet soil. Nocturnal; sheltering during the day under pieces of wood and dead leaves. Biology. Seasonality: January, July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning pieces of wood; pitfall trapping.

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Pterostichus (Hypherpes) vandykei Schaeffer, 1910 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Gastrosticta) ventralis (Say, 1823) Ecology. Lowlands. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Cryobius) ventricosus ventricosus (Eschscholtz, 1823) Ecology. Lowlands. Mostly coniferous forests: along river banks. Also heaths and meadows (in the tundra). Shaded or open ground; more or less moist soil. Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: July-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. References. Ball, 1966 (ecology); Lindroth, 1966 (ecology, dispersal power). Pterostichus (Lenapterus) vermiculosus (Ménétriés, 1851) Ecology. Tundra. Open ground; moist, muddy soil, rich in organic debris, covered with continuous thin vegetation (e.g., Eriophorum, Carex). Biology. Seasonality: June-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Reference. Bousquet, 1999 (taxonomy). Pterostichus (Pseudoferonina) vexatus Bousquet, 1985 Ecology. Unknown. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Pterostichus (Hypherpes) vicinus Mannerheim, 1843 Ecology. Mountains. Forest ravines: along streams. Wet soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: February-August, October. Adult food, in the field: sowbugs (isopods). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning stones; pitfall trapping. Pterostichus (Cryobius) woodi Ball and Currie, 1997 Ecology. Dry rocky tundra. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Reference. Ball and Currie, 1997 (taxonomy, ecology). Pterostichus (Hypherpes) ybousqueti Berlov, 1999 (Pterostichus (Hypherpes) parens Casey, 1913) Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Reference. Berlov, 1999 (taxonomy).

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Pterostichus (Hypherpes) yosemitensis Hacker, 1968 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Reference. Bousquet, 1999 (taxonomy). Pterostichus (Hypherpes) zunianus Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Rhadine anthicoides Casey, 1913 Ecology. Uplands. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Rhadine albamontana Dajoz, 1998 Ecology. Mountains. Forests (Pinus). Shaded ground; dry rocky soil. Nocturnal; sheltering during the day in soil crevices. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Inspecting soil crevices. Reference. Dajoz, 1998 (taxonomy, ecology). Rhadine araizai babcocki (Barr, 1960) Ecology. Caves. Biology. Seasonality: April, August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine austinica Barr, 1974 Ecology. Caves. Biology. Seasonality: May. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine balesi (Gray, 1937) Ecology. A moist underground inlet of waterworks (WA). Biology. Seasonality: May. Dispersal power. Brachypterous, incapable of flight. Rhadine caudata (LeConte, 1863) Ecology. Lowlands and mountains. Deciduous forests (e.g., Quercus, Carya, Ostrya). Shaded ground; moderately moist soil. Caves (occasionally). Nocturnal; sheltering during the day under well-embedded stones (mostly; as deep as 20 cm), in soil crevices, and under debris. Biology. Seasonality: January-December. Tenerals: July and September (outside caves); all months (in caves). Adult food: cave-cricket eggs (in the field); meat (in captivity). Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Strong burrower. Collecting techniques. Turning embedded stones; pitfall trapping. References. Blatchley, 1910 (ecology); Erwin, 1981 (natural history). Rhadine constricta Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Rhadine dissecta (LeConte, 1863) Ecology. Uplands. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Rhadine exilis (Barr and Lawrence, 1960) Ecology. Caves. Climbing on ceilings and walls. Biology. Seasonality: April, July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Occasional climber. Rhadine gracilenta Casey, 1913 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Rhadine howdeni (Barr and Lawrence, 1960) Ecology. Caves: associated with bat guano. Biology. Seasonality: February-April, June, August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine infernalis ewersi (Barr, 1960) Ecology. Caves. Biology. Seasonality: April-May. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine infernalis infernalis (Barr and Lawrence, 1960) Ecology. Caves. Biology. Seasonality: June-July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine insolita Barr, 1974 Ecology. Caves. Biology. Seasonality: March. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine jejuna (LeConte, 1878) Ecology. Mountains. Damp situations surrounded by semi-arid country. Shrub-steppes. Open ground; damp soil covered with some vegetation (e.g., Artemisia, Chrysothamnus,

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Bromus). Nocturnal; sheltering during the day under stones. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting techniques. Turning stones; pitfall trapping. Rhadine koepkei koepkei (Barr, 1960) Ecology. Caves: along streams. Soil sometimes clayish. Climbing on walls or occurring on the ground. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Occasional climber (on cave walls). Rhadine koepkei privata Barr, 1974 Ecology. Caves. Biology. Seasonality: May. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine lanei (Gray, 1937) Ecology. Macrohabitat unknown. Under a pile of stones. Biology. Seasonality: February. Dispersal power. Brachypterous, incapable of flight. Rhadine larvalis LeConte, 1848 Ecology. Forests and caves: along river banks. Bare soil composed of clay or of clay and sand. Biology. Seasonality: June, October. Defense-mechanism: when alarmed, the adult escapes by running into soil crevices or along plant roots. Dispersal power. Brachypterous, incapable of flight. Fast runner. Strong burrower. Collecting technique. Inspecting soil crevices. Rhadine lindrothi Barr, 1965 Ecology. Uplands. Burrows of Ground squirrels; soil crevices along river banks. Cultivated fields (e.g., corn, oat); gravel pits. Nocturnal; sheltering during the day in Ground squirrel burrows, cracks in the soil, and under stones. Biology. Seasonality: May, JulySeptember. Dispersal power. Brachypterous, incapable of flight. Fast runner. Strong burrower. Collecting technique. Inspecting Ground squirrel burrows and soil crevices. References. Lindroth, 1966 (natural history); Bell, 1971 (ecology). Rhadine longiceps Van Dyke, 1949 Ecology. Caves and cellars. Sheltering under stones. Biology. Seasonality: November. Dispersal power. Brachypterous, incapable of flight. Fast runner. Collecting technique. Turning stones.

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Rhadine longicollis Benedict, 1927 Ecology. Caves (mostly in the dark zone; also in the twilight and entrance zones). Associated with bat guano. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine longipes Casey, 1913 Ecology. Uplands. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Rhadine myrmecodes (G.H. Horn, 1892) Ecology. Uplands. Ant nests. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine nivalis (G.H. Horn, 1881) Ecology. Uplands. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Rhadine noctivaga Barr, 1974 Ecology. Caves. Biology. Seasonality: January, March. Tenerals: March. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine ozarkensis Sanderson and Miller, 1941 Ecology. Caves. Biology. Seasonality: January, November. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine perlevis Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Rhadine persephone Barr, 1974 Ecology. Caves. Biology. Seasonality: May. Tenerals: May. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine pertenuis Casey, 1920 Ecology. Unknown. Biology. Seasonality: April-May, July. Dispersal power. Brachypterous, incapable of flight.

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Rhadine plumasensis Casey, 1920 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Rhadine rossi Van Dyke, 1949 Ecology. Burrows of Pocket gophers. Nocturnal; sheltering during the day in Pocket gopher burrows. Biology. Seasonality: November. Dispersal power. Brachypterous, incapable of flight. Fast runner. Strong burrower. Collecting technique. Inspecting Pocket gopher burrows. Rhadine rubra (Barr, 1960) Ecology. Rodent burrows; caves. Biology. Seasonality: August, October. Dispersal power. Brachypterous, incapable of flight. Fast runner. Strong burrower. Collecting technique. Inspecting rodent burrows. Rhadine russelli Barr, 1974 Ecology. Caves. Biology. Seasonality: May. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine specus crinicollis Barr, 1974 Ecology. Caves. Biology. Seasonality: March. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine specus gentilis Barr, 1974 Ecology. Caves: among stalagmites. Wet, silty soil. Biology. Seasonality: May. Tenerals: May. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine specus specus (Barr, 1960) Ecology. Caves: along stream banks. Loose, wet silty soil. Biology. Seasonality: March, August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine sublustris Casey, 1913 Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Rhadine subterranea mitchelli Barr, 1974 Ecology. Caves. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Rhadine subterranea subterranea (Van Dyke, 1918) Ecology. Caves. Rather dry silty soil. Biology. Seasonality: March, December. Adult food, in the field: cave-cricket eggs. Dispersal power. Brachypterous, incapable of flight. Fast runner. Reference. Mitchell, 1971a (biology), 1971b (dispersal), and 1971c (ecology). Rhadine tenebrosa mckenziei Barr, 1974 Ecology. Caves. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine tenebrosa tenebrosa (Barr, 1960) Ecology. Caves. Biology. Seasonality: March-April. Dispersal power. Brachypterous, incapable of flight. Fast runner. Rhadine tenuipes Casey, 1920 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Rhadine testacea Casey, 1920 Ecology. Uplands. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Rhadine umbra Casey, 1913 Ecology. Uplands. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Scaphinotus (Steniridia) aeneicollis (Beutenmüller, 1903) Ecology. Mountains (highest zones). Coniferous forests (e.g., Picea, Abies): along ravines and streams. Shaded ground; moist soil. Nocturnal; sheltering during the day under loose tree bark. Biology. Seasonality: May-October. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Lifting loose bark; pitfall trapping. References. Valentine, 1935 (ecology); Barr, 1969 (ecology). Scaphinotus (Steniridia) andrewsii amplicollis (Casey, 1920) Ecology. Mountains. Forests. Shaded ground; moderately moist soil. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping.

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Scaphinotus (Steniridia) andrewsii andrewsii (T.W. Harris, 1839) Ecology. Mountains. Deciduous and mixed forests: along slopes and streams. Shaded ground; moist soil. Nocturnal; sheltering during the day under logs, the loose bark of fallen trees, in rotten wood, and tree hollows. Gregarious (in autumn; up to 25 individuals under the loose bark of a fallen tree). Biology. Seasonality: May-June, August, October. Adult food: snails and berries (Smilax) (in the field); snails (in captivity). Predators: thrushes. Defense-mechanism: when alarmed, the adult emits a defensive fluid from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Regular climber (on tree trunks). Collecting techniques. Turning logs; lifting loose bark; pitfall trapping; night collecting with a headlamp; pitfall trapping. References. Valentine, 1935 (ecology); Wheeler et al. 1970 (biology). Scaphinotus (Steniridia) andrewsii barksdalei (Valentine, 1936) Ecology. Mountains. Forests. Shaded ground; moderately moist soil. Biology. Seasonality: June, October. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Scaphinotus (Steniridia) andrewsii darlingtoni (Valentine, 1935) Ecology. Mountains. Coniferous forests (e.g., Picea, Abies): along ridges and ravines. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under the loose bark of fallen trees. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Lifting loose bark; pitfall trapping. Reference. Valentine, 1935 (ecology). Scaphinotus (Steniridia) andrewsii germari (Chaudoir, 1861) Ecology. Mountains. Deciduous forests (e.g., Quercus, Acer, Ostrya): along slopes and streams. Shaded ground; moist soil. Caves (occasionally). Nocturnal; sheltering during the day under the loose bark of fallen trees and logs. Biology. Seasonality: June-July, September. Adult food, in the field: snails. Defense-mechanism: when alarmed, the adult may emit a defensive fluid from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Regular climber (on pieces of wood and cave walls). Collecting techniques. Lifting loose bark; turning logs; pitfall trapping. Reference. Valentine, 1935 (ecology). Scaphinotus (Steniridia) andrewsii montanus (Valentine, 1935) Ecology. Mountains. Deciduous and mixed forests: along ravines and streams. Shaded ground; moist, rocky soil. Nocturnal; sheltering during the day under loose bark. Biology. Seasonality: May-June, August-September. Defense-mechanism: when alarmed, the adult may emit a defensive fluid from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Lifting loose bark; pitfall trapping; baiting traps with molasses. Reference. Valentine, 1935 (ecology).

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Scaphinotus (Steniridia) andrewsii mutabilis (Casey, 1920) Ecology. Mountains. Deciduous forests (e.g., Quercus, Acer). Shaded ground; moderately moist soil covered with dead leaves, stones, and logs. Nocturnal; sheltering during the day in rotten logs. Gregarious (in winter). Biology. Seasonality: May-December. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Dismembering rotten logs; pitfall trapping. Scaphinotus (Steniridia) andrewsii nantahalae (Valentine, 1936) Ecology. Mountains. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day in old tree-stumps and logs (mostly), sometimes under moss carpets. Biology. Seasonality: May-June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Frequent climber (on tree-stumps and logs). Collecting techniques. Dismembering old tree-stumps and logs; pitfall trapping. Scaphinotus (Steniridia) andrewsii parvitarsalis (Valentine, 1935) Ecology. Mountains. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under the loose bark of tree-stumps and fallen trees. Biology. Seasonality: June-October. Defense-mechanism: when alarmed, the adult may emit a smoke with musty ammoniac odor from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Frequent climber (on tree-stumps and fallen trees). Collecting techniques. Lifting loose bark; pitfall trapping. Scaphinotus (Steniridia) andrewsii saludae (Valentine, 1936) Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Scaphinotus (Steniridia) andrewsii waldensius (Valentine, 1935) Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Scaphinotus (Neocychrus) angulatus (T.W. Harris, 1839) Ecology. Lowlands and mountains. Temperate rainforests (coniferous e.g., Pseudotsuga). Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under old logs and leaf litter (mostly), sometimes in rotten tree trunks. Biology. Seasonality: AprilNovember. Tenerals: mid-June (WA). Adult food, in the field: snails. Predators: spiders. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on tree trunks). Collecting techniques. Turning old logs; raking the leaf litter; pitfall trapping; baiting traps with honey and fruit. References. Exline and Hatch, 1934 (biology); Lindroth, 1961 (natural history); Weber and Kavanaugh, 1992 (ecology).

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Scaphinotus (Stenocantharus) angusticollis (Fischer von Waldheim, 1823) Ecology. Lowlands, mountains, and subalpine zone. Mostly forests (coniferous, mixed, deciduous): often along streams. Cultivated fields. Human habitations (occasionally). Shaded ground; moist soil. Nocturnal; sheltering during the day mostly in rotten logs and tree-stumps, under the loose bark and moss of dead standing and fallen trees, in hollow tree roots, also under logs and boards, among dead leaves and other debris. Gregarious (up to 50 individuals in or under a single log). Biology. Seasonality: MarchNovember. Tenerals: May-July, September. Adults found overwintering in rotten halfrotten tree-stumps and logs. Adult food: snails, slugs, earthworms, and spiders (in the field); slugs and juvenile salamanders (in captivity). Defense-mechanism: when alarmed, the adult may emit a malodorous musky defensive fluid from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Frequent climber (on trees, up to 20 m above ground; also on shrubs, tree-stumps, and logs). Moderate runner. Collecting techniques. Dismembering rotten logs and tree-stumps; pitfall trapping; baiting traps with sugar. References. Exline and Hatch, 1934 (biology); Spencer, 1942 (ecology); Van Dyke, 1944 (ecology, dispersal power); Lindroth, 1961 (natural history); Johnson et al. 1966 (biology); Kavanaugh, 1977 (ecology) and 1992 (natural history); Nelson, 1988 (biology); Ovaska and Smith, 1988 (biology). Scaphinotus (Neocychrus) behrensi (Roeschke, 1907) Ecology. Mountains. Temperate rainforests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day under old logs and leaf litter (mostly), in rotten tree trunks. Biology. Seasonality: May. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on tree trunks). Collecting techniques. Turning old logs; raking the leaf litter; pitfall trapping. Reference. Weber and Kavanaugh, 1992 (ecology). Scaphinotus (Nomaretus) bilobus (Say, 1823) Ecology. Lowlands, mountains, and subalpine zone. Coniferous (e.g., Picea, Abies), mixed forests (e.g., Alnus, Acer, Betula, Abies), and their edges, clearings, and roadsides: often along rivers, lakes, and the seashore. Gravel pits surrounded by spruce forests; Sphagnum-bogs and fens. Shaded (mostly) or open ground; moist or wet soil covered with thick leaf litter or moss. Nocturnal; sheltering during the day under dead leaves (mostly), logs, stones, or the loose bark of fallen trees. Larval habitat: in leaf litter. Biology. Seasonality: May-September. Tenerals: July (mostly), September. Adult food, in the field: snails. Adult and larval food, in captivity: snails and slugs. Predators: toads and shrews. Also eaten by insectivorous plants (Sarracenia). Phoretic mites found on adult body. Defense-mechanism: when alarmed, the adult stridulates or escapes by burrowing into the leaf litter. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Frequent climber. Good burrower. Collecting techniques. Raking the leaf litter; pitfall trapping; using malaise traps. References. Lindroth, 1961 (natural history); Larochelle, 1973j (ecology); Bousquet and Pilon, 1977 (biology).

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Scaphinotus (Brennus) bullatus Van Dyke, 1924 Ecology. Mountains. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Scaphinotus (Nomaretus) cavicollis (LeConte, 1859) Ecology. Lowlands. Coniferous forests (e.g., Pinus) and tree-covered stony knolls. Shaded ground; moderately moist soil. Biology. Seasonality: April-June, August-October, December. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Scaphinotus (Brennus) cordatus (LeConte, 1853) Ecology. Mountains. Forests. Biology. Seasonality: May-June, September, November. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Scaphinotus (Brennus) crenatus (Motschulsky, 1859) Ecology. Mountains. Forests: along stream banks. Nocturnal; sheltering during the day under stones and logs. Biology. Seasonality: February, August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning stones and logs; pitfall trapping. Scaphinotus (Brennus) cristatus (T.W. Harris, 1839) Ecology. Mountains. Coniferous forests (Pseudotsuga). Shaded ground; moderately moist soil. Biology. Seasonality: May-July, September-November. Tenerals: May-June (CA). Predators: shrews and Scaphinotus striatopunctatus. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. References. Tilden, 1955 (biology); Weber and Kavanaugh, 1992 (ecology). Scaphinotus (Maronetus) debilis alpinus (Beutenmüller, 1903) Ecology. Mountains. Forests. Shaded ground; moderately moist soil, rich in leaf litter and humus. Nocturnal; sheltering during the day under stones and the loose bark of trees (mostly), and also in leaf litter. Biology. Seasonality: May-June, August-September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on trees). Collecting techniques. Turning stones; lifting loose bark; pitfall trapping. Scaphinotus (Maronetus) debilis debilis (LeConte, 1853) Ecology. Mountains. Coniferous forests (e.g., Picea, Abies). Shaded ground; moderately moist soil rich in leaf litter and humus. Nocturnal; sheltering during the day in thick

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leaf litter (mostly), under logs, stones, and the loose bark of fallen trees. Larval habitat: under dead leaves. Biology. Seasonality: June-July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber. Collecting techniques. Raking or sifting the leaf litter; pitfall trapping. References. Roeschke, 1907 (ecology); Emden, 1942 (biology); Barr, 1969 (ecology). Scaphinotus (Scaphinotus) elevatus coloradensis Van Dyke, 1907 Ecology. Lowlands and mountains. Mostly forests (e.g., Populus, Picea): along stream beds. Sometimes in cultivated fields (e.g., corn) and prairies. Shaded or open ground; moist soil. Nocturnal; sheltering during the day under stones, logs, and other debris. Biology. Seasonality: February, May-August. Adults found overwintering in rotten logs. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning stones and logs; pitfall trapping. Reference. Potts, 1943 (biology). Scaphinotus (Scaphinotus) elevatus elevatus (Fabricius, 1787) Ecology. Lowlands. Mostly forests. Also cultivated fields, meadows, and clay pits. Shaded or open ground; moderately moist soil. Nocturnal; sheltering during the day under stones, fallen tree-stumps, logs, the loose bark of trees and tree-stumps, dead leaves, boards, in animal burrows, rotten logs and fallen trees, and scarabaeid (Xylorictes) borings at the base of trees. Gregarious in winter (up to 30 individuals in the same shelter). Larval habitat: in a pupation cell dug under a log. Biology. Seasonality: March-December. Tenerals: April-May. Adult food, in the field: snails, slugs, lepidopterous caterpillars and pupae, and maggots. Predators: crows, owls, and raccoons. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting techniques. Turning stones, fallen tree-stumps, and logs; pitfall trapping; sugaring trees. References. Wood, 1894 (biology); Wickham, 1895 (biology); Lindroth, 1961 (natural history); Graves and Graves, 1978 (ecology). Scaphinotus (Scaphinotus) elevatus flammeus Haldeman, 1844 Ecology. Lowlands. Deciduous forests: often along rivers. Shaded ground; moist soil. Nocturnal; sheltering during the day under dead leaves. Biology. Seasonality: AprilNovember. Tenerals: May-June (TN). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; pitfall trapping. Scaphinotus (Scaphinotus) elevatus lengi Van Dyke, 1938 Ecology. Lowlands. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Scaphinotus (Scaphinotus) elevatus neomexicanus Van Dyke, 1924 Ecology. Mountains. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight.

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Scaphinotus (Scaphinotus) elevatus tenebricosus Roeschke, 1907 Ecology. Lowlands and mountains. Open forests. Shaded ground; wet soil. Nocturnal; sheltering during the day under the loose bark of fallen trees. Biology. Seasonality: MayJune, October. Tenerals: May (VA). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber. Collecting techniques. Lifting loose bark; pitfall trapping. Reference. Valentine, 1935 (ecology). Scaphinotus (Nomaretus) fissicollis (LeConte, 1853) Ecology. Lowlands and mountains. Deciduous (e.g., Quercus, Carya) and coniferous forests (e.g., Pinus). Shaded ground; moderately moist soil. Biology. Seasonality: MarchMay, August-November. Tenerals: April (AR), May (IA). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Scaphinotus (Steniridia) guyotii (LeConte, 1866) Ecology. Mountains. Coniferous (e.g., Picea, Abies), mixed and deciduous forests: along slopes and ravines. Also mountain tops. Cool, shaded ground; moist soil. Nocturnal; sheltering during the day under the loose bark and pieces of wood. Biology. Seasonality: May-September. Adult food, in the field: snails. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber. Collecting techniques. Lifting loose bark; pitfall trapping. References. Valentine, 1935 (ecology); Barr, 1969 (biology). Scaphinotus (Stenocantharus) hatchii Beer, 1971 Ecology. Mountains. Coniferous forests (e.g., Tsuga). Shaded ground; moderately moist soil covered with sparse vegetation (e.g., Vaccinium). Nocturnal; sheltering during the day under old logs, the loose bark of fallen trees, and stones. Biology. Seasonality: JuneSeptember. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber. Collecting techniques. Turning old logs; pitfall trapping. Scaphinotus (Maronetus) hubbardi (Schwarz, 1895) Ecology. Mountains. Coniferous forests (e.g., Picea, Abies). Shaded ground; moderately moist soil covered with thick leaf litter and sticks. Nocturnal; sheltering during the day in leaf litter (mostly), under small sticks and the loose bark of fallen trees. Biology. Seasonality: May-June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber. Collecting techniques. Pitfall trapping; baiting traps with molasses mixed with a few drops of asafoetida; raking or sifting the leaf litter. References. Nicolay and Weiss, 1934 (ecology); Barr, 1969 (ecology).

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Scaphinotus (Maronetus) imperfectus (G.H. Horn, 1860) Ecology. Mountains. Deciduous forests, coniferous forests, and thickets (e.g., Rhododendron): along streams. Shaded ground; moist soil, rich in leaf litter and humus. Nocturnal; sheltering during the day in leaf litter (mostly), under stones and the loose bark of fallen trees. Biology. Seasonality: June-November. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber. Collecting techniques. Pitfall trapping; raking the leaf litter. Reference. Roeschke, 1907 (ecology). Scaphinotus (Maronetus) incompletus (Schwarz, 1895) Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Scaphinotus (Nomaretus) infletus Allen and Carlton, 1988 Ecology. Mountains. Biology. Seasonality: May-July. Dispersal power. Brachypterous, incapable of flight. Scaphinotus (Brennus) interruptus (Ménétriés, 1843) Ecology. Mountains. Deciduous forests (e.g., Quercus), coniferous forests (e.g., Pseudotsuga), river banks, open grassy knolls, gardens, and cultivated fields. Shaded or open ground; moderately moist soil covered with some vegetation. Nocturnal; sheltering during the day under the loose bark of trees, clods of earth, fallen branches, boards, and in logs. Gregarious. Biology. Seasonality: March-July, December. Tenerals: June and December (CA). Adult food: snails, slugs, and sowbugs (in the field); snails and slugs (in captivity). Economic importance: may play an important role in the control of the Garden and Green House slugs (Deroceras agreste (Linné), Milax gagetes (Müller)). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Frequent climber (on tree trunks and logs). Collecting techniques. Lifting the loose bark of trees; pitfall trapping; night collecting with a headlamp. References. Lange, 1941 (biology); Ingram, 1946 (biology); Kavanaugh, 1977 (ecology); Weber and Kavanaugh, 1992 (ecology). Scaphinotus (Brennus) johnsoni Van Dyke, 1924 Ecology. Mountains and subalpine zone. Forests (e.g., Tsuga). Shaded ground; moderately moist soil covered with dense vegetation (e.g., moss, Oxalis, Polystictum). Nocturnal; sheltering during the day under and in rotten logs. Gregarious. Biology. Seasonality: April-May, July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Inspecting old logs; pitfall trapping. References. Van Dyke, 1944 (ecology); Lindroth, 1961 (natural history).

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Scaphinotus (Scaphinotus) kelloggi (Dury, 1912) Ecology. Mountains. In a canyon: along river banks. Moist soil. Biology. Seasonality: July-August. Dispersal power. Brachypterous, incapable of flight. Scaphinotus (Nomaretus) liebecki Van Dyke, 1936 Ecology. Lowlands. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Scaphinotus (Steniridia) lodingi lodingi (Valentine, 1935) Ecology. Lowlands. Forests (mostly). Caves (occasionally). Shaded ground; moderately moist soil. Biology. Seasonality: July-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on cave walls). Collecting technique. Pitfall trapping. Scaphinotus (Steniridia) lodingi obscurus (Valentine, 1935) Ecology. Lowlands. Forests. Shaded ground; wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Scaphinotus (Neocychrus) longiceps Van Dyke, 1924 Ecology. Mountains. Temperate rainforests (coniferous, e.g., Sequoia, Pseudotsuga). Shaded ground; moderately moist soil. Nocturnal; sheltering under old logs and in leaf litter (mostly), and in rotten tree trunks. Biology. Seasonality: May, July, October-December. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on tree trunks). Collecting techniques. Turning logs; raking the leaf litter; pitfall trapping. Reference. Weber and Kavanaugh, 1992 (ecology). Scaphinotus (Pseudonomaretus) mannii Wickham, 1919 Ecology. Lowlands. Fragments of forests and brushes: often along rivers and canyons. Shaded ground; moist soil. Nocturnal; sheltering during the day in cavities dug under stones. Gregarious (in small groups under large stones). Larval habitat: similar to adult, in burrows dug in the soil. Biology. Seasonality: March-July, October-November. Copulating pairs: June (WA). Tenerals: March-May (WA). Adults found overwintering in soil crevices. Adult food, in captivity: slugs, crushed snails, and earthworms. Larval food, in captivity: snails. Predators: spiders. Defense-mechanism: when alarmed, the adult stridulates or emits an acrid secretion from the pygidial glands, at a range of up to 30 cm. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting techniques. Turning stones; pitfall trapping. Reference. Greene, 1975 (ecology, biology).

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Scaphinotus (Brennus) marginatus (Fischer von Waldheim, 1820) Ecology. Lowlands, mountains, subalpine and alpine zones. Coniferous (e.g., Pseudotsuga, Pinus), mixed, and deciduous forests: often along streams, lakes, the seashore, and canyons. Forest clearings, fragments of forests and brushes, and cultivated fields. Vicinity of human habitations (urban and rural sites; e.g., gardens). Supratidal meadows, open coastal toundra (Aleutian Islands). Vicinity of snow-patches (above timber line). Shaded or open ground; moist soil. Nocturnal; sheltering during the day in rotten logs and tree-stumps, under stones, under the loose bark of logs, in hollow tree roots, in leaf litter and moss. Gregarious (up to 50 individuals in the same shelter). Larval habitat: under stones. Biology. Seasonality: March-November. Tenerals: March to early June (BC, WA), mostly May (WA); also late July (AK). Adults found overwintering in logs, halfrotten logs and tree-stumps, and in crevices in the soil. Adult food: snails and slugs (in the field); slugs, snails, dipterous eggs, mealworms, and earthworms (in captivity). Larval food, in captivity: snails and mealworms. Predators: shrews and spiders. Mites infesting the adult body. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Excellent climber (on logs and tree-stumps). Collecting techniques. Dismembering rotten logs and tree-stumps; pitfall trapping; light trapping. References. Exline and Hatch, 1934 (biology); Lindroth, 1961 (natural history); Greene, 1975 (ecology, biology); Kavanaugh, 1977 (ecology) and 1992 (natural history); Nelson, 1988 (biology); Niemelä and Spence, 1991 (ecology); Niemelä et al. 1992 (ecology, biology); Digweed, 1994 (biology). Scaphinotus (Pseudonomaretus) merkelii (G.H. Horn, 1890) Ecology. Mountains. Mixed forests: along brooks. Shaded ground; rather dry soil. Nocturnal; sheltering during the day. Biology. Seasonality: April, June-August. Tenerals: late June (ID), late July (BC). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Lindroth, 1961 (natural history). Scaphinotus (Brennus) obliquus (LeConte, 1868) Ecology. Mountains. Biology. Seasonality: May-June. Predators: robins. Dispersal power. Brachypterous, incapable of flight. Scaphinotus (Brennus) oreophilus (Rivers, 1890) Ecology. Mountains. Biology. Seasonality: May, September, November. Dispersal power. Brachypterous, incapable of flight. Scaphinotus (Scaphinotus) parisiana Allen and Carlton, 1988 Ecology. Mountains. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight.

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Scaphinotus (Scaphinotus) petersi biedermani Roeschke, 1907 Ecology. Mountains. Coniferous forests: along slopes. Shaded ground; moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: July-September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Scaphinotus (Scaphinotus) petersi catalinae Van Dyke, 1924 Ecology. Mountains. Coniferous forests. Shaded ground; moist soil. Sheltering during the day. Biology. Seasonality: June-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Scaphinotus (Scaphinotus) petersi corvus (Fall, 1910) Ecology. Mountains. Coniferous forests: along brooks. Shaded ground; moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: June-September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Scaphinotus (Scaphinotus) petersi grahami Van Dyke, 1938 Ecology. Mountains. Coniferous and mixed forests. Shaded ground; moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: June-September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Scaphinotus (Scaphinotus) petersi kathleenae Ball, 1966 Ecology. Mountains. A mixed forest, on shaded ground with moist, gravelly-loamy soil. Nocturnal; sheltering during the day under logs, small pieces of wood, and in leaf litter. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Scaphinotus (Scaphinotus) petersi petersi Roeschke, 1907 Ecology. Mountains. Coniferous forests. Shaded ground; moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: June-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Scaphinotus (Brennus) punctatus (LeConte, 1859) Ecology. Mostly forests: along lakes and streams. Also dry fields. Shaded (mostly) or open ground; moist or dry soil. Nocturnal; sheltering during the day under stones, logs, and dead leaves. Gregarious. Biology. Seasonality: January-April, June-July, September-October, December. Tenerals: early June. Adult food, in the field: snails (mostly), live and dead lepidopterous caterpillars, a carabid (Platynus brunneomarginatus), meat, and a berry. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning stones and logs; pitfall trapping; trapping with

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meat placed under stones, chips, and dead leaves. Reference. Noonan, 1967b (ecology, biology). Scaphinotus (Pseudonomaretus) regularis (LeConte, 1884) Ecology. Mountains. Various habitats, e.g., forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day. Larval habitat: under logs. Biology. Seasonality: April-August, October-November. Tenerals: late spring to early summer (WA). Adults found overwintering in soil crevices. Adult food, in captivity: slugs and earthworms. Larval food, in captivity: crushed snails. Predators, in captivity: spiders and shrews. Defense-mechanism: when alarmed, the adult stridulates. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Greene, 1975 (ecology, biology). Scaphinotus (Pseudonomaretus) relictus (G.H. Horn, 1881) Ecology. Mountains. Forests (deciduous, mixed, and coniferous). Gardens (occasionally). Shaded ground; moderately moist soil, with thick humus layer. Nocturnal; sheltering during the day under stones and fallen trees. Larval habitat: under stones. Biology. Seasonality: April-November. Copulating pairs: August-September (WA). Tenerals: May-July (BC, ID, WA); mostly in late June (WA). Adult food, in captivity: slugs and earthworms. Larval food, in captivity: crushed snails. Mites infesting the adult body. Defense-mechanism: when alarmed, the adult stridulates. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning stones and fallen trees; pitfall trapping. References. Lindroth, 1961 (natural history); Greene, 1975 (ecology, biology). Scaphinotus (Steniridia) ridingsii intermedius (Valentine, 1935) Ecology. Mountains. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Scaphinotus (Steniridia) ridingsii monongahelae Leng, 1917 Ecology. Mountains. Coniferous forests (e.g., Picea): often along gorges. Shaded ground; moderately moist, stony soil. Nocturnal; sheltering during the day. Biology. Seasonality: May-September. Tenerals: mid-July (PA). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on tree trunks). Collecting technique. Pitfall trapping. Scaphinotus (Steniridia) ridingsii ridingsii (Bland, 1863) Ecology. Mountains. Forests: along streams. Shaded ground; moist, rocky soil. Nocturnal; sheltering during the day in thick leaf litter and under stones. Biology. Seasonality: JuneSeptember. Tenerals: early July (VA). Adult food, in the field: snails. Dispersal power.

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Brachypterous, incapable of flight. Moderate runner. Frequent climber (on stones and boulders). Collecting techniques. Raking the leaf litter; pitfall trapping; baiting traps with molasses. References. Valentine, 1935 (ecology); Erwin, 1981 (natural history). Scaphinotus (Brennus) riversi (Roeschke, 1907) Ecology. Mountains. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: October. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Scaphinotus (Brennus) rugiceps incipiens (Casey, 1897) Ecology. Mountains. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Scaphinotus (Brennus) rugiceps rugiceps (G.H. Horn, 1872) Ecology. Mountains. Coniferous forests (e.g., Pinus). Nocturnal; sheltering during the day. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Scaphinotus (Maronetus) schwarzi (Beutenmüller, 1913) Ecology. Mountains. Forests. Shaded ground; moderately moist soil, rich in leaf litter, humus, and moss. Nocturnal; sheltering during the day in leaf litter, humus, and under large stones well embedded in moss. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting techniques. Pitfall trapping; raking leaf litter and humus. Scaphinotus (Scaphinotus) snowi roeschkei Van Dyke, 1907 Ecology. Mountains. Biology. Seasonality: June-August. Dispersal power. Brachypterous, incapable of flight. Scaphinotus (Scaphinotus) snowi snowi (LeConte, 1881) Ecology. Mountains. Coniferous forests: along brooks and springs. Shaded ground; moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: May-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Scaphinotus (Brennus) striatopunctatus (Chaudoir, 1844) Ecology. Lowlands and mountains. Mostly deciduous forests (e.g., Quercus). Also canyons and grassy knolls. Shaded or open ground; moderately moist, sandy soil. Nocturnal;

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sheltering during the day in old logs and under the loose bark of fallen trees. Gregarious. Biology. Seasonality: February, April, June, November. Adult food, in the field: a Scaphinotus cristatus. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Frequent climber (on logs and fallen trees). Collecting techniques. Dismembering old logs; pitfall trapping. References. Tilden, 1955 (biology); Kavanaugh, 1977 (ecology). Scaphinotus (Brennus) subtilis (Schaum, 1863) Ecology. Lowlands. Vicinity of water bodies. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: January-February. Dispersal power. Brachypterous, incapable of flight. Scaphinotus (Maronetus) tenuis (Casey, 1914) Ecology. Mountains. Forests. Shaded ground; moderately moist soil, rich in leaf litter and humus. Nocturnal; sheltering during the day in leaf litter and humus. Biology. Seasonality: June-July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking leaf litter and humus. Scaphinotus (Steniridia) tricarinatus (Casey, 1914) Ecology. Alpine zone. Coniferous forests (e.g., Picea, Abies): often along ravines and streams. Shaded ground; moist soil. Nocturnal; sheltering during the day under moss carpets. Biology. Seasonality: June-September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on tree trunks). Collecting techniques. Turning moss carpets; pitfall trapping. References. Valentine, 1935 (ecology); Barr, 1969 (ecology). Scaphinotus (Scaphinotus) unicolor floridanus Leng, 1915 Ecology. Lowlands. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: October. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Scaphinotus (Scaphinotus) unicolor heros T.W. Harris, 1839 Ecology. Mountains. Forests: along rivers and ravines. Shaded ground; moist soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: May-July, September-November. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning stones; pitfall trapping. Scaphinotus (Scaphinotus) unicolor shoemakeri Leng, 1914 Ecology. Mountains. Forests: often along streams and springs. Shaded ground; wet, rocky soil. Nocturnal; sheltering during the day under stones, logs, and dead leaves. Biology. Seasonality: June, August-October. Tenerals: June (MD). Dispersal power. Brachypter-

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ous, incapable of flight. Moderate runner. Collecting techniques. Turning stones and logs; pitfall trapping; baiting bottles. Reference. Erwin, 1981 (natural history; as unicolor (Fabricius). Scaphinotus (Scaphinotus) unicolor unicolor (Fabricius, 1787) Ecology. Lowlands. Forests: along rivers. Shaded ground; wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: March-July, September. Adult food, in the field: scarabaeid larvae. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Löding, 1945 (ecology). Scaphinotus (Maronetus) unistriatus (Darlington, 1931) Ecology. Mountains. Mixed forests: often along slopes. Shaded ground; moderately moist soil, rich in leaf litter and humus. Nocturnal; sheltering during the day in leaf litter and humus. Biology. Seasonality: March-April, June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; raking leaf litter and humus. Scaphinotus (Scaphinotus) vandykei Roeschke, 1907 Ecology. Mountains. Coniferous forests: often along water bodies. Wet soil. Nocturnal; sheltering during the day. Biology. Seasonality: September-October. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping. Scaphinotus (Stenocantharus) velutinus (Ménétriés, 1843) Ecology. Lowlands and mountains. Mature coniferous forests (Sequoia). Biology. Seasonality: March-October. Dispersal power. Brachypterous, incapable of flight. Reference. Van Dyke, 1925 (ecology). Scaphinotus (Brennus) ventricosus (Dejean, 1831) Ecology. Mostly deciduous forests (e.g., Quercus) and poison-oak thickets. Also open grassy knolls. Shaded or open ground; moderately moist soil. Nocturnal; sheltering during the day in logs and under their loose bark. Gregarious. Biology. Seasonality: February-April, June-July, September, November-December. Tenerals: late March (CA). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on logs). Collecting techniques. Turning logs; lifting loose bark; pitfall trapping. Reference. Kavanaugh, 1977 (ecology). Scaphinotus (Scaphinotus) viduus (Dejean, 1826) Ecology. Forests (mostly deciduous, e.g., Acer; also mixed and coniferous); adjoining fields. Usually close to water. Shaded ground; wet soil, which thick leaf litter. Noctur-

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nal; sheltering during the day under stones, dead leaves, the loose bark of fallen trees and logs, under fallen trees, pieces of wood, and moss carpets. Apparently more active at the surface of the ground in rainy weather. Larval habitat (in winter): under logs. Gregarious (in winter). Biology. Seasonality: January-December; mostly late July to midSeptember (PA). Tenerals: July and September (QU). Adults found overwintering under the loose bark of trees, and in small cells dug under logs, fallen tree-stumps, and pieces of wood. Adult food: snails (mostly), slugs, and lepidopterous caterpillars (in the field); snails, molasses (in captivity). Larval food, in the field: snails. Defense-mechanism: when alarmed, the adult produces a strong stridulation (similar to the noise produced by a rattlesnake or to the rustling of dry leaves) or emits a strong smoky defensive gas or fluid from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning stones; pitfall trapping; sugaring trees; baiting traps with molasses mixed with asafoetida; pinning live snails under boards; light trapping. References. Klages, 1898 (biology); Joutel, 1904 (biology); Lindroth, 1961 (natural history); Barr, 1969 (ecology; as irregularis (Beutenmüller)); Wheeler et al. 1970 (biology). Scaphinotus (Steniridia) violaceus carolinae (Valentine, 1935) Ecology. Mountains. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Scaphinotus (Steniridia) violaceus violaceus (LeConte, 1863) Ecology. Mountains. Deciduous and mixed forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day in rotten tree-stumps (mostly; near their base) and under the loose bark of fallen trees. Biology. Seasonality: May, July, September-November. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on tree-stumps). Collecting techniques. Dismembering rotten tree-stumps; pitfall trapping; baiting traps with molasses. Reference. Valentine, 1935 (ecology). Scaphinotus (Scaphinotus) webbi Bell, 1959 Ecology. Mountains. Deciduous forests (e.g., Quercus): often along brooks. Shaded ground; moist soil. Nocturnal; sheltering during the day under large stones. Biology. Seasonality: June-July. Tenerals: July (VA). Adult food, in the field and in captivity: snails. Defense-mechanism: when alarmed, the adult may emit a defensive fluid from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning large stones; pitfall trapping. Reference. Wheeler et al. 1970 (biology). Scarites (Scarites) lissopterus Chaudoir, 1880 Ecology. Lowlands and mountains. Biology. Seasonality: March, May-June, October. Dispersal power. Unknown.

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Scarites (Scarites) marinus Nichols, 1986 Ecology. Coastal lowlands. Sea beaches: intertidal area. Open ground; wet, sandy soil covered with washed up seaweed. Nocturnal; sheltering during the day under rotten seaweed. Biology. Seasonality: December and March (USA); throughout the year (Mexico). Tenerals: January (Mexico). Dispersal power. Brachypterous, incapable of flight. Slow runner. Strong burrower. Collecting technique. Turning rotten seaweed. References. Nichols, 1986 (ecology) and 1988 (ecology, dispersal power). Scarites (Scarites) ocalensis Nichols, 1986 Ecology. Lowlands. Biology. Seasonality: May. Dispersal power. Brachypterous, incapable of flight. Scarites (Scarites) patruelis LeConte, 1845 Ecology. Macrohabitat unknown. Nocturnal; sheltering during the day under pieces of wood. Biology. Seasonality: April, June. Dispersal power. Brachypterous, incapable of flight. Scarites (Scarites) quadriceps Chaudoir, 1843 (Scarites substriatus Haldeman, 1844) Ecology. Lowlands. Cultivated fields (e.g., corn, wheat, hay, soybean, alfalfa, flax, oat, asparagus), pastures, prairies, wastelands, fencerows, and pond borders; also forests and their ravines. Open ground; moderately moist, sandy soil. Nocturnal; sheltering during the day in burrows dug in the soil, under logs, boards, and other debris. Solitary. Biology. Seasonality: January-November. Copulating pairs: June (MI); coitus lasting 5-8 minutes. Adult food: lepidopterous caterpillars and scarabeids (in the field); weevils, live and dead invertebrates (e.g., lepidopterous caterpillars) (in captivity). Reared on larvae from the Southern Cornstalk Borer. Intraspecific aggressivity is well developped in captivity. Predators: alligators, raccoons, crows, ducks, and woodpeckers. Defense-mechanism: when alarmed, the adult feigns death (tricking in its antennae and legs and raising the pronotum upward, in a comatose position for up to 5 minutes). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Slow runner. Strong burrower. Strongly favored by human activities. Collecting techniques. Pitfall trapping; light trapping; turning logs and pieces of wood. References. Leng, 1915 (ecology, dispersal power; as substriatus Haldeman); Hlavac, 1967 (biology; as substriatus Haldeman); Kirk, 1969 (biology, dispersal power; as substriatus Haldeman) and 1975c (ecology); as substriatus Haldeman); Best et al. 1981 (biology, dispersal power); Nichols, 1988 (ecology, dispersal power); Will et al. 1995 (dispersal power). Scarites (Scarites) subterraneus Fabricius, 1775 Ecology. Lowlands and mountains. Borders of rivers, lakes, ponds, and irrigation canals; roadsides, gardens, lawns, urban and rural parks, vacant lots, old fields, wastelands, pastures, cultivated fields (e.g., corn, soybean, peanut, alfalfa, cabbage, cotton, tobacco), field

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edges, seed beds, orchards (e.g., citrus, apple), forest edges, and open forests. Open ground; moderately moist, sandy, clayish or muddy, rarely gravelly soil covered with some vegetation. Nocturnal; sheltering during the day in burrows dug in the soil (mostly; as deep as 15 cm), under stones, logs, boards, dead leaves, and fallen dead grass; also in the burrows of the Southern Dung Beetle (Copris carolina Linné). Solitary. Larval habitat: in burrows dug in the soil. Biology. January-December. Eggs are laid singly on the leaves of trees and shrubs; they are encased in a covering of clay and mud. Tenerals: April (rarely; FL), JulyAugust, October. Adults found overwintering in woods; under well-embedded stones and the loose bark of fallen trees. Adult food: lepidopterous caterpillars (frequently), cetoniid larvae, coccinellids, elaterids, other insects, earthworms, an injured salamander, and young seedlings in seed beds (in the field); lepidopterous caterpillars, ichneumonid larvae, and mealworms (in captivity). Adults are cannibalistic in captivity. Larval food, in captivity: mealworms. Economic importance: plays an important role in the control of lepidopterous caterpillars; often a pest of young seedlings. Predators: toads, frogs, alligators, blackbirds, bluebirds, cardinals, crows, ducks, woodpeckers, grackles, lapwings, meadowlarks, quails, robins, sparrows, starlings, thrushes, turkeys, and spiders. Parasites: mites (paratennulid, paramegistid, pyemotoid) and sarcophagid flies. Defense-mechanism: when alarmed, the adult feigns death (tricking in its antennae and legs and raising the pronotum upward, in a comatose position for up to 5 minutes). Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in drift material, which indicates flight ability. Slow runner. Good climber. Strong burrower. Strongly favored by human activities. Collecting techniques. Pitfall trapping; light trapping; turning stones and logs. References. Riley, 1886 (biology); Blatchley, 1910 (biology); Leng, 1915 (ecology, dispersal power); Lindroth, 1961 (natural history); Hlavac, 1967 (ecology, biology); Kirk, 1969 (natural history); Erwin, 1981 (natural history); Lesiewicz et al. 1983 (ecology, biology); Ferguson and McPherson, 1985 (biology); Nichols, 1988 (ecology, dispersal power); Will et al. 1995 (ecology, dispersal power). Scarites (Scarites) texanus Chaudoir, 1880 Natural History. Unknown. Schizogenius (Schizogenius) amphibius (Haldeman, 1843) Ecology. Lowlands and mountains. Banks of rivers and brooks. Open ground; wet, gravelly, sand-mixed, bare soil. Nocturnal; sheltering during the day among gravel (mostly), under stones and debris. Biology. Seasonality: May-September. Tenerals: late May (VT). Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting techniques. Raking the loose gravel; pouring the water over the ground. Reference. Whitehead, 1972 (natural history). Schizogenius (Schizogenius) auripennis Bates, 1881 Ecology. Stream banks. Open ground; gravelly, bare soil. Nocturnal; sheltering during the day among gravel. Biology. Seasonality: January-December. Dispersal power. Mac-

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ropterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. Collecting techniques. Raking the loose gravel; pouring water over the ground. Reference. Whitehead, 1972 (natural history). Schizogenius (Schizogenius) brevisetosus Whitehead, 1972 Ecology. Stream banks. Open ground; gravelly, bare soil. Nocturnal; sheltering during the day among gravel. Biology. Seasonality: April-August. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Slow runner. Strong burrower. Collecting techniques. Raking the loose gravel; pouring water over the ground; light trapping. Reference. Whitehead, 1972 (natural history). Schizogenius (Schizogenius) chiricahuanus Whitehead, 1972 Ecology. Mountains. Stream banks. Open ground; gravelly, bare soil. Nocturnal; sheltering during the day among gravel. Biology. Seasonality: June-August. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting techniques. Raking the loose gravel; pouring water over the ground. Reference. Whitehead, 1972 (natural history). Schizogenius (Genioschizus) crenulatus crenulatus LeConte, 1852 Ecology. River banks. Open ground; bare soil. Nocturnal; sheltering during the day in burrows dug in the soil. Biology. January-December. Dispersal power. Macropterous. Occasional flier (to artificial artificial lights at night). Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Whitehead, 1972 (possible extinction in southwestern USA). Schizogenius (Schizogenius) depressus LeConte, 1852 Ecology. Lowlands and mountains. Stream banks and lake shores. Close to water. Open ground; gravelly, bare soil. Nocturnal; sheltering during the day among gravel. Biology. Seasonality: January-December. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting techniques. Raking the loose gravel; pouring water over the ground. References. Lindroth, 1961 (natural history); Whitehead, 1972 (natural history). Schizogenius (Schizogenius) falli Whitehead, 1972 Ecology. Lowlands and mountains. Stream banks. Open ground; gravelly, bare soil. Nocturnal; sheltering during the day among gravel. Biology. Seasonality: April-October. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Slow runner. Strong burrower. Collecting techniques. Raking the loose gravel; pouring water over the ground; light trapping. Reference. Whitehead, 1972 (natural history).

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Schizogenius (Schizogenius) ferrugineus Putzeys, 1846 Ecology. Lowlands and mountains. Banks of rivers and brooks, lake shores, and sea beaches. At some distance from water or close to it. Open ground; moist, sandy (mostly) or gravelly, bare soil. Nocturnal; sheltering during the day in burrows dug in the soil under sticks, boards, stones, and logs. Biology. Seasonality: March-October. Tenerals: August-September (MD). Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Regular flier (to artificial lights at night). Occasional in drift material, which indicates flight ability. Slow runner. Strong burrower. Collecting techniques. Raking the sand; pouring water over the ground; light trapping. References. Schwarz, 1876 (ecology); Lindroth, 1961 (natural history); Whitehead, 1972 (ecology); Kirk, 1975c (dispersal power); Erwin, 1981 (natural history). Schizogenius (Schizogenius) lindrothi Whitehead, 1972 Ecology. Lowlands. Margins of a small spring-fed pond, on open ground with sandy, bare soil. Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Reference. Whitehead, 1972 (ecology). Schizogenius (Schizogenius) lineolatus (Say, 1823) Ecology. Lowlands and mountains. Bars and banks: along rivers and brooks. Close to water. Open ground; wet, bare, gravelly soil, sometimes mixed with clay or sand. Nocturnal; sheltering during the day mostly in burrows dug under gravel and in soil crevices, also under stones and vegetal debris. Gregarious (up to 10 individuals in the same shelter). Biology. Seasonality: April-October. Gravids: early June (QU). Tenerals: JulySeptember. Predators: frogs and toads. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Occasional flier (at dusk; to artificial lights at night; to artifical lights in captivity). Slow runner. Strong burrower. Collecting techniques. Raking the loose gravel; pouring water over the ground. References. Lindroth, 1961 (natural history); Whitehead, 1972 (natural history); Kirk, 1975c (dispersal power); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Schizogenius (Schizogenius) litigiosus Fall, 1901 Ecology. Lowlands and mountains. Stream banks. Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: January-December. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pouring water over the ground. Reference. Whitehead, 1972 (natural history). Schizogenius (Schizogenius) longipennis Putzeys, 1866 Ecology. Mountains. Stream banks. Open ground; gravelly, bare soil. Nocturnal; sheltering during the day among gravel. Biology. Seasonality: January-December (USA-Mexico). Ten-

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erals: June-August (Mexico). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. Collecting techniques. Raking the loose gravel; pouring water over the ground. Reference. Whitehead, 1972 (natural history). Schizogenius (Schizogenius) neovalidus Whitehead, 1972 Ecology. Stream banks. Open ground; gravelly, bare soil. Nocturnal; sheltering during the day among gravel. Biology. Seasonality: May-August. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting techniques. Raking the loose gravel; pouring water over the ground. Reference. Whitehead, 1972 (natural history). Schizogenius (Schizogenius) ochthocephalus Whitehead, 1972 Ecology. Mountains. Biology. Seasonality: January, March-April, June. Dispersal power. Macropterous, probably capable of flight. Schizogenius (Schizogenius) ozarkensis Whitehead, 1972 Ecology. Mountains. Stream banks. Open ground; gravelly, bare soil. Nocturnal; sheltering during the day among gravel. Biology. Seasonality: June-September. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting techniques. Raking the loose gravel; pouring water over the ground. Reference. Whitehead, 1972 (natural history). Schizogenius (Schizogenius) planulatus LeConte, 1863 Ecology. Lowlands and mountains. A stream bank, on open ground with gravelly, bare soil. Nocturnal; sheltering during the day among gravel. Biology. Seasonality: MaySeptember. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting techniques. Raking the loose gravel; pouring water over the ground. Reference. Whitehead, 1972 (natural history). Schizogenius (Schizogenius) planuloides Whitehead, 1972 Ecology. A stream bank, on open ground with gravelly bare soil. Nocturnal; sheltering during the day among gravel. Biology. Seasonality: March-August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Reference. Whitehead, 1972 (natural history). Schizogenius (Schizogenius) pluripunctatus LeConte, 1852 Ecology. Stream banks. At some distance from water. Open ground; moderately moist, gravelly, bare soil. Biology. Seasonality: March-September. Dispersal power. Macropterous. Regular flier (to artificial lights at night). Slow runner. Strong burrower. Collecting

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techniques. Raking the loose gravel; pouring water over the ground; light trapping. Reference. Whitehead, 1972 (natural history). Schizogenius (Schizogenius) pygmaeus Van Dyke, 1925 Ecology. Lowlands and mountains. Stream banks and lake shores. Open ground; gravelly, bare soil. Nocturnal; sheltering during the day among gravel. Biology. Seasonality: January-December. Adults observed overwintering. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. Collecting techniques. Raking the loose gravel; pouring water over the ground. References. Van Dyke, 1925 (ecology); Whitehead, 1972 (natural history). Schizogenius (Schizogenius) sallei Putzeys, 1866 Ecology. Banks of rivers and brooks. Open ground; gravelly, bare soil. Nocturnal; sheltering during the day among gravel. Biology. Seasonality: March-September. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. Collecting techniques. Raking the loose gravel; pouring water over the ground. Reference. Whitehead, 1972 (natural history). Schizogenius (Schizogenius) scopaeus Whitehead, 1972 Ecology. Mountains. Banks of brooks and rivers. Open ground; gravelly, bare soil. Nocturnal; sheltering during the day in burrows dug under gravel. Nocturnal; sheltering during the day among gravel. Biology. Seasonality: April-October. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Slow runner. Strong burrower. Collecting techniques. Raking the loose gravel; pouring water over the ground; light trapping. Reference. Whitehead, 1972 (natural history). Schizogenius (Schizogenius) seticollis seticollis Fall, 1901 Ecology. Mountains. Stream banks. Open ground; gravelly, bare soil. Nocturnal; sheltering during the day among gravel. Biology. Seasonality: March-October. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. Collecting techniques. Raking the loose gravel; pouring water over the ground. Reference. Whitehead, 1972 (natural history). Schizogenius (Schizogenius) sulcifrons Putzeys, 1846 Ecology. Lowlands and mountains. Banks of rivers and brooks. Open ground; wet, gravelly, bare soil. Nocturnal; sheltering during the day in burrows dug under gravel, stones, and logs. Semi-gregarious. Biology. Seasonality: March-October. Tenerals: August-September. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. Collecting techniques. Raking the loose gravel; pouring water over the ground. References. Lindroth, 1961 (natural history); Whitehead, 1972 (natural history).

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Schizogenius (Schizogenius) tibialis Whitehead, 1972 Ecology. A deep cave, along a brook. Biology. Seasonality: January-December (USAMexico). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Reference. Whitehead, 1972 (natural history). Selenophorus (Celiamorphus) adjunctus (Casey, 1914) Natural History. Unknown. Selenophorus (Selenophorus) aeneopiceus Casey, 1884 Natural History. Unknown. Selenophorus (Selenophorus) blanchardi Manee, 1915 Natural History. Unknown. Selenophorus breviusculus G.H. Horn, 1880 Ecology. Lowlands. Vacant lots, cultivated fields, and roadsides. Open ground; dry, sandy soil covered with sparse or moderate vegetation. Nocturnal; sheltering during the day in burrows dug in the soil at the base of plants (mostly), rarely under dead leaves and stones. Biology. Seasonality: February, July. Tenerals: late July (AL, TX). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Strong burrower. Collecting techniques. Digging at the base of plants; pitfall trapping. Selenophorus chaparralus Purrington, 2000 Ecology. Uplands. A dry grassy acacia savanna. Nocturnal; sheltering during the day under a small stone. Biology. Seasonality: March, October. Dispersal power. Unknown. Reference. Purrington, 2000 (taxonomy, ecology). Selenophorus (Selenophorus) concinnus Schaeffer, 1910 Natural History. Unknown. Selenophorus (Celiamorphus) contractus (Casey, 1914) Ecology. Unknown. Biology. Seasonality: March-April. Dispersal power. Unknown. Selenophorus (Selenophorus) cupreolus Casey, 1914 Natural History. Unknown.

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Selenophorus (Selenophorus) depressulus (Casey, 1914) Ecology. Lowlands. Gardens and cultivated fields. Open ground; rather dry soil. Nocturnal; sheltering during the day. Biology. Seasonality: January-March, November-December. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Pitfall trapping. Selenophorus (Selenophorus) discoderoides Schaeffer, 1910 Natural History. Unknown. Selenophorus (Celiamorphus) discopunctatus Dejean, 1829 Ecology. Lowlands. Open situations. Biology. Seasonality: March. Dispersal power. Unknown. Selenophorus (Celiamorphus) ellipticus Dejean, 1829 Ecology. Lowlands and mountains. Abandoned fields, dry fields, cultivated fields (e.g., tobacco), forest clearings; sidewalks. Open ground; dry, sandy soil covered with some vegetation. Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day under dead leaves, plant rosettes (Verbascum), and in burrows dug in the soil. Biology. Seasonality: January-December. Adults found overwintering in clumps of broomsedges. Dispersal power. Macropterous. Occasional in sea- and lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants and trees). Strongly favored by human activities. Collecting techniques. Raking the leaf litter; lifting plant rosettes; pitfall trapping; using tanglefoot screens; sweeping plants. References. Blatchley, 1910 (ecology); Lindroth, 1968 (natural history); Kirk, 1969 (biology) and 1970 (ecology); Erwin, 1981 (natural history). Selenophorus (Selenophorus) elongatus (LeConte, 1848) Ecology. Lowlands. Biology. Seasonality: January, December. Adults found overwintering under fallen tree bark. Adult food, in captivity: lepidopterous caterpillars and beetle larvae. Dispersal power. Unknown. Reference. Löding, 1945 (biology). Selenophorus (Selenophorus) famulus Casey, 1914 Natural History. Unknown. Selenophorus (Selenophorus) fatuus (LeConte, 1863) Ecology. Lowlands. Vacant lots, cultivated fields, roadsides, and sand pits. Open ground: dry, sandy soil covered with some vegetation. Nocturnal; sheltering during the day in burrows dug in the soil at the base of plants (mostly), under heaps of hay, stones, and

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other debris. Gregarious. Biology. Seasonality: January-August, October. Tenerals: January and July (AL), August (TX). Adults found overwintering under dead leaves. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Strong burrower. Collecting techniques. Digging at the base of plants; pitfall trapping; light trapping. Selenophorus (Celiamorphus) fossulatus Dejean, 1829 Ecology. Lowlands. Sand pits. Open ground; dry, sandy soil covered with sparse vegetation. Nocturnal; sheltering during the day in burrows dug in the soil at the base of plants. Biology. Seasonality: February, May, July. Adults found overwintering in sand pits; in burrows dug in the soil. Dispersal power. Wing-condition unknown. Moderate runner. Strong burrower. Collecting techniques. Digging at the base of plants; pitfall trapping. Selenophorus (Selenophorus) gagatinus Dejean, 1829 Ecology. Lowlands and mountains. Hills, dry fields, old fields, cultivated fields, pastures, meadows, roadsides, moraines, sand pits, gravel pits, thickets (e.g., Juniper), orchards (e.g., peach), forest edges and clearings, light woods. Open ground; well-drained, dry, sandy soil covered with sparse vegetation. Mostly nocturnal; usually sheltering during the day mostly under embedded stones, dead leaves, reindeer moss (Cladonia), also at the base of plants (e.g., Solidago) and under pieces of wood. Semi-gregarious (in winter). Biology. Seasonality: January-October. Tenerals: April-May (rarely; QU), JulyOctober. Adults found overwintering on hills, in meadows, pastures, moraines, and gravel pits; under stones. Adult food, in captivity: meat. Predators: toads. Defense-mechanism: when alarmed, the adult may emit a malodorous smell. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Occasional climber (on trees). Strong burrower. Strongly favored by human activities. Collecting techniques. Turning embedded stones; raking the leaf litter; lifting reindeer moss; pitfall trapping; baiting traps with molasses. References. Blatchley, 1910 (ecology); Lindroth, 1968 (biology, dispersal power); Larochelle, 1974a (ecology); Erwin, 1981 (natural history); Anderson et al. 1995 (ecology, biology). Selenophorus (Celiamorphus) granarius Dejean, 1829 Ecology. Lowlands. Vacant lots, cultivated fields (e.g., corn), roadsides, sand pits, edges of forests, and forest clearings. Open ground; dry, sandy soil covered with sparse vegetation. Nocturnal; sheltering during the day mostly in burrows dug in the soil at the base of grass tufts and under mats of reindeer moss (Cladonia), also under dead fallen plants and leaves. Semi-gregarious. Biology. Seasonality: March, May, August. Copulating pairs: May (RI). Tenerals: March (NC). Adults found overwintering in vacant lots, in the upper zone of roadsides and in wood clearings; under moss and vegetal debris. Dispersal power. Wing-dimorphic: macropterous (capable of flight) or brachypterous (incapable of flight). Occasional in seashore drift material, which indicates

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flight ability. Moderate runner. Strong burrower. Collecting techniques. Digging at the base of grass turfs; lifting reindeer moss; pitfall trapping. Selenophorus (Selenophorus) houstoni Casey, 1914 Natural History. Unknown. Selenophorus (Selenophorus) hylacis (Say, 1823) Ecology. Lowlands and mountains. Deciduous forests (e.g., Quercus, Carya) and their edges. Shaded ground. Nocturnal; sheltering during the day under the loose bark of dead standing trees (mostly) and in the soil. Biology. Seasonality: January-December. Tenerals: August (VT). Adults found overwintering in woods; in leaf litter. Predators: woodpeckers. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Frequent climber (on tree trunks). Collecting techniques. Lifting loose bark; light trapping; pyrethrum spraying. References. Blatchley, 1910 (ecology); Lindroth, 1968 (dispersal power); Will et al. 1995 (dispersal power). Selenophorus (Selenophorus) implicans Casey, 1914 Natural History. Unknown. Selenophorus (Selenophorus) laesus (LeConte, 1858) Natural History. Unknown. Selenophorus (Selenophorus) maritimus Casey, 1914 Natural History. Unknown. Selenophorus (Celiamorphus) municeps (Casey, 1924) Natural History. Unknown. Selenophorus (Selenophorus) mustus (Casey, 1914) Ecology. Lowlands. Biology. Seasonality: January, May-August. Adults found overwintering in clumps of broomsedges. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Inspecting clumps of broomsedges; light trapping. Reference. Kirk, 1969 (biology, dispersal power). Selenophorus (Celiamorphus) nanulus (Casey, 1924) Natural History. Unknown.

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Selenophorus (Selenophorus) opalinus (LeConte, 1863) Ecology. Lowlands and mountains. Hills, meadows, pastures, dry fields, relict prairies, vacant lots, wastelands, old fields, cultivated fields (e.g., soybean, alfalfa), roadsides, gardens, fencerows, sand pits, gravel pits, orchards (e.g., peach), forest edges and clearings, open forests. Open ground; dry, sandy soil covered with sparse or patchy vegetation (e.g., grass, Fragaria). Nocturnal; sheltering during the day under dead leaves and stones. Semi-gregarious. Biology. Seasonality: January, March-December. Tenerals: May (rarely; QU), mid-July to October. Adults found overwintering on hills, in meadows, vacant lots, gravel pits, at the edge of woods, and in open woods; under stones, logs, dead leaves, and in wood debris. Adult food, in captivity: an acridid. Predators: toads. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on tree trunks). Strongly favored by human activities. Collecting techniques. Raking the leaf litter; turning stones; pitfall trapping; baiting traps with syrup, molasses or fermented sugar; light trapping; turning drift material. References. Leng, 1915 (ecology); Lindroth, 1968 (natural history); Kirk, 1969 (biology); Larochelle, 1974a (ecology); Kirk, 1975c (ecology, dispersal power); Erwin, 1981 (natural history); Anderson et al. 1995 (ecology, biology); Will et al. 1995 (dispersal power). Selenophorus (Selenophorus) otiosus Casey, 1914 Natural History. Unknown. Selenophorus (Selenophorus) palliatus (Fabricius, 1798) Ecology. Lowlands. Vacant lots, cultivated fields (e.g., alfalfa), vegetable gardens (e.g., cabbage, beet), and sand dunes. Open ground; dry, sandy soil covered with some vegetation. Nocturnal; sheltering during the day in burrows dug in the soil at the base of plants (mostly), under dead leaves and pieces of wood. Biology. Seasonality: JanuaryDecember. Tenerals: late July (AL). Adults found overwintering under pieces of wood. Predators: blackbirds and meadowlarks. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Strong burrower. Collecting techniques. Digging at the base of plants; pitfall trapping; light trapping; using Japanese beetle traps. References. Leng, 1915 (ecology, dispersal power); Kirk, 1969 (dispersal power). Selenophorus (Selenophorus) pedicularius Dejean, 1829 Ecology. Lowlands, mountains, subalpine and alpine zones. Meadows, pastures, rangelands (short- and mid-grass), wastelands, cultivated fields (e.g., tobacco), and roadsides. Open ground; dry, sandy soil covered with sparse vegetation (e.g., grass). Mostly nocturnal; sometimes active in the sunshine; usually sheltering during the day in burrows dug in the soil at the base of plants, under dry cattle dung, under depressed vegetation (e.g., grass), and in clumps of broomsedges. Biology. Seasonality: February-October. Adults found overwintering in meadows; under heaps of hay. Adult food, in cap-

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tivity: grasshopper eggs and a small tenebrionid. Larval food, in captivity: grasshopper eggs. Predators: robins and starlings. Parasites: tapeworms. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Frequent in sea- and lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants and trees). Strong burrower. Strongly favored by human activities. Collecting techniques. Digging at the base of plants; pitfall trapping; using tanglefoot screens; light trapping; turning drift material; sweeping plants. References. Leng, 1915 (dispersal power); Lindroth, 1968 (dispersal power); Kirk, 1969 (ecology, dispersal power); Lavigne, 1977 (ecology, biology); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Selenophorus (Selenophorus) perpolitus Casey, 1884 Natural History. Unknown. Selenophorus (Selenophorus) planipennis LeConte, 1848 Ecology. Lowlands and mountains. Grasslands, relict prairies, cultivated fields (e.g., corn, oat), rangelands, vacant lots, hills, and fencerows. Open ground; dry, sandy soil covered with sparse vegetation (e.g., grass). Mostly nocturnal; sometimes active in the sunshine on plants (e.g., Artemisia; usually sheltering during the day under wellembedded pieces of wood. Biology. Seasonality: April-October. Adult food, in the field: plant tissue. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in drift material (lake shores), which indicates flight ability. Moderate runner. Occasional climber (on plants). Strong burrower. Strongly favored by human activities. Collecting techniques. Turning embedded pieces of wood; pitfall trapping; sweeping plants. References. Kirk, 1975c (ecology, dispersal power); Lavigne, 1978 (ecology). Selenophorus (Selenophorus) riparius Casey, 1914 Natural History. Unknown. Selenophorus (Selenophorus) schaefferi Csiki, 1932 Natural History. Unknown. Selenophorus (Selenophorus) scolopaceus Casey, 1914 Natural History. Unknown. Selenophorus (Selenophorus) sinuaticollis Notman, 1922 Natural History. Unknown.

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Selenophorus (Celiamorphus) subtropicus (Casey, 1924) Natural History. Unknown. Selenophorus (Selenophorus) trepidus (Casey, 1924) Natural History. Unknown. Selenophorus (Selenophorus) vigilans (Casey, 1914) Natural History. Unknown. Semiardistomis puncticollis (Dejean, 1831) Ecology. Lowlands. Mostly river banks and lake shores; also wet meadows. Open ground; soft, wet, clayish or muddy, rarely sandy soil covered with some vegetation (e.g., grass). Nocturnal; sheltering during the day in burrows dug in the soil (mostly), in humus, under dead leaves, and among fallen dead grass. Gregarious. Biology. Seasonality: February-December. Adults found overwintering (IN). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet. References. Blatchley, 1910 (biology); Nichols, 1988 (ecology, dispersal power) Semiardistomis viridis (Say, 1823) Ecology. Lowlands. Banks of river and brooks (mostly), lakes, and ponds. Open (mostly) or half-shaded ground; soft, wet, clayish or muddy soil (sometimes sand- or gravelmixed) covered with some vegetation. Nocturnal; sheltering during the day in burrows dug in the soil (mostly), under vegetal debris and stones. Gregarious. Biology. Seasonality: February-November. Tenerals: mid-July (WV) to late August (TX). Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Frequent flier (to artificial lights at night; sometimes swarming). Slow runner. Occasional climber (on trees). Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet; light trapping; using tanglefoot screens. References. Smith, 1910 (ecology); Erwin, 1981 (natural history); Nichols, 1988 (ecology, dispersal power). Sericoda bembidioides Kirby, 1837 Ecology. Lowlands and mountains. Coniferous forests (e.g., Thuja, Pseudotsuga, Pinus) and recently burnt forest sites. Shaded or open ground; moderately moist, often burnt soil. Nocturnal and diurnal; sheltering during the day under the loose bark of trees or active on burnt tree-stumps and branches of dead trees, burnt logs and pieces of wood, or near charcoal. Biology. Seasonality: March, May-August, October. Eggs are laid on peat moss. Defense-mechanism: when alarmed, the adult escapes by flying; iridescent color-pattern of dorsum matching shine of surrounding pieces of charcoal. Dispersal

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power. Macropterous. Frequent flier (when alarmed; to forest fires; to artificial lights at night). Moderate runner. Frequent climber (on trees, tree-stumps, and logs). Long-distance disperser. Strongly favored by forest fires. Collecting techniques. Lifting loose bark; beating trees; light trapping. References. Lindroth, 1966 (natural history); Liebherr, 1991a (natural history). Sericoda bogemannii (Gyllenhal, 1813) Ecology. Lowlands and mountains. Coniferous forests (e.g., Picea, Pseudotsuga) and recently burnt sites. Shaded or open ground; moderately moist, often recently burnt soil. Nocturnal and diurnal; sheltering during the day under the loose bark of trees damaged by forest fires or active on hot ashes. Biology. Seasonality: July-September. Tenerals: September (BC). Dispersal power. Macropterous. Frequent flier (to forest fires; artificial lights at night). Moderate runner. Frequent climber (on trees). Long-distance disperser. Strongly favored by forest fires. Collecting techniques. Lifting loose bark; beating trees; light trapping; using malaise traps. References. Lindroth, 1945, 1966, and 1986 (ecology, dispersal power); Koch, 1989 (ecology); Liebherr, 1991a (ecology). Sericoda obsoleta (Say, 1823) Ecology. Lowlands, mountains, subalpine and alpine zones. Coniferous forests (mostly; e.g., Picea, Pinus), mixed forests, recently burnt forest sites, human habitations (with wood-burning stoves), and sawmills. Shaded or open ground; moderately moist, often burnt soil. Nocturnal and diurnal; sheltering during the day under the loose bark of standing trees (live or partially burnt), under branches, stones, other debris, house floor lineoleum and tiles, in heaps of wood chips, or, active on hot ashes and plants. Adults and larvae occur among ashes after forest fires. Gregarious (in winter). Biology. Seasonality: January-November. Tenerals: late May (rarely; QU), July to late August. Adults found overwintering in woods, on hills, and in the upper zone of river banks; under the loose bark of trees (e.g., Pinus) and under stones. Adult and larval food, in captivity: mealworms. Dispersal power. Macropterous. Frequent flier (to forest fires; to artificial lights at night; sometimes swarming). Frequent in sea- and lakeshore drift material, which indicates flight ability. Moderate runner. Frequent climber (on trees and plants). Long-distance disperser. Strongly favored by forest fires and human activities. Sometimes becoming a nuisance when swarming into human habitations. Collecting techniques. Lifting loose bark; beating trees; pitfall trapping; light trapping; turning drift material; using malaise traps. References. Lindroth, 1966 (natural history); Larochelle, 1975k (ecology, dispersal power); Liebherr, 1991a (ecology, dispersal power); Dearborne and Donahue, 1993 (ecology). Sericoda quadripunctata (DeGeer, 1774) Ecology. Lowlands, mountains, subalpine and alpine zones. Coniferous forests (e.g., Pinus, Abies, Picea), mixed forests, recently burnt forest sites, sawmills, and human habitations (with wood-burning stoves). Shaded or open ground; moderately moist, often

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burnt soil. Nocturnal and diurnal; sheltering during the day under the loose bark of trees damaged by forest fires, active on tree-stumps, logs and branches of dead burnt trees, or hiding under logs, stones, and in moss. Gregarious. Biology. Seasonality: AprilOctober. Tenerals: July. Adult and larval food, in captivity: mealworms. Larvae are cannibalistic in captivity. Defense-mechanism: color-pattern of dorsum matching surrounding pieces of charcoal. Dispersal power. Macropterous. Frequent flier (to forest fires; to artificial lights at night). Frequent in seashore drift material, which indicates flight ability. Moderate runner. Frequent climber (on trees, e.g., Picea, Abies). Long-distance disperser. Strongly favored by forest fires and human activities. Remaining on a recently burnt site no more than two years. Collecting techniques. Lifting loose bark; beating trees; light trapping; turning drift material; using malaise traps. Reference. Lindroth, 1945, 1955a, 1963a, 1966, and 1986 (natural history); Holliday, 1984 (ecology) and 1991 (ecology, biology); Burakowski, 1986 (biology); Koch, 1989 (ecology); Liebherr, 1991a (ecology); Dearborne and Donahue, 1993 (ecology). Serranillus jeannelli Barr, 1995 Ecology. Mountains. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Reference. Barr, 1995 (taxonomy). Somotrichus unifasciatus (Dejean, 1831) Ecology. Warehouses. In stored products; intercepted from a shipment of nuts. Biology. Unknown. Dispersal power. Macropterous. Comments. Doubtfully established in North America. Sphaeroderus bicarinatus (LeConte, 1853) Ecology. Mountains. Forests (deciduous, mixed, coniferous). Shaded ground; moderately moist soil covered with thick leaf litter. Nocturnal; sheltering during the day in leaf litter (mostly), under the loose bark of fallen trees, in and under rotten logs, under stones and moss carpets. Biology. Seasonality: April-October. Tenerals: late August (TN). Adult and larval food, in captivity: mealworms. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; pitfall trapping. Reference. Barr, 1969 (ecology; as multicarinatus Darlington). Sphaeroderus canadensis canadensis Chaudoir, 1861 Ecology. Lowlands and mountains. Forests (mostly deciduous and mixed; also coniferous): often along rivers, brooks, the seashore, and slopes. Shaded ground; moderately moist (mostly) or wet soil, which thick leaf litter. Mostly nocturnal; sheltering during the day in leaf litter (mostly), under fallen trees, stones, and moss carpets. Larval habitat: similar to adult, in leaf litter. Biology. Seasonality: March-November. Copulating pairs: April to early July; mostly May (QU); coitus lasting 101-142 minutes. Gravids: late May to late July (QU). Tenerals: May (rarely; QU), July-September. Adults found

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overwintering on higher and drier ground, at the summit of wooded slopes and hillocks; in leaf litter. Adult food: snails, lepidopterous caterpillars and pupae (in the field); crushed snails, sawfly cocoons, mealworms, a live carabid, earthworms, and fruit (in captivity). Larval food, in captivity: slugs and snails without shells. Predators: toads (in the field) and shrews (in captivity). Defense-mechanism: when alarmed, the adult stridulates or escapes by burrowing deep into the leaf litter. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on tree trunks). Collecting techniques. Pitfall trapping; raking the leaf litter; pinning snails under boards. References. Lindroth, 1961 (natural history); Barr, 1969 (ecology); Larochelle, 1975j (ecology); Levesque et al. 1979 (biology); Bousquet and Pilon, 1980 (ecology, biology); Reeves et al. 1983 (biology); Dearborne and Donahue, 1993 (biology). Sphaeroderus canadensis lengi Darlington, 1933 Ecology. Mountains. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: March, May-June, August-October. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Sphaeroderus indianae (Blatchley, 1910) Ecology. Mountains. Forests. Shaded ground; moderately moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: February, April, September. Adults found overwintering in leaf litter. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Pitfall trapping. Sphaeroderus nitidicollis brevoorti LeConte, 1848 Ecology. Lowlands, mountains, and subalpine zone. Forests (coniferous, mixed) and their edges: often along rivers, brooks, the seashore, and slopes. Shaded ground; wet (mostly) or moist soil covered with thick leaf litter. Mostly nocturnal; sheltering during the day in leaf litter (mostly), under logs and stones; sometimes active on cloudy days. Larval habitat: similar to the adult; in leaf litter. Biology. Seasonality: April-November. Copulating pairs: late June to November (QU); coitus lasting up to 72 hours. Gravids: spring (ON); late August to late November (QU). Tenerals: mostly August (QU); autumn (ON). Adults found overwintering on higher and drier ground, in woods; under moss and in leaf litter, at the base of tree-stumps and trees, under the loose bark of fallen trees, in fallen rotten branches, and under stones. Adult and larval food, in the field: snails. Adult food, in captivity: mealworms. Larval food, in captivity: snails (without shells) and pieces of slugs. Predators: toads (in the field) and shrews (in captivity). Phoretic mites found on adult body (sometimes up to 300 mites). Defense-mechanism: when alarmed, the adult stridulates, escapes by burrowing deep into the leaf littter, or emits a molasses-like defensive fluid from the mouth. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on tree trunks, in malaise traps). Collecting techniques. Pitfall trapping; raking the leaf litter; using malaise traps. Ref-

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erences. Lindroth, 1961 (natural history); Larochelle, 1975j (ecology); Bousquet and Pilon, 1977 (biology; as nitidicollis Chevrolat) and 1980 (ecology, biology); Holliday, 1991 (ecology, biology). Sphaeroderus nitidicollis nitidicollis Guérin-Méneville, 1829 Ecology. Lowlands, mountains, and subalpine zone. Forests (deciduous, mixed, coniferous) and thickets: often along rivers, brooks, and the seashore. Shaded ground; wet (mostly) or moist soil covered with thick leaf litter. Nocturnal; sheltering during the day under dead leaves (mostly), moss, logs, and stones. Biology. Seasonality: June-September. Copulating pairs: mid-July (QU); coitus lasting 22 hours. Tenerals: mid-July and mid-August (QU). Adult food, in the field: snails. Predators: thrushes. Defense-mechanism: when alarmed, the adult escapes by burrowing deep into the leaf litter, or emits a molasses-like defensive fluid from the mouth. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (on tree trunks). Collecting techniques. Raking the leaf litter; pitfall trapping. References. Lindroth, 1955a, 1961, and 1963a (natural history). Sphaeroderus schaumii Chaudoir, 1861 Ecology. Lowlands and mountains. Deciduous forests (e.g., Acer, Celtis). Shaded ground; moderately moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: June, August-September. Tenerals: August (OH). Adult food, in the field: snails. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Pitfall trapping; night collecting with a headlamp. Reference. Will et al. 1995 (natural history) Sphaeroderus stenostomus aequalis Casey, 1920 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Sphaeroderus stenostomus lecontei Dejean, 1826 Ecology. Lowlands, mountains, and subalpine zone. Forests (mostly mixed and deciduous; also coniferous): often along rivers, brooks, lakes, and the seashore. Forest clearings and edges; adjoining roadsides, sand pits, gravel pits, fields, swamps, and marshes. Shaded (mostly) or half-shaded ground; moist or wet soil covered with thick leaf litter. Mostly nocturnal; sheltering during the day in leaf litter (mostly), under logs, stones, moss, and the loose bark of fallen trees; may invade river banks and other open sites at night (NF); sometimes active on cloudy days. Gregarious (in winter). Larval habitat: in leaf litter. Biology. Seasonality: January-December. Copulating pairs: AprilJuly (mostly), September-November; coitus lasting 72 hours on average. Oviposition: mid-May to early July (QU). Tenerals: late July to September. Adults found overwintering on higher and drier ground, in woods and at their edges; in leaf litter (mostly), in rotten tree-stumps and fallen trees, under clods of earth, and under stones. Adult food:

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snails, slugs, and lepidopterous caterpillars (in the field); snails, lepidopterous caterpillars, sawfly cocoons, mealworms, earthworms, meat, and fruit (in captivity). Larval food, in captivity: snails (without shells) and pieces of slugs. Predators: toads, frogs, crows, grouses, owls, and robins. Phoretic mites found on adult body. Defense-mechanism: when alarmed, the adult stridulates, escapes by burrowing deep into the leaf litter, or emits a Brachinus-like smoke or a defensive fluid from the pygidial glands, at a range of up to 15 cm. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Occasional climber (in malaise traps). Collecting techniques. Pitfall trapping; raking the leaf litter; turning stones; using malaise traps; baiting traps with molasses; pinning live snails under boards; night collecting with a headlamp. References. Blatchley, 1910 (ecology); Lindroth, 1955a, 1961, and 1963a (natural history); Larochelle, 1975j (ecology); Bousquet and Pilon, 1977 (biology); Levesque et al. 1979 (biology); Bousquet and Pilon, 1980 (ecology, biology); Levesque et al. 1980 (biology); Reeves et al. 1983 (biology); Levesque and Levesque, 1986 (biology); Bousquet, 1987b (biology). Sphaeroderus stenostomus stenostomus (Weber, 1801) Ecology. Lowlands and mountains. Deciduous forests (mostly), coniferous forests, and borders of eutrophic marshes. Shaded ground; dry soil. Mostly nocturnal; sheltering during the day under dead leaves, stones, and logs; sometimes active on cloudy days. Biology. Seasonality: February-November. Copulating pairs: April, October-November. Tenerals: July (DC, VA). Adults found overwintering (DC). Adult food: snails and slugs (in the field); snails and meat (in captivity). Predators: toads, crows, and thrushes. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; turning stones; pitfall trapping; light trapping. References. Ulke, 1903 (ecology, biology); Erwin, 1981 (natural history); Anderson et al. 1995 (ecology). Stenocrepis (Stenous) cuprea (Chaudoir, 1843) Ecology. Lowlands and uplands. Borders of rivers (mostly), brooks, lakes, and ponds; bayous. Open ground; soft, wet, muddy or clayish soil (sometimes sand-mixed) without vegetation. Nocturnal; sheltering during the day in burrows dug in the soil, under well-embedded debris and partly submerged logs. Biology. Seasonality: January-November. Tenerals: March (FL), July (TX), September (IN). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Strong burrower. Good swimmer. Collecting techniques. Pouring water over the ground; treading the soil with the feet; turning embedded debris. References. Löding, 1945 (ecology); Erwin, 1981 (natural history); Bousquet, 1996a (taxonomy, ecology). Stenocrepis (Stenous) duodecimstriata (Chevrolat, 1835) Ecology. Lowlands. Borders of marshes, ponds, lakes, slow rivers and brooks, and irrigation canals; swamps. Open ground; soft, wet, muddy, clayish or silty soil, rich in vegetal debris (e.g., dead leaves), covered with some vegetation. Nocturnal; sheltering dur-

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ing the day in burrows dug in the soil and under embedded debris (e.g., boards, rubbbish). Biology. Seasonality: January-December. Tenerals: March-April, June-July. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strong burrower. Good swimmer. Collecting techniques. Pouring water over the ground; treading the soil with the feet; turning embedded debris; light trapping. Reference. Bousquet, 1996a (taxonomy). Stenocrepis (Stenous) elegans (LeConte, 1851) Ecology. River banks. Open ground; soft, wet, muddy or silty soil. Nocturnal; sheltering during the day in burrows dug in the soil or under embedded debris. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet; turning embedded debris. Reference. Bousquet, 1996a (taxonomy). Stenocrepis (Stenocrepis) insulana (Jacquelin du Val, 1857) (Oodes texanus LeConte, 1863) Ecology. Lowlands. Swamp borders. Wet soil. Biology. Seasonality: June-August, November. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Strong burrower. Reference. Bousquet, 1996a (taxonomy). Stenocrepis (Stenous) mexicana (Chevrolat, 1835) (Stenocrepis (Crossocrepis) quatuordecimstriata (Chaudoir, 1843)) Ecology. Lowlands. Banks of rivers (mostly) and brooks; lake shores and wet pastures. Open ground; soft, wet, muddy or silty soil, bare or covered with sparse to moderate vegetation. Nocturnal; sheltering during the day in burrows dug in the soil and under embedded debris (mostly), under dead leaves, pieces of wood, logs, and stones. Biology. Seasonality: January-November. Tenerals: late July and August (KY). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Slow runner. Moderate runner. Strong burrower. Good swimmer. Collecting techniques. Pouring water over the ground; treading the soil with the feet; turning embedded debris. References. Kirk, 1969 (dispersal power; as quatuordecimstriata (Chaudoir)); Bousquet, 1996a (taxonomy). Stenocrepis (Crossocrepis) quatuordecimstriata (Chaudoir, 1843), see Stenocrepis (Stenous) mexicana (Chevrolat, 1835). Reference. Bousquet, 1996a (taxonomy). Stenocrepis (Stenocrepis) texana (LeConte, 1863), see Stenocrepis (Stenocrepis) insulana (Jacquelin du Val, 1857). Reference. Bousquet, 1996a (taxonomy).

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Stenocrepis (Stenous) tibialis (Chevrolat, 1834) Ecology. Lowlands. Borders of salt marshes. Open ground; soft, wet, saline soil. Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: AprilJune, October. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Reference. Bousquet, 1996a (taxonomy). Stenolophus (Stenolophus) abstinens Casey, 1914 Natural History. Unknown. Stenolophus (Stenolophus) anceps LeConte, 1857 Ecology. Lowlands and mountains. Borders of eutrophic marshes, pools, ponds, lakes, reservoirs, slow rivers, and irrigation canals. Open ground; wet, clayish soil, rich in organic debris (e.g., dead leaves), covered with dense vegetation (e.g., grass, Carex). Nocturnal; sheltering during the day mostly under layers of vegetal debris (e.g., dead leaves, grass), rarely under stones. Biology. Seasonality: April-July. Tenerals: mid-June and early July. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Treading the vegetation into the water. Reference. Lindroth, 1968 (natural history). Stenolophus (Agonoderus) binotatus (Casey, 1914) Ecology. Lowlands. Biology. Seasonality: July. Dispersal power. Macropterous, probably capable of flight. Stenolophus (Stenolophus) carbo Bousquet, 1993 (Stenolophus carbonarius (Dejean, 1829)) Ecology. Lowlands. A swamp (Typha). A lake shore. Vicinity of a hillside spring. An old field. A deciduous forest. Open or shaded ground; wet soil. Nocturnal; sheltering during the day under debris and stones. Biology. Seasonality: March-October. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on trees). Collecting techniques. Turning debris and stones; using tanglefoot screens; sifting debris. References. Blatchley, 1910 (ecology; as carbonarius (Dejean)); Lindroth, 1968 (natural history; as carbonarius (Dejean)); Kirk, 1975c (dispersal power; as carbonarius (Dejean)). Stenolophus (Stenolophus) cincticollis LeConte, 1858 Ecology. River banks. Biology. Unknown. Dispersal power. Macropterous, probably capable of flight.

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Stenolophus (Agonoderus) comma (Fabricius, 1775) Ecology. Lowlands and mountains. Borders of marshes, pools, lakes, reservoirs, rivers, brooks, and irrigation canals; swamps. Cultivated fields (mostly corn; also wheat, barley, tobacco, carrot, oat, soybean, dry bean, alfalfa, asparagus, cabbage, rutabaga, radish), plowed fields, rangelands, pastures, vacant lots, roadsides, sand pits, gravel pits, and orchards (e.g., apple). Open ground; moderately moist, mostly sandy soil covered with some vegetation. Mostly nocturnal; quite active in the spring sunshine; usually sheltering during the day in ground tunnels (mostly; 2-10 cm long), in soil cracks, under silt cakes, dead leaves, crust of dead plants, pieces of wood, well-embedded stones, and other debris. Larval habitat: tunnels in the soil. Gregarious. Biology. Seasonality: January-December. Oviposition: late April to late July (SD). Eggs are laid singly in cells in the soil. Tenerals: mid-June to October. Adults found overwintering on higher and drier ground, in fields, in the upper zone of sand dunes, sand pits, and marshes, at the edge of woods and in woods; mostly in burrows dug in the soil (2-50 cm deep), also in leaf litter and rotten logs, under stones and the loose bark of fallen trees. Adult food: flies, lygaeids, ants, aphids, grasshopper eggs, corn seed, and fungi (in the field); lepidopterous caterpillars, dipterous immature stages, chrysomelid eggs and larvae, grasshopper eggs, dead lygaeid bugs, and plant seeds (oats, barley, wheat) (in captivity). Larval food, in captivity: lepidopterous caterpillars, dipterous larvae and adults, beetles, trichopterans, and homopterans. Adults and larvae are cannibalistic in captivity. This species is mostly carnivorous, but will eat vegetal matter if necessary. Economic importance: sometimes a severe pest of corn. Predators: frogs, toads, bluebirds, and racerunners. Parasites: mites. Defense-mechanism: when alarmed, the adult escapes by digging into the soil; body color-pattern matching the ground. Dispersal power. Macropterous. Frequent flier (sometimes swarming by the thousands; in the spring sunshine; to artificial lights at night). Dispersal flights: mid-April to mid-October (SD). Frequent in lake- and seashore drift material, which indicates flight ability. Slow runner. Occasional climber (on plants and trees). Strong burrower. Strongly favored by human activities. Spring swarms sometimes becoming a nuisance around illuminated human habitations. Collecting techniques. Pouring water over the ground; treading the soil with the feet; pitfall trapping; light trapping; turning drift material; sweeping plants. References. Johnson, 1949 (biology); Lindroth, 1955a, 1963a, and 1968 (natural history); Kirk, 1969 (ecology, dispersal power), 1975a (natural history), and 1975c (ecology, dispersal power); Tomlin, 1975b (biology); Wyman et al. 1976 (biology, dispersal power); Hsin et al. 1979 (biology); Pausch, 1979 (biology); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Stenolophus (Agonoleptus) conjunctus (Say, 1823) Ecology. Lowlands, mountains, subalpine and alpine zones. Grasslands, prairies, sagebrush areas, meadows, rangelands, pastures, cultivated fields (e.g., corn, wheat, clover), old fields, pastures, lawns, vacant lots, roadsides, sand pits, gravel pits, ski fields, orchards (e.g., peach), forest edges and clearings, and light woods. Greenhouses (occasionally). Open ground; dry, usually sandy soil covered with sparse vegetation. Mostly

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nocturnal; quite active in the spring sunshine; usually sheltering during the day under stones, dead leaves, plant debris (e.g., corn), reindeer moss (Cladonia), heaps of hay, dry cow dung, and in cavities in the soil at the base of plants. Gregarious (in winter). Biology. Seasonality: January-December. Copulating pairs: May (QU). Gravids: early May and late June (QU). Tenerals: March-April (rarely), July to late November; mostly in July-August. Adults found overwintering on higher and drier ground, in vacant lots, pastures, on hills, in the upper zone of sand pits and gravel pits, at the edge of woods and in woods; mostly under well-embedded stones, plant rosettes (Verbascum), and dead leaves, also under logs, wood chips, in grass and clumps of moss, and in burrows dug in the soil (as deep as 7.5 cm). Adult food: coccinellids (in the field); meat (in captivity). Predators: toads (regularly), pintails, robins, starlings, and trashers. Parasites: tapeworms. Phoretic mites found on adult body. Dispersal power. Wing-dimorphic: macropterous (mostly) or brachypterous (incapable of flight). Frequent flier (in the spring sunshine; at dusk; to artificial lights at night; into rotary traps). Frequent in lake- and seashore drift material, which indicates fligh ability. Moderate runner. Occasional climber (on plants). Strong burrower. Strongly favored by human activities and spring floods along large rivers. Collecting techniques. Turning stones; raking the leaf litter; pitfall trapping; baiting traps with molasses and fermented sugar; light trapping; turning drift material; using rotary traps; sweeping plants. References. Blatchley, 1910 (ecology, dispersal power); Lindroth, 1968 (natural history); Kirk, 1970 (ecology) and 1975c (ecology, dispersal power); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Stenolophus (Stenolophus) dissimilis Dejean, 1829 Ecology. Lowlands. Lake shores. Open ground; soft, wet, muddy soil, somewhat mixed with dead leaves. Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: March-August, October-November. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Strong burrower. Collecting technique. Pitfall trapping. Stenolophus (Stenolophus) extensicollis Casey, 1924 Natural History. Unknown. Stenolophus (Stenolophus) flavipes LeConte, 1858 Ecology. Unknown. Biology. Seasonality: spring to fall. Dispersal power. Macropterous, probably capable of flight. Stenolophus (Stenolophus) fuliginosus Dejean, 1829 Ecology. Lowlands, mountains, subalpine and alpine zones. Borders of pools, ponds, lakes, marshes (freshwater, saline), flood-plain forests, slow brooks and rivers, and roadside ditches; wet meadows, beaver houses, mud flats (in sand pits and gravel pits), and salt flats. In habitats often highly polluted by human activities. Open or slightly shaded

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ground; wet soil, rich in clay and vegetal debris (e.g., dead leaves), covered with rather thin depressed vegetation (e.g., Agrostis, Carex). Mostly nocturnal; active in the spring sunshine; usually sheltering during the day mostly under dead leaves and fallen plants (e.g., grass), also under stones and in cracks in the soil. Gregarious (in winter). Biology. Seasonality: January-December. Copulating pairs: May (QU). Gravids: late May and late June (NY, VT). Tenerals: early May (rarely; QU), mid-June to mid-August, November. Adults found overwintering on higher and drier ground, in the upper banks of marshes, swamps, sand pits, gravel pits, on top of sand dunes, in vacant lots, at the edge of woods, and in woods; mostly in leaf litter and under stones, also in clumps of grass and moss, under pieces of wood, the loose bark of fallen tree-stumps, in rotten logs, and in burrows dug in the soil (as deep as 10 cm). Adult food, in captivity: pieces of earthworms and a corn kernel. Predators: frogs, toads, and flycatchers. Parasites: Laboulbeniales (Fungi). Defense-mechanism: body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (int the spring sunshine; at sunset; to artificial lights at night; into rotary traps). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Strong burrower. Occasional climber (on plants and trees). Moderately favored by human activities. Collecting techniques. Raking the leaf litter; turning fallen dead plants; using rotary traps; light trapping; turning drift material; baiting traps with molasses and fermented sugar; sweeping plants. References. Blatchley, 1910 (biology); Lindroth, 1955a, 1963a, and 1968 (natural history); Kirk, 1975c (ecology, dispersal power). Stenolophus (Stenolophus) fuscatus Dejean, 1829 Ecology. Lowlands and mountains. Mostly borders of lakes, ponds, pools, and marshes; also swamps, meadows, vacant lots, roadsides, sand pits, and gravel pits. Not restricted to the vicinity of water. Ground partly shaded by trees or bushes; moderately dry, sandy soil covered with some vegetation. Nocturnal; sheltering during the day under dead leaves, fallen dead plants (e.g., Juncus, grass), at the base of standing plants (e.g., Juncus), under stones, and in burrows dug in the soil. Gregarious (in winter). Biology. Seasonality: February-August, November. Tenerals: August (ON, QU). Adults found overwintering on higher and drier ground, in the upper zone of marsh borders, lake shores, sand pits and gravel pits, on hills, in pastures and vacant lots, at the edge of woods and in woods; in burrows dug in the soil (as deep as 10 cm), under stones, and in clumps of dead leaves and grass. Defense-mechanism: body color-pattern matching the soil. Dispersal power. Macropterous. Occasional flier (at dusk). Occasional in lake- and seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Raking the leaf litter; pouring water over the ground. References. Blatchley, 1910 (ecology); Lindroth, 1968 (natural history). Stenolophus (Stenolophus) humidus Hamilton, 1893 Ecology. Lowlands. Borders of small eutrophic marshes: along brooks. Open ground; wet, sandy soil, clayish or muddy, sometimes mixed with sand, covered with some vegetation (Salix, Typha, Juncus, Equisetum). Nocturnal; sheltering during the day under

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dead leaves and fallen dead plants (e.g., Juncus). Biology. Seasonality: April-August. Predators: toads. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; turning vegetal debris; pouring water over the ground. References. Larochelle, 1976c (ecology); Erwin, 1981 (natural history). Stenolophus (Stenolophus) incultus Casey, 1914 Ecology. Lowlands and mountains. Border of an open mixed forest, on moist soil consisting of fine sand, covered with sparse vegetation, except for a few mossy patches (BC). Biology. Seasonality: April-August. Dispersal power. Macropterous, probably capable of flight. Reference. Lindroth, 1968 (natural history). Stenolophus (Agonoderus) infuscatus (Dejean, 1829) Ecology. Lowlands. Macrohabitat unknown. Moderately moist soil. Nocturnal; sheltering during the day under vegetal debris. Biology. Seasonality: January-September, November-December. Adults found overwintering. Dispersal power. Macropterous. Frequent flier (sometimes swarming by the thousands; to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Slow runner. Occasional climber (on trees). Strong burrower. Collecting techniques. Turning vegetal debris; light trapping; using tanglefoot screens. References. Leng, 1915 (dispersal power); Kirk, 1969 and 1970 (dispersal power). Stenolophus (Agonoderus) lecontei (Chaudoir, 1868) Ecology. Lowlands and uplands. Borders of rivers (mostly), brooks, lakes, ponds, and irrigation canals; mud flats, wet meadows, and flood-plain forests. Cultivated fields (mostly corn; also sorghum, soybean, tobacco, cabbage, rutabaga, radish), wastelands, vacant lots, roadsides, orchards (e.g., peach), and tree plantations (e.g., Pinus). Not restricted to the vicinity of water. Open ground; wet or moist soil consisting of clay or mud or sandy loam, sometimes pure sand, covered with sparse to moderate vegetation. Nocturnal; sheltering during the day mostly in burrows dug in the soil, also under dead leaves and stones. Biology. Seasonality: January, March-October. Tenerals: July-August. Adults found overwintering in fields; under dead leaves. Adult food: insects (e.g., lygaeid bugs) and corn seeds (in the field); mostly animal food, e.g., chrysomelids, lepidopterous eggs and larvae, but also to a lower degree, corn seeds (in captivity). Larval food, in captivity: animal matter. This species is mostly carnivorous, but will eat vegetal matter if necessary. Economic importance: sometimes a severe pest of corn. Predators: bluebirds, quails, catbirds, prairie chickens, crows, flycatchers, larks, quails, robins, starlings, swallows, trashers, thrushes, and woodpeckers. Parasites: mites. Defense-mechanism: body color matching the soil. Dispersal power. Macropterous. Frequent flier (swarming to artificial lights at night). Frequent in lakeshore drift material, which indicates flight ability. Slow runner. Occasional climber (on plants). Strong burrower. Strongly favored by human activities. Collecting techniques. Pitfall trapping; pouring water

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over the ground; treading the soil with the feet; light trapping; turning drift material; sweeping plants. References. Herne, 1963 (biology); Wyman et al. 1976 (biology, dispersal power); Pausch, 1979 (biology); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power). Stenolophus (Stenolophus) limbalis LeConte, 1857 Ecology. Lowlands. Damp places. Nocturnal; sheltering during the day. Biology. Seasonality: June. Predators: phalaropes. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting technique. Pitfall trapping. Reference. Hatch, 1953 (ecology). Stenolophus (Agonoderus) lineola (Fabricius, 1775) Ecology. Lowlands and uplands. Borders of rivers, ponds, reservoirs, and lakes. Cultivated fields (e.g., corn), golf courses (tunnelling in golf greens), and tree plantations (e.g., Pinus). Not restricted to the vicinity of water. Open ground; moist or dry, usually sandy soil covered with some vegetation. Nocturnal; sheltering during the day in soil tunnels (mostly) and under stones. Biology. Seasonality: March-October. Adults found overwintering in vacant lots; under dead leaves and heaps of hay. Adult food: lepidopterous caterpillars, other insects, and corn seeds (in the field); dipterous larvae and corn seeds (in captivity). Larval food, in captivity: animal food. Economic importance: sometimes a pest of corn. Predators: flycatchers and robins. Defense-mechanism: body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (sometimes swarming; to artificial lights at night). Frequent in lake- and seashore drift material, which indicates flight ability. Slow runner. Strong burrower. Long-distance disperser (over the sea up to 100 km from the mainland). Strongly favored by human activities. Collecting techniques. Pitfall trapping; pouring water over the ground; treading the soil with the feet; light trapping; turning drift material. References. Lindroth, 1968 (natural history); Kirk, 1969 (ecology), 1970, and 1975c (ecology, dispersal power). Stenolophus (Agonoderus) maculatus (LeConte, 1868) Ecology. Cultivated fields (e.g., lettuce). Nocturnal; sheltering during the day in burrows dug in the soil. Biology. Seasonality: May, July. Adult food, in the field: germinating lettuce seeds. Adult and larval food, in captivity: lepidopterous eggs and larvae. Dispersal power. Macropterous, probably capable of flight. Slow runner. Strong burrower. Collecting technique. Pitfall trapping. Stenolophus (Stenolophus) megacephalus Lindroth, 1968 Ecology. Lowlands and mountains. Borders of Sphagnum-bogs: often along lakes and ponds. Also swamps. Open ground; soft, mostly floating soil covered with moss (Sphagnum). Nocturnal; sheltering during the day in moss. Biology. Seasonality: April-August. Adults found overwintering on higher and drier ground, in the upper zone of lake

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shores; in leaf litter. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting technique. Treading moss carpets into the water. References. Lindroth, 1968 (natural history); Will et al. 1995 (dispersal power). Stenolophus (Stenolophus) ochropezus (Say, 1823) Ecology. Lowlands and mountains. Borders of eutrophic marshes, ponds, lakes, and pools; swamps, swamp-forests, flood-plain forests, mud flats, wet meadows; borders of of slow rivers, brooks, irrigation canals, and roadsides; sand pits and gravel pits with temporary water; beaver houses. Open (mostly) or shaded ground; poorly drained, soft, wet, clayish or muddy soil covered with dense vegetation (e.g., grass, Typha). Nocturnal; sheltering during the day mostly in burrows and cracks in the soil, and under wellembedded pieces of wood, also under dead leaves, mats of dead fallen plants, in moss, in clumbs of broomsedges, and under stones. Gregarious (in winter). Biology. Seasonality: January-December. Gravids: late June (QU, VT). Tenerals: May (rarely; QU), JulySeptember; mostly after mid-July. Adults found overwintering on higher and drier ground, in the upper zone of lake shores, pond edges, marshes, river banks, sand pits, dunes, gravel pits, and roadside ditches, in vacant lots and hawthorn hedges, at the edge of woods and in woods; in the soil and leaf litter, under stones, logs, in rotten logs and tree-stumps, and under heaps of hay. Adult food, in captivity: a corn kernel. Predators: frogs and robins. Parasites: Laboulbeniales (Fungi). Phoretic mites found on adult body. Dispersal power. Macropterous. Frequent flier (sometimes swarming; to artificial lights at night; into rotary traps). Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants and trees; up to 2 m above ground). Strong burrower. Favored by spring floods along rivers. Collecting techniques. Treading the vegetation into the water; pouring water over the ground; turning embedded pieces of wood; light trapping; using rotary traps; turning drift material; baiting traps with molasses and fermented sugar; using tanglefoot screens; sweeping plants. References. Blatchley, 1910 (biology); Leng, 1915 (ecology); Lindroth, 1968 (natural history); Kirk, 1969 (ecology, dispersal power), 1970 (dispersal power), and 1975c (ecology, dispersal power); Erwin, 1981 (natural history); Bousquet, 1997a (taxonomy). Stenolophus (Agonoleptus) parviceps (Casey, 1914) Ecology. Uplands. Muddy shores of a saline lake (MN). Nocturnal. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Stenolophus (Stenolophus) peregrinus Casey, 1914 Natural History. Unknown. Stenolophus (Stenolophus) plebejus Dejean, 1829 Ecology. Lowlands, mountains, and subalpine zone. Borders of eutrophic marshes, ponds, and lakes; cranberry bogs, swamps, low forests, mud flats, wet meadows; borders of slow

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rivers and brooks; roadside ditches and beaver houses. Open ground; soft, wet, clayish or muddy soil covered with dense vegetation (e.g., Juncus, Typha, Carex). Nocturnal; sheltering during the day mostly in burrows dug in the soil and under well-embedded pieces of wood, also under dead leaves and mats of dead fallen plants. Biology. Seasonality: January-September, December. Tenerals: April (rarely; FL), July (mostly), September. Adults found overwintering on higher and drier ground, in the upper zone of pond edges and roadside ditches, in vacant lots, and woods; in burrows dug in the soil (as deep as 4 cm), in leaf litter, and moss. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Strong burrower. Collecting techniques. Treading the vegetation into the water; turning embedded pieces of wood; baiting traps with molasses or fermented sugar. References. Kirk, 1969 (dispersal power); Erwin, 1981 (natural history). Stenolophus (Stenolophus) remissus Casey, 1914 Natural History. Unknown. Stenolophus (Agonoleptus) rotundatus LeConte, 1863 Ecology. Lowlands and mountains. Pastures, cultivated fields (e.g., alfalfa), old fields, golf courses, lawns, and river banks. Open ground; sandy soil covered with some vegetation. Nocturnal; sheltering during the day under heaps of grass. Biology. Seasonality: April-August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Fast runner. Occasional climber (on trees). Collecting techniques. Turning heaps of grass; pitfall trapping. References. Lindroth, 1968 (dispersal power); Kirk, 1970 (dispersal power); Lavigne, 1978 (ecology); Will et al. 1995 (dispersal power). Stenolophus (Agonoleptus) rotundicollis (Haldeman, 1843) Ecology. Lowlands and mountains. A garden, in heaps of grass (MA). A mixed forest (CT). Nocturnal; sheltering during the day. Biology. Seasonality: March-June, August. Adults found overwintering (SC). Dispersal power. Macropterous, capable of flight. Occasional in lakeshore drift material, which indicates flight ability. Slow runner. Occasional climber (on plants and trees). Collecting technique. Sweeping plants. References. Lindroth, 1968 (natural history); Erwin, 1981 (natural history). Stenolophus (Agonoderus) rugicollis (LeConte, 1859) Ecology and Biology. Unknown. Dispersal power. Macropterous, probably capable of flight. Stenolophus (Stenolophus) semitinctus Casey, 1914 Natural History. Unknown.

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Stenolophus (Stenolophus) splendidulus Motschulsky, 1864 Natural History. Unknown. Reference. Bousquet, 1997a (taxonomy). Stenolophus (Stenolophus) spretus Dejean, 1831 Ecology. Lowlands. Edge of a pond (SC). Biology. Seasonality: March-August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). References. Leng, 1915 (dispersal power); Kirk, 1969 (ecology). Stenolophus (Agonoleptus) unicolor dolosus Casey, 1914 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Stenolophus (Agonoleptus) unicolor unicolor Dejean, 1829 Ecology and Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Stenomorphus californicus californicus (Ménétriés, 1843) Ecology. Desert grasslands areas. Nocturnal; sheltering during the day. Biology. Seasonality: August-October. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Reference. Ball et al. 1991 (natural history). Stenomorphus californicus rufipes LeConte, 1858 Ecology. Lowlands. Pastures, cultivated fields (e.g., oats), and grasslands (Acacia). Open ground; moderately moist, clay soil. Nocturnal; sheltering during the day under debris. Biology. Seasonality: January-December. Predators: toads. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Turning debris; pitfall trapping. Reference. Ball et al. 1991 (natural history). Stenomorphus convexior Notman, 1922 Ecology. Lowlands and mountains. Disturbed areas close to water (e.g., pond borders, chicken yards). Open ground; moist soil. Nocturnal; sheltering during the day under debris (e.g., dead leaves) and in burrows dug in the soil under stones. Biology. Seasonality: January, May-September, November. Defense-mechanism: when alarmed, the adult escapes by retreating into its burrow. Dispersal power. Macropterous, probably capable of flight. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Turning debris; pitfall trapping. Reference. Ball et al. 1991 (natural history).

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Stenomorphus sinaloae Darlington, 1936 Ecology. Lowlands and mountains. Pastures. Mostly nocturnal; sheltering during the day. Biology. Seasonality: January, June-November. Dispersal power. Macropterous. Frequent flier (swarming during the day; to artificial lights at night). Moderate runner. Collecting techniques. Pitfall trapping; light trapping. Reference. Ball et al. 1991 (natural history). Stereocerus haematopus (Dejean, 1831) Ecology. Lowlands, mountains, subalpine and alpine zones. Moraines, spruce-bogs, and alpine meadows. Ground shaded by bushes (e.g., dwarf Betula, Alnus); rather dry, often sandy, sometimes gravelly soil covered with scattered stones and some vegetation (e.g., Empetrum, Ledon, Kalmia). Nocturnal; sheltering during the day under stones and in leaf litter. Biology. Seasonality: May-September. Copulating pairs: mid-June (NH). Gravids: June and early July (NH, QU). Tenerals: late July to September. Adult food, in captivity: mealworms and shrimps. Larval food, in captivity: mealworms. Parasites: tachinid larvae. Dispersal power. Wing-dimorphic: macropterous (mostly) or brachypterous (incapable of flight). Moderate runner. Collecting techniques. Turning stones; pitfall trapping. References. Lindroth, 1955a, 1963a, and 1966 (natural history); Bousquet, 1999 (biology). Stereocerus rubripes (Motschulsky, 1860) Ecology. Tundra. A dry, sandy slope covered with heath-type vegetation. Nocturnal; sheltering during the day. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Reference. Lindroth, 1966 (ecology, dispersal power; as kryzhanovskii Lindroth, 1966). Stomis (Stomis) pumicatus (Panzer, 1795) Ecology. Lowlands. Grasslands, cultivated fields, open deciduous forests, forest edges and clearings. Europe: moist biotopes along rivers and brooks, open lands, deciduous forests, parks, gardens, and rodent nests. Open ground; moderately moist, usually clay soil (sometimes sand-mixed) covered with some vegetation (e.g., grass) or leaf litter. Nocturnal; sheltering during the day among dead leaves. Biology. Seasonality: JuneJuly, September. Tenerals: September (NS). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strongly favored by human activities. Collecting techniques. Raking the leaf litter; pitfall trapping. References. Larsson, 1939 (biology); Lindroth, 1945 and 1966 (natural history); Sharova, 1981 (ecology, biology); Lindroth, 1986 (natural history); Koch, 1989 (ecology). Stomis (Neostomis) termitiformis (Van Dyke, 1925) Ecology. Lowlands and mountains. Forests (e.g., Picea, Pseudotsuga, Quercus) and thickets: along streams, sea beaches, and sea dunes. Shaded (mostly) or open ground. Nocturnal; sheltering during the day under well-embedded logs. Biology. Seasonality: June.

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Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting techniques. Turning embedded logs; pitfall trapping. Reference. Hacker, 1968 (ecology). Syntomus americanus (Dejean, 1831) Ecology. Lowlands, mountains, subalpine and alpine zones. Abandoned fields, pastures, meadows, hills, cultivated fields (e.g., wheat, barley, carrot), roadsides, grassy borders of fields, sand pits, gravel pits, ski fields, thickets (Juniper), forest clearings and roads, tree plantations (e.g., Pinus), and light forests. Open ground; dry, usually sandy, rarely peaty soil covered with sparse low vegetation (e.g., Rumex). Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days under dead leaves, stones, reindeer moss (Cladonia), in the soil at the base of plants, under heaps of hay, plant rosettes (Verbascum), and pieces of wood. Biology. Seasonality: February-December. Gravids: mid-July (AB). Tenerals: April (rarely; QU); late June to September. Adults found overwintering on higher and drier ground, in the upper zone of sand pits and roadsides, on hills, in vacant lots, at the edge of woods, and in open woods; mostly under stones and leaf litter, also in moss or under fallen trees. Adult food, in captivity: lygaeid larvae and lepidopterous eggs. Predators: toads and gulls. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (in the bright sunshine; to artificial lights at night). Occasional in lake- and seashore drift material, which indicates flight ability. Fast runner. Occasional climber (on trees, e.g., Pinus, Prunus). Strongly favored by human activities. Collecting techniques. Raking the leaf litter; turning stones and reindeer moss; pitfall trapping. References. Lindroth, 1955a, 1963a, and 1969a (natural history); Frank, 1971b (biology); Larochelle, 1973e (dispersal power); Levesque et al. 1979 (biology). Synuchus (Pristodactyla) dubius (LeConte, 1854) Ecology. Mountains. Coniferous forests (e.g., Pinus); their edges and adjoining slopes. Shaded or open ground; rocky soil. Nocturnal; sheltering during the day under logs (mostly). Biology. Seasonality: May-June, August. Tenerals: late June (AZ). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning logs; pitfall trapping. Synuchus (Pristodactyla) impunctatus (Say, 1823) Ecology. Lowlands, mountains, and subalpine zone. Light forests, their clearings and edges, tree plantations (e.g., Pinus) and thickets (e.g., Alnus, Juniper). Adjoining moraines, vacant lots, sand pits, cultivated fields (e.g., barley). Vicinity of human habitations (urban and rural sites). Ground shaded by trees, shrubs and bushes (e.g., Acer, Alnus, Rubus) or open; rather dry, sandy soil covered with thick leaf litter. Mostly nocturnal; usually sheltering during the day among dead leaves (mostly), under pieces of wood, logs, stones, reindeer moss (Cladonia), moss, the loose bark of standing trees, heaps of hay, and pieces of cardboard; sometimes active on cloudy days. Biology. Sea-

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sonality: April-November. Copulating pairs: September (QU). Gravids: mostly JulyAugust (QU). Tenerals: April (rarely; QU), June-September; mostly mid-June to midJuly. Adult food: lepidopterous immature stages and plant seeds (Melampyron lineare Desrousseaux) (in the field); mealworms and seeds (Melampyron) (in captivity). Beetles kept alive for 3 months on nothing but a seed diet. Seed-caching is exhibited by adults. Larval food, in captivity: mealworms. Predators: toads (frequently). Parasites: mites and nematodes. Defense-mechanism: when alarmed, the adult escapes by running and digging into the leaf litter. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (to artificial lights at night). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on trees, e.g., Picea, Abies). Collecting techniques. Pitfall trapping; raking the leaf litter; using interception traps. References. Lindroth, 1955a, 1963a, and 1966 (natural history); Frank, 1971b (biology); Manley, 1971 (biology); Bousquet and Pilon, 1977 (biology); Larochelle, 1977c (dispersal power); Levesque et al. 1979 (biology); Liebherr and Mahar, 1979 (biology); Erwin, 1981 (natural history); Reeves et al. 1983 (biology); Levesque and Levesque, 1986 (biology); Epstein and Kulman, 1990 (ecology, biology); Holliday, 1991 (biology); Niemelä and Spence, 1991 (ecology). Tachys bradycellinus Hayward, 1900 Ecology. Lowlands. Biology and Dispersal power. Unknown. Tachys bryanti Lindroth, 1966 Ecology. Unknown. Biology. Seasonality: September. Dispersal power. Macropterous, probably capable of flight. Tachys corax LeConte, 1852 Ecology. Lowlands and uplands. Borders of alkaline lakes, ponds, marshes, and slow rivers; salt flats. Open ground; soft, wet, muddy, alkaline soil, bare or covered with sparse vegetation. Nocturnal; sheltering during the day in burrows and cracks in the soil, under well-embedded pieces of wood, and at the base of plants. Biology. Seasonality: June-September. Tenerals: late July (UT). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Strong burrower. Collecting techniques. Pouring water over the ground; treading the soil with the feet; turning embedded pieces of wood. Tachys halophilus Lindroth, 1966 Ecology. Margins of alkaline lakes and ponds. Open ground; firm, moderately wet clay soil covered with sparse vegetation (e.g., Glaux, Spergularia, Triglochin, Distichlis). Biology. Seasonality: April-August, October. Tenerals: mid-July (MB). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Reference. Lindroth, 1966 (natural history).

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Tachys litoralis Casey, 1884 Ecology. Coastal lowlands. Sea beaches; borders of salt marshes; salt meadows. Nocturnal; sheltering during the day under pieces of wood or dry cow dung. Biology. Seasonality: June-September. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet; turning debris. References. Leng, 1915 (ecology, dispersal power; as occultator Casey); Löding, 1945 (ecology; as occultator Casey). Tachys misellus LaFerté-Sénectère, 1841 Ecology. Coastal lowlands. Beaches. Nocturnal; sheltering during the day under pieces of wood or dry cow dung. Biology and Dispersal power. Unknown. Tachys mordax LeConte, 1852 Natural History. Unknown. Tachys pallidus Chaudoir, 1868 Ecology. Borders of salt marshes and ponds. Open ground; wet, sandy, saline soil, bare or covered with sparse vegetation (e.g., Salicornia). Nocturnal; sheltering during the day in cracks in the soil. Biology. Seasonality: June-July. Dispersal power. Wing-condition unknown. Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Tachys pulchellus LaFerté-Sénectère, 1841 Ecology. Lowlands. Sea beaches, borders of salt marshes, and salt flats. Open ground; wet, sandy, saline soil, bare or covered with sparse vegetation (e.g., Salicornia). Nocturnal; sheltering during the day in cracks in the soil or under vegetal debris. Biology. Seasonality: July. Tenerals: late July (TX). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Long-distance disperser (over the sea up to 60 km from the mainland). Collecting techniques. Pouring water over the ground; treading the soil with the feet; turning debris. Tachys virgo LeConte, 1852 Ecology. Coastal lowlands. Saline places. Open ground; moist, muddy, saline soil. Biology. Seasonality: August. Dispersal power. Wing-condition unknown. Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet.

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Tachys vittiger LeConte, 1852 Ecology. Lowlands. Salt meadows. Open ground; moist, muddy, saline soil. Biology. Seasonality: May, August. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Collecting techniques. Pouring water over the ground; treading the soil with the feet. Tachyta angulata Casey, 1918 Ecology. Lowlands and mountains. Forests. Open ground. Nocturnal; sheltering during the day under the loose bark of fallen or standing, dead or dying trees (e.g., Populus, Acer, Betula, Picea, Pinus, Pseudotsuga). Biology. Seasonality: January-December. Tenerals: April (rarely; MB), August. Dispersal power. Macropterous. Occasional in drift material, which indicates flight ability. Moderate runner. Frequent climber (on trees). Collecting techniques. Lifting loose bark; using malaise traps. Reference. Lindroth, 1966 (natural history). Tachyta falli (Hayward, 1900) Ecology. Lowlands and mountains. Forests. Open ground. Nocturnal; sheltering during the day under the loose bark of fallen or standing, dead or dying trees (e.g., Pinus, Abies, Pseudotsuga). Biology. Seasonality: April-September, December. Tenerals: July-August. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Frequent climber (on trees). Collecting technique. Lifting loose bark. References. Lindroth, 1966 (natural history); Erwin, 1975 (natural history). Tachyta (Tachyta) nana inornata (Say, 1823) Ecology. Lowlands and mountains. Forests. Open ground. Mostly nocturnal; sheltering during the day under the loose bark of fallen or standing, dead or dying trees (e.g., Abies, Celtis, Picea, Pinus, Quercus, Ulmus). Biology. Seasonality: January-December. Tenerals: June, August. Adults found overwintering in wood debris (SC). Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Frequent climber (on trees). Collecting technique. Lifting loose bark. References. Kirk, 1969 (dispersal power) and 1970 (biology); Erwin, 1975 and 1981 (natural history). Tachyta (Tachyta) nana kirbyi Casey, 1918 Ecology. Lowlands and mountains. Forests. Open ground. Nocturnal; sheltering during the day under the loose bark of fallen or standing, dead or dying, trees (e.g., Pinus, Abies, Picea, Pseudotsuga). Biology. Seasonality: March-October. Tenerals: August (BC). Dispersal power. Macropterous, probably capable of flight. Moderate runner. Frequent climber (on trees). Collecting technique. Lifting loose bark. References. Lindroth, 1966 (natural history); Erwin, 1975 (natural history).

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Tachyta (Tachyta) parvicornis Notman, 1922 Ecology. Lowlands and mountains. Forests. Open ground. Nocturnal; sheltering during the day under the loose bark of fallen or standing, dead or dying trees (e.g., Pinus). Biology. Seasonality: January-December. Tenerals: April-July, September. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Frequent climber (on trees). Collecting technique. Lifting loose bark. Reference. Erwin, 1975 (natural history). Tanystoma cuyama Liebherr, 1985 Ecology. Lowlands and mountains. Open oak-grasslands and chaparrals. Biology. Seasonality: January-June, August, October-December. Dispersal power. Brachypterous, incapable of flight. Reference. Liebherr, 1985 (ecology). Tanystoma maculicolle (Dejean, 1828) Ecology. Lowlands and mountains. Mostly open grasslands. Marshlands: along rivers. Orchards (e.g., apple), forests, and human habitations. Mostly open ground; moist soil covered with some vegetation (e.g., Scirpus). Crepuscular and nocturnal; sheltering during the day under logs and the loose bark of trees (e.g., Eucalyptus). Gregarious (in winter). Biology. Seasonality: January-August, October-December. Tenerals: JanuaryJune, November-December. Adult food: lepidopterous caterpillars and meat (in captivity); stored products (in human habitations). Predators: woodpeckers (flickers). Defensemechanism: when alarmed, the adult emits a nasty smell (sometimes a nuisance in human habitations). Dispersal power. Wing-dimorphic: macropterous or brachypterous (incapable of flight). Frequent flier (mostly in June-July; swarming at dusk following rains in the fall; to artificial lights at night). Moderate runner. Strongly favored by human activities. Collecting techniques. Turning logs; lifting loose bark; light trapping; pitfall trapping. References. Essig, 1958 (biology, dispersal power); Liebherr, 1984a and 1985 (natural history); Liebherr and Hajek, 1986 (dispersal power). Tanystoma striatum (Dejean, 1828) Ecology. Lowlands and mountains. Grasslands. Open ground; moist soil covered with some vegetation (e.g., grass). Biology. Seasonality: January-December. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Reference. Liebherr, 1985 (ecology). Tanystoma sulcatum (Dejean, 1828) Ecology. Lowlands and mountains. Edge of coniferous forests; roadsides. Soil covered with dense vegetation (e.g., grass, Senecio). Nocturnal; sheltering during the day under stones. Biology. Seasonality: February-June, August, October-December. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Reference. Liebherr, 1985 (ecology).

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Technophilus croceicollis croceicollis (Ménétriés, 1843) Ecology. Lowlands and uplands. Margins of alkaline playas; salt flats. Open ground; dry, saline or alkaline soil composed of sand or mud cakes, covered with sparse vegetation (e.g., Salicornia). Diurnal. Biology. Seasonality: April-September, November. Tenerals: May-August (CA). Adults found overwintering in the soil (as deep as 15 cm) under Salicornia-mats. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Frequent climber (on low plants and shrubs). Collecting techniques. Pitfall trapping; sweeping or beating the vegetation; light trapping. Reference. Larson, 1969 (natural history). Technophilus croceicollis peigani Larson, 1969 Ecology. Bases of eroded coulees; alluvial fens. Open ground; dry, alkaline, clayish sandy soil covered with sparse vegetation (e.g., Bouteloua, Atriplex, Opuntia). Mostly diurnal; active in the bright sunshine on the ground or low plants; sheltering on cloudy or cool days in cracks in the soil and under debris. Biology. Seasonality: May-July. Copulating pairs: May-June (WY). Tenerals: July (WY). Larval food, in captivity: weevil larvae, lepidopterous caterpillars, sawflies, ant larvae and pupae, and mealworms. Dispersal power. Macropterous, probably capable of flight. Moderate runner. Frequent climber (on low plants). Collecting techniques. Pitfall trapping; sweeping the vegetation. Reference. Larson, 1969 (natural history). Technophilus pilatei Chaudoir, 1877 Ecology. Coastal lowlands. Borders of salt marshes close to the sea. Open ground; saline soil covered with some vegetation. Diurnal and nocturnal; active on plants during the day or sheltering under debris. Biology. Seasonality: June. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Moderate runner. Regular climber (on plants). Collecting techniques. Pitfall trapping; sweeping the vegetation. Reference. Larson, 1969 (ecology, dispersal power) Tetragonoderus (Tetragonoderus) fasciatus (Haldeman, 1843) Ecology. Lowlands. Shores of lakes, rivers, and brooks. Also abandoned fields, cultivated fields (e.g., corn, cotton), and sand pits. Not restricted to the vicinity of water. Open ground; dry sandy soil covered with sparse vegetation. Mostly diurnal; active in the sunshine; sheltering on cloudy or cool days at the base of clumps of grass and other plants, under dead leaves, stones, logs, and pieces of wood. Gregarious. Biology. Seasonality: February-November. Tenerals: August (VT). Adults found overwintering in leaf litter. Defense-mechanism: when alarmed, the adult escapes by running and flying readily like a tiny Cicindela; body color-pattern matching the soil (beetle visible only when moving). Dispersal power. Macropterous. Frequent flier (when alarmed; to artificial lights night). Occasional in lakeshore drift material, which indicates flight ability. Fast runner. Moderately favored by human activities. Collecting techniques. Hunting

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with a butterfly net; inspecting the base of clumps of grass; pitfall trapping; light trapping. References. Ulke, 1903 (ecology); Blatchley, 1910 (ecology, biology); Everly and Everly, 1936 (ecology); Lindroth, 1969a (natural history); Erwin, 1981 (natural history); Bousquet, 1997a (taxonomy). Tetragonoderus (Tetragonoderus) intersectus (Germar, 1824) Ecology. Lowlands. Cultivated fields (e.g., tobacco, peanut), pastures, abandoned fields, gardens, and roadsides. Open ground; dry, sandy soil, bare or covered with sparse low vegetation. Diurnal and nocturnal; active in the sunshine or sheltering in leaf litter, under hay and weeds. Biology. Seasonality: January-December. Adults found overwintering in dead leaves. Defense-mechanism: when alarmed, the adult escapes by running and flying readily like a tiny Cicindela; body color-pattern matching the soil. Dispersal power. Macropterous. Frequent flier (when alarmed; often swarming; to artificial lights at night). Fast runner. Strongly favored by human activities. Collecting techniques. Raking the leaf litter; pitfall trapping; light trapping. References. Leng, 1915 (ecology, dispersal power); Kirk, 1969 (biology, dispersal power) and 1970 (dispersal power). Tetragonoderus (Peronoscelis) latipennis LeConte, 1874 Ecology. Lowlands. River banks. Biology. Seasonality: August. Dispersal power. Macropterous, probably capable of flight. Fast runner. Tetragonoderus pallidus G.H. Horn, 1868 Ecology. Lowlands. Sand dunes. Not associated with bodies of water. Open ground; dry, sandy, bare or covered with sparse vegetation. Nocturnal; sheltering during the day in rodent burrows dug in the soil, under plant debris, and pieces of wood. Biology. Seasonaliy: January-December. Defense-mechanism: body color-pattern matching the soil. Dispersal power. Macropterous, probably capable of flight. Fast runner. Strong burrower. Collecting techniques. Inspecting rodent burrows; turning plant debris; pitfall trapping. Tetraleucus picticornis (Newman, 1844) Ecology. Lowlands and mountains. Low deciduous forests, flood-plain forests, and swamp-forests (e.g., Taxodium): often along lakes, ponds, rivers, and brooks. Shaded ground; wet, clayish or muddy soil, which thick leaf litter. Nocturnal; sheltering during the day in leaf litter (mostly) and under pieces of wood. Biology. Seasonality: JanuaryDecember. Adult and larval food, in captivity: mealworms. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous. Regular flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Raking the leaf litter; pitfall trapping; night collecting with a headlamp; light trapping. References. Ulke, 1903 (dispersal power); Leng, 1915 (ecology); Lindroth, 1966 (dispersal power); Liebherr, 1991a (natural history).

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Thalassotrechus barbarae (G.H. Horn, 1892) Ecology. Coastal lowlands. Upper intertidal zone of the seashore. Open ground; wet, saline soil consisting of rock or sand with rocky outcrops. Sheltering in intertidal rock crevices (e.g., below the high tide mark). Adults are active over stones and sandy soil at low tide. Gregarious. Associated with staphylinids (Diaulota, Liparopohalus). Larval habitat: similar to the adult, in cracked rocks and crevice sediments. Biology. Seasonality: March, May, July, October-November. Adult food, in the field: dipterous larvae, small crustaceans, and stranded zooplankton. Defense-mechanism: when alarmed, the adult escapes by running for shelter or by slowly emitting a discharge from the pygidial glands (at a range less shorter than its body length). Dispersal power. Brachypterous, incapable of flight. Moderate runner (larva slower than adult). Collecting techniques. Inspecting intertidal rock crevices at low tide; splitting open the cracked rocks with a crowbar; night collecting with a headlamp. References. Van Dyke, 1918 (ecology; as nigripennis Van Dyke); Moore, 1956 (natural history); Evans, 1970, 1976, 1977, 1982, 1983, 1985, and 1986 (natural history); Bousquet and Laplante, 1997 (taxonomy, ecology). Comments. Endangered by oil spills. Thalpius cubanus (Chaudoir, 1877) Ecology. Lowlands.Vicinity of a flooded pool. Nocturnal; sheltering during the day under flood debris. Biology. Seasonality: January-June. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Thalpius deceptor (Darlington, 1934) Ecology. Lowlands. In flood debris. Biology. Seasonality: June. Dispersal power. Macropterous, probably capable of flight. Thalpius dorsalis (Brullé, 1834) Ecology. Lowlands. An interdunal swale (VA). Biology. Seasonality: January-March, May-June, August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). References. Ulke, 1903 (dispersal power); Kirk, 1975c (dispersal power); Davidson, 1995 (ecology, dispersal power). Thalpius hoegei (Bates, 1883) Ecology. Lowlands. Biology. Unknown. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Thalpius horni (Chaudoir, 1872) Ecology. River banks. Moist soil. Biology. Seasonality: May-August; spring-fall (CA). Tenerals: late July (TX). Dispersal power. Macropterous. Occasional flier (to artificial lights at night).

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Thalpius microcephalus Van Dyke, 1926 Ecology. A hill, under a stone. Biology. Seasonality: January. Dispersal power. Macropterous, probably capable of flight. Thalpius pygmaeus (Dejean, 1826) Ecology. Lowlands. Shores of lakes and canals. Moist soil. Biology. Seasonality: April, June-July. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Reference. Davidson, 1995 (ecology, dispersal power). Thalpius rufulus (LeConte, 1851) Ecology. Moist situations. Nocturnal; sheltering during the day under stones. Biology. Seasonality: June-July. Parasites: Laboulbeniales (Fungi). Dispersal power. Macropterous, probably capable of flight. Collecting technique. Turning stones. Reference. LaBonte, 1996 (ecology). Trechoblemus westcotti Barr, 1972 Ecology. Unknown. Biology. Seasonality: July-August. Dispersal power. Macropterous. Occasional flier (to artificial lights at night). Trechus (Microtrechus) aduncus aduncus Barr, 1962 Ecology. Mountains. Forests: along seepages or small streams. Shaded ground; wet soil covered with thick leaf litter. Nocturnal; sheltering during the day in wet leaf litter between small stones. Biology. Seasonality: September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. Reference. Barr, 1979 (ecology). Trechus (Microtrechus) aduncus coweensis Barr, 1979 Ecology. Mountains. Forests: along seepages or small streams. Shaded ground; wet soil covered with thick leaf litter. Nocturnal; sheltering during the day in wet leaf litter between small stones. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. Trechus (Microtrechus) aduncus howellae Barr, 1979 Ecology. Mountains. Forests: along seepages or small streams. Shaded ground; wet soil covered with thick leaf litter. Nocturnal; sheltering during the day in wet leaf litter between small stones. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter.

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Trechus (Microtrechus) aduncus toxawayi Barr, 1979 Ecology. Mountains. Forests: along seepages or small streams. Shaded ground; wet soil covered with thick leaf litter. Nocturnal; sheltering during the day in wet leaf litter between small stones. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. Reference. Barr, 1979 (ecology). Trechus (Trechus) alinae Dajoz, 1990 Ecology. Subalpine zone. Coniferous forests (e.g., Pinus, Abies, Juniperus). Shaded ground; moist soil. Nocturnal; sheltering during the day. Biology. Seasonality: April. Tenerals: April. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Trechus (Trechus) apache Dajoz, 1990 Ecology. Mountains. A wooded canyon (Juniperus, Quercus), near a pool. Nocturnal; sheltering during the day under vegetal debris. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Trechus (Trechus) apicalis Motschulsky, 1845 Ecology. Lowlands, mountains, subalpine and alpine zones. Mostly forests (deciduous, mixed, coniferous): often along rivers, brooks, lakes, and ponds; forest clearings and edges, thickets (e.g., Alnus, Betula). Also meadows, moraines, grasslands, and vacant lots (coastal humid areas of NF, LB, northeastern QU). Sometimes drier parts of Carex-marshes and ski fields. Ground shaded by trees, shrubs or bushes; moderately moist, gravelly, sandy or peaty soil, mostly covered with thick leaf litter, sometimes covered with some vegetation. Nocturnal; sheltering during the day in leaf litter (mostly), also under stones, pieces of wood, in moss, heaps of hay, and pieces of cardboard. Biology. Seasonality: March-October. Gravids: July and early August (AB, QU). Tenerals: April, early June to late September; mostly in July. Defense-mechanism: when alarmed, the adult may emit a strong smell from the pygidial glands. Dispersal power. Wing-dimorphic: brachypterous (mostly; incapable of flight) or macropterous. Occasional flier (at sunset). Moderate runner. Moderately favored by human activities. Collecting techniques. Raking the leaf litter; pitfall trapping. References. Lindroth, 1955a, 1963a, and 1963b (natural history). Trechus (Trechus) arizonae Casey, 1918 Ecology. Mountains. Biology and Dispersal power. Unknown. Trechus (Microtrechus) balsamensis Barr, 1962 Ecology. Mountains. Coniferous forests (e.g., Picea, Abies): at the base of low cliffs. Nocturnal; sheltering during the day under moss carpets and in conifer-needle litter. Biolo-

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gy. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning moss carpets; raking conifer-needle litter. Reference. Barr, 1979 (ecology). Trechus (Microtrechus) barberi (Jeannel, 1931) Ecology. Mountains. Coniferous (e.g., Picea, Abies) and deciduous forests: along crests, slopes, and ravines. Shaded ground; moist soil covered with thin leaf litter and moss carpets. Nocturnal; sheltering during the day under stones. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning leaf litter. Reference. Barr, 1969 (ecology). Trechus (Microtrechus) bowlingi Barr, 1962 Ecology. Mountains. Coniferous forests (Picea, Abies). Shaded ground; moist soil covered with thin leaf litter and moss carpets. Nocturnal; sheltering during the day under moss carpets, leaf litter, and stones. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning moss carpets. Reference. Barr, 1979 (ecology). Trechus (Trechus) caliginis Barr, 1985 Ecology. Mountains. Macrohabitat unknown. Moist soil. Nocturnal. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Trechus (Trechus) carolinae Schaeffer, 1901 Ecology. Mountains. Coniferous forests (e.g., Picea, Abies): along seepages, small streams, slopes, and low cliffs. Shaded ground; wet soil covered with thick leaf litter. Nocturnal; sheltering during the day in leaf litter and moss. Biology. Seasonality: May, July-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; turning moss carpets. Reference. Barr, 1979 (ecology). Trechus (Trechus) chalybeus Dejean, 1831 Ecology. Lowlands, mountains, subalpine and alpine zones. Coniferous (e.g., Pinus) and decidous forests: along streams, lakes, and ponds; thickets (e.g., Salix, Alnus). Grasslands (coastal humid areas). Margins of snow-patches. Shaded (mostly) or open ground; moderately moist soil covered with thick leaf litter or moss. Nocturnal; sheltering during the day in leaf litter and moss, under stones and pieces of wood. Biology. Seasonality: May-August, November. Tenerals: June, August. Adult food, in captivity: dipterous eggs. Parasites: Laboulbeniales (Fungi). Defense-mechanism: when alarmed, the adult may emit a nasty smell from the pygidial glands. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; turn-

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ing moss carpets. References. Lindroth, 1961 (natural history); Kavanaugh, 1992 (natural history). Trechus (Trechus) coloradensis Schaeffer, 1915 Ecology. Mountains, subalpine and alpine zones. Vicinity of cold streams and snow fields. Moist soil. Nocturnal; sheltering during the day under stones. Biology. Seasonality: April-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning stones. Trechus (Trechus) conformis Jeannel, 1927 Ecology. Coastal lowland. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Trechus (Trechus) crassiscapus Lindroth, 1955 Ecology. Lowlands, mountains, subalpine and alpine zones. Small swamp-forests, swamps, low forests, bogs, and marshes: along rivers, lakes, and ponds. Also forest clearings and alpine meadows. Ground somewhat shaded by Alnus- or Betula-bushes; wet, clayish, muddy or peaty soil covered with thick leaf litter and moss carpets. Nocturnal; sheltering during the day mostly among dead leaves and moss, rarely under stones. Biology. Seasonality: May-October. Gravids: early July (QU). Tenerals: August (NF, VT, QU). Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; turning moss carpets. References. Lindroth, 1955a, 1961, and 1963a (natural history). Trechus (Trechus) cumberlandus Barr, 1962 Ecology. Lowlands and mountains. Forests: along deep ravines, sinkholes, and cold springs. Shaded ground; moist soil. Caves (occasionally). Nocturnal; sheltering during the day. Biology. Seasonality: October. Dispersal power. Brachypterous, incapable of flight. Moderate runner. References. Barr, 1969 and 1979 (ecology). Trechus (Trechus) dietriechi Barr, 1962 Ecology. Mountains. Forests. Moist soil. Nocturnal; sheltering during the day. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Trechus (Microtrechus) haoe Barr, 1979 Ecology. Mountains. A forest, on a large boulder. Shaded ground; moist soil covered with thin leaf litter and moss carpets. Nocturnal; sheltering during the day under moss carpets. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight.

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Trechus (Trechus) humboldti Van Dyke, 1945 Ecology. A dense coniferous forest (Sequoia). Biology. Seasonality: July. Dispersal power. Unknown. Trechus (Trechus) hydropicus avus Barr, 1962 Ecology. Mountains. Coniferous forests (Picea, Abies). Shaded ground; moist soil covered with thin leaf litter and moss carpets. Biology. Seasonality: June, August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; turning moss carpets. Trechus (Trechus) hydropicus beutenmuelleri Jeannel, 1931 Ecology. Mountains. Coniferous forests (Picea, Abies): along ravines and seepages. Shaded ground; moist soil covered with thin leaf litter and moss carpets. Nocturnal; sheltering during the day under stones, under wood chips, under and inside rotten logs. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning stones and wood chips. Reference. Barr, 1969 (ecology). Trechus (Trechus) hydropicus canus Barr, 1962 Ecology. Mountains. Coniferous forests (Picea, Abies). Shaded ground; moist soil covered with thin leaf litter and moss carpets. Nocturnal; sheltering during the day. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Trechus (Trechus) hydropicus hydropicus G.H. Horn, 1883 Ecology. Mountains. Forests: along slopes. Shaded ground; moist soil covered with thin leaf litter and moss carpets. Nocturnal; sheltering during the day among dead leaves and moss. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; turning moss carpets. Trechus (Microtrechus) inexpectatus Barr, 1985 Ecology. Mountains. Forests: along seepages or small streams. Shaded ground; wet soil covered with thick leaf litter. Nocturnal; sheltering during the day in wet leaf litter between small stones. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. Trechus (Microtrechus) luculentus luculentus Barr, 1962 Ecology. Mountains. Deciduous forests: along streams, seepages, and small ravines. Shaded ground; wet, silty soil covered with thick leaf litter. Nocturnal; sheltering during the day under stones (mostly), in wet dead leaves, sticks, and moss carpets. Biology.

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Seasonality: May-July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning stones. Reference. Barr, 1979 (ecology). Trechus (Microtrechus) luculentus unicoi Barr, 1979 Ecology. Mountains. Deciduous forests: along ravines, seepages, and springs. Shaded ground; wet soil. Nocturnal; sheltering during the day under stones and moss. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning stones and moss carpets. Reference. Barr, 1979 (ecology). Trechus (Microtrechus) luculentus wayahensis Barr, 1979 Ecology. Mountains. Macrohabitat unknown. Moist soil. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Reference. Barr, 1979 (ecology). Trechus (Trechus) mitchellensis Barr, 1962 Ecology. Mountains. Coniferous forests (e.g., Picea, Abies). Shaded ground; moist soil covered with thin leaf litter and moss carpets. Nocturnal; sheltering during the day under moss carpets. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning moss carpets. Reference. Barr, 1979 (ecology). Trechus (Microtrechus) nantahalae Barr, 1979 Ecology. Mountains. Forests: along seepages and small streams. Shaded ground; wet soil covered with thick leaf litter. Nocturnal; sheltering during the day in leaf litter between stones. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. Trechus (Microtrechus) nebulosus Barr, 1962 Ecology. Mountains. Forests: along cliffs and boulders. Shaded ground; moist soil covered with thin leaf litter and moss carpets. Nocturnal; sheltering during the day under moss carpets. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning moss carpets. Reference. Barr, 1979 (ecology). Trechus (Microtrechus) novaculosus Barr, 1962 Ecology. Mountains. Coniferous forests (Picea, Abies): along seepages and small streams. Shaded ground; wet soil covered with thick leaf litter. Nocturnal; sheltering during the day in leaf litter between stones. Biology. Seasonality: May, August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. Reference. Barr, 1979 (ecology).

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Trechus (Trechus) obtusus Erichson, 1837 Ecology. Lowlands. Cultivated fields, wastelands, roadsides, gravel pits, city lawns, gardens, parks, orchards (e.g., apple), and forest edges. Greenhouses (occasionally). Vicinity of urban and rural sites. Open ground; moderately moist, sandy, loamy or silty soil covered with sparse vegetation. Nocturnal; sheltering during the day in leaf litter and under stones. Biology. Seasonality: March-December. Adult food, in the field and in captivity: dipterous eggs. Defense-mechanism: when alarmed, the adult may emit a discharge from the pygidial glands (at a range shorter than its body length). Dispersal power. Wing-dimorphic: macropterous (mostly) or brachypterous (incapable of flight). Occasional flier (to artificial lights at night). Moderate runner. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Raking the leaf litter; turning stones; pitfall trapping. References. Lindroth, 1945, 1961, and 1985 (natural history); Kavanaugh and Erwin, 1985 (natural history); Koch, 1989 (ecology); Kavanaugh, 1992 (natural history); Luff, 1993 (biology). Trechus (Trechus) oregonensis Hatch, 1951 Ecology. Lowlands and mountains. Mature mixed forests. Somewhat shaded ground; slightly moist soil covered with thick leaf litter and humus. Nocturnal; sheltering during the day in leaf litter. Biology. Seasonality: May-August. Tenerals: late July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. Reference. Lindroth, 1963b (natural history). Trechus (Trechus) ovipennis Motschulsky, 1845 Ecology. Coastal lowlands. Sea cliffs, sea beaches, and borders of adjoining streams. Caves (occasionally). Shaded ground; wet, sandy or gravelly soil. Nocturnal; sheltering during the day in soil crevices, under driftwood, dead algae, stones, among gravel, and at the base of plants. Biology. Seasonality: May, July-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning driftwood, dead algae, and stones; raking the gravel. References. Hatch, 1953 (ecology); Kavanaugh, 1992 (natural history). Trechus (Microtrechus) pisgahensis Barr, 1979 Ecology. Mountains. Coniferous forests: along seepages and small streams. Shaded ground; wet soil covered with thick leaf litter and moss carpets. Nocturnal; sheltering during the day in leaf litter and under moss carpets growing on boulders. Biology. Seasonality: March, May-August. Adults found overwintering in moss growing on boulders. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; turning moss carpets. Trechus (Trechus) pomonae Fall, 1901 Ecology. Mountains. Wooded canyons. Biology. Seasonality: January, March, October. Dispersal power. Wing-condition unknown. Moderate runner.

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Trechus (Trechus) quadristriatus (Schrank, 1781) Ecology. Lowlands. Cultivated fields (e.g., onion, corn), field edges, gardens, and vacant lots. Open (mostly) or shaded ground; rather dry, sandy, gravelly or loamy soil covered with sparse vegetation. Diurnal and nocturnal; active in the shade of plants or sheltering under dead leaves, stones, and other debris. Biology. Seasonality: May-September. Tenerals: early July (MI). Adults found overwintering in the soil. Adult food, in the field (Europe): insect eggs (e.g., flies), scarabaeid larvae. Larval food, in the field (Europe): scarabaeid larvae, worms, and small molluscs. Economic importance (Europe): may play an important role in the control of some insect pests (e.g., the Cabbage Root Fly). Predators: toads. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Moderate runner. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Raking the leaf litter; turning stones; pitfall trapping; light trapping. References. Lindroth, 1945 (natural history); Mitchell, 1963a (biology) and 1963b (natural history); Sharova, 1981 (ecology, biology); Bousquet et al. 1984 (ecology, dispersal power); Lindroth, 1985 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology). Trechus (Trechus) roanicus Barr, 1962 Ecology. Mountains. Coniferous forests: along seepages and small streams (e.g., on northfacing cliffs). Shaded ground; wet soil covered with moss carpets and leaf litter. Nocturnal; sheltering during the day under moss carpets. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning moss carpets. Reference. Barr, 1979 (ecology). Trechus (Microtrechus) rosenbergi Barr, 1962 Ecology. Mountains. Coniferous forests (Picea, Abies): along seepages, small streams, and low cliffs. Shaded ground; wet soil covered with thick leaf litter and moss carpets. Nocturnal; sheltering during the day in conifer-needle litter. Biology. Seasonality: JulyAugust. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the conifer-needle litter. Trechus (Trechus) rubens (Fabricius, 1801) Ecology. Lowlands, mountains, subalpine and alpine zones. Borders of lakes, ponds, pools, and marshes; swamps; borders of slow rivers, brooks, and roadside ditches; low woods and beaver houses. Open (mostly) or shaded ground; wet or moist, clayish (mostly) or peaty soil, often rich in organic debris, covered with dense vegetation (e.g., Carex, grass). Crepuscular and nocturnal; sheltering during the day in humus, under wellembedded pieces of wood, stones and in cracks in the soil, also in leaf litter, moss, and under pieces of cardboard. Biology. Seasonality: May-October. Gravids: late July and early August (QU). Tenerals: late June (NB, QU), August (QU). Dispersal power. Macropterous. Frequent flier (often swarming; at dusk; to artificial lights at night). Occa-

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sional in sea- and lakeshore drift material, which indicates flight ability. Fast runner. Strong burrower. Good swimmer. Clearly effective as a colonist. Strongly favored by human activities. Collecting techniques. Turning embedded pieces of wood and stones; treading the soil with the feet; pouring water over the ground; light trapping; inspecting the walls of beaver houses. References. Lindroth, 1945, 1961, and 1963b (natural history); Mitchell, 1963a and 1963b (biology); Sharova, 1981 (ecology, biology); Lindroth, 1985 (natural history); Koch, 1989 (ecology); Luff, 1993 (biology). Trechus (Microtrechus) satanicus Barr, 1962 Ecology. Mountains. Open thickets (blackberry): along seepages and small streams. Shaded ground; wet soil covered with thick leaf litter and moss carpets. Nocturnal; sheltering during the day in leaf litter and moss in contact with rock surface. Biology. Seasonality: May, July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; turning moss carpets. Trechus (Trechus) schwarzi saludae Barr, 1979 Ecology. Mountains. Forests: along cool ravines near small spring-fed streams. Shaded ground; wet soil. Nocturnal; sheltering during the day under moss and liverworts growing on stones. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning moss- and liverwort carpets. Reference. Barr, 1979 (ecology). Trechus (Trechus) schwarzi schwarzi Jeannel, 1931 Ecology. Mountains. Forests: along cool ravines near seepages and small streams. Shaded ground; wet soil covered with thick leaf litter. Nocturnal; sheltering during the day in leaf litter between stones. Biology. Seasonality: September. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. Trechus (Trechus) schwarzi scopulosus Barr, 1979 Ecology. Mountains. Forests: along cool ravines near seepages and small streams. Shaded ground; wet soil covered with thick leaf litter. Nocturnal; sheltering during the day in leaf litter between stones. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. Trechus (Microtrechus) stupkai Barr, 1979 Ecology. Mountains. Forests. Shaded ground; moist soil. Nocturnal; sheltering during the day in moss growing on stones. Biology. Seasonality: May. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning moss carpets.

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Trechus (Microtrechus) subtilis Barr, 1962 Ecology. Mountains. Forests: along slopes. Wet soil covered with thick leaf litter. Nocturnal; sheltering during the day in leaf litter between stones. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. Trechus (Microtrechus) talequah Barr, 1962 Ecology. Mountains. Forests: along slopes. Wet soil covered with thick leaf litter. Nocturnal; sheltering during the day in leaf litter between stones. Biology. Seasonality: September-August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Raking the leaf litter. Trechus (Microtrechus) tennessensis tauricus Barr, 1962 Ecology. Lowlands. A big sink surrounding a cave mouth. Moist soil. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Trechus (Microtrechus) tennessensis tennessensis Barr, 1962 Ecology. Lowlands. A big sink surrounding a cave mouth. Moist soil. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Trechus (Trechus) tenuiscapus Lindroth, 1961 Ecology. Mountains. Mixed forests. Moderately shaded ground; rather moist soil covered with thick humus layer. Nocturnal; sheltering during the day under stones and logs. Biology. Seasonality: May-August. Tenerals: late July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning stones and logs. Reference. Lindroth, 1961 (natural history). Trechus (Microtrechus) tonitru Barr, 1962 Ecology. Mountains. A heath thicket. Moist soil. Nocturnal; sheltering during the day in moss. Biology. Seasonality: July. Dispersal power. Brachypterous, incapable of flight. Trechus (Microtrechus) tuckaleechee Barr, 1962 Ecology. Caves. Moist soil. Biology. Seasonality: April, August. Dispersal power. Brachypterous, incapable of flight. Fast runner.

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Trechus (Microtrechus) tusquitee Barr, 1979 Ecology. Mountains. Forests: summit slopes and cliffs. Shaded ground; moist or wet soil covered with thin leaf litter and moss carpets. Nocturnal; sheltering during the day under moss carpets, dead leaves, and stones. Biology. Seasonality: August. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Turning moss carpets; raking the leaf litter. Reference. Barr, 1979 (ecology). Trechus (Microtrechus) uncifer Barr, 1962 Ecology. Mountains. Forests (coniferous, deciduous): along seepages and small streams. Shaded ground; wet soil covered with thick leaf litter. Nocturnal; sheltering during the day under stones. Biology. Seasonality: June-July. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning stones. Reference. Barr, 1979 (ecology). Trechus (Microtrechus) valentinei Barr, 1979 Ecology. Mountains. Coniferous forests (Picea, Abies): along streams, springs, and shallow ravines. Shaded ground; wet soil covered with thick leaf litter. Nocturnal; sheltering during the day under stones. Biology. Unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting technique. Turning stones. Reference. Barr, 1979 (ecology). Trechus (Microtrechus) vandykei (Jeannel, 1927) Ecology. Mountains. Coniferous forests (Picea, Abies). Shaded ground; moist soil covered with thin leaf litter and moss carpets. Nocturnal; sheltering during the day in leaf litter and under moss carpets. Biology. Seasonality: unknown. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Collecting techniques. Raking the leaf litter; turning moss carpets. References. Barr, 1969 and 1979 (ecology). Trechus (Microtrechus) verus Barr, 1962 Ecology. Mountains. Forests: along seepages and small streams. Shaded ground; wet soil covered with thick leaf litter. Nocturnal; sheltering during the day in deep moss, conifer-needle litter, and under well-embedded stones. Biology. Seasonality: June. Dispersal power. Brachypterous, incapable of flight. Moderate runner. Strong burrower. Collecting techniques. Turning moss carpets; raking the conifer-needle litter. Reference. Barr, 1979 (ecology). Trichocellus (Trichocellus) cognatus (Gyllenhal, 1827), see Dicheirotrichus (Trichocellus) cognatus (Gyllenhal, 1827). References. Kryzhanovskij et al. 1995 (taxonomy); Ball and Bousquet, 2001 (taxonomy).

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Trichocellus (Oreoxenus) mannerheimii (R.F. Sahlberg, 1844), see Dicheirotrichus (Oreoxenus) mannerheimii (R.F. Sahlberg, 1844). References. Kryzhanovskij et al. 1995 (taxonomy); Ball and Bousquet, 2001 (taxonomy). Trichotichnus (Iridessus) autumnalis (Say, 1823) Ecology. Lowlands and mountains. Flood-plain forests (deciduous; e.g., Liriodendron, Liquidambar); upland deciduous forests (e.g., Quercus). Shaded ground; wet or moderately moist soil covered with leaf litter. Nocturnal; sheltering during the day in leaf litter (mostly) and under silt cakes. Biology. Seasonality: January-November. Gravids: late September (QU). Tenerals: September and October (MI, NH). Adults found overwintering in woods; among debris. Predators: ducks. Dispersal power. Macropterous. Frequent in lake- and seashore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Raking the leaf litter; pitfall trapping; turning drift material; using sticky boards. References. Blatchley, 1910 (ecology); Lindroth, 1968 (biology, dispersal power); Kirk, 1969 (biology); Erwin, 1981 (natural history); Anderson et al. 1995 (ecology, biology); Ball and Bousquet, 2001 (taxonomy). Trichotichnus (Trichochichnus) dichrous (Dejean, 1829) Ecology. Lowlands and mountains. Deciduous forests. Shaded ground; wet or dry soil. Nocturnal; sheltering during the day under logs, stones, and in leaf litter. Biology. Seasonality: March-November. Tenerals: mid-June (IN, NY). Adult food, in the field: insects. Predators: frogs and starlings. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Turning logs and stones; pitfall trapping; light trapping. References. Blatchley, 1910 (ecology); Lindroth, 1968 (dispersal power); Erwin, 1981 (natural history). Trichotichnus (Iridessus) fulgens (Csiki, 1932). Ecology. Lowlands and mountains. Open deciduous forests, pastures, cultivated fields (e.g., alfalfa), old fields, grasslands, vacant lots, and roadsides. Open or shaded ground; moderately dry, sandy or gravelly soil covered with dead leaves or grass. Mostly diurnal; active in the sunshine; sheltering on cloudy and cool days under dead leaves (mostly) and plants, stones, and other debris. Biology. Seasonality: January-December. Tenerals: January, March, May-July, September-October. Adults found overwintering in woods; in leaf litter, under dead fallen plants, among grass roots, and in the soil. Dispersal power. Macropterous. Frequent flier (in the sunshine; to artificial lights at night). Frequent in sea- and lakeshore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Collecting techniques. Raking the leaf litter; pitfall trapping; light trapping; turning drift material; sweeping plants. References. Erwin, 1981 (natural history); Noonan, 1991 (natural history); Anderson et al. 1995 (ecology); Ball and Bousquet, 2001 (taxonomy).

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Trichotichnus (Trichochichnus) vulpeculus (Say, 1823) Ecology. Lowlands and mountains. Low forests and flood-plain forests (deciduous, mixed, coniferous): along rivers, brooks, pools, and ravines. Shaded ground; wet, clayish or muddy soil covered with thick leaf litter. Nocturnal; sheltering during the day under the loose bark of logs and fallen trees (mostly), under logs, stones, fallen bark, boards, and in leaf litter. Biology. Seasonality: March-November. Gravids: September (QU). Tenerals: late May (rarely; QU), early July (WV), September (MD). Predators: frogs. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional in lakeshore drift material, which indicates flight ability. Moderate runner. Collecting techniques. Lifting loose bark; pitfall trapping; light trapping. References. Lindroth, 1968 (dispersal power); Larochelle, 1974a (ecology); Erwin, 1981 (natural history); Will et al. 1995 (dispersal power) Troglanillus valentinei Jeannel, 1963, see Anillinus valentinei (Jeannel, 1963). Reference. Barr, 1995 (taxonomy). Xenotrechus condei Barr and Krekeler, 1967 Ecology. A cave: along a stream. Dry or moist soil. Sheltering under stones. Biology. Seasonality: May-July. Dispersal power. Brachypterous, incapable of flight. Fast runner. Xenotrechus denticollis Barr and Krekeler, 1967 Ecology. Caves: along streams. Wet soil. Sheltering under stones or occurring near castings of oligochaetes. Biology. Seasonality: March, December. Parasites: fungi. Dispersal power. Brachypterous, incapable of flight. Fast runner. Xestonotus lugubris (Dejean, 1829) Ecology. Lowlands and mountains. Light deciduous forests (mostly); their clearings, roadsides, and edges; adjoining fields (e.g., corn), sand pits, and sand dunes. Also hedges and beaver houses. Shaded ground; moderately moist, gravelly, stony or muddy soil, rich in humus, covered with thick leaf litter. Nocturnal; sheltering during the day mostly in thick leaf litter and under well-embedded stones, also in the walls of beaver houses. Gregarious (in winter; in beaver houses). Biology. Seasonality: March-December. Copulating pairs: mid-May (QU); coitus lasting 13 minutes. Gravids: mid-June and early July (QU). Tenerals: April-May, July-September. Adults found overwintering on higher and drier ground, in woods, at the edge of woods, on hills, and in vacant lots; mostly in the soil under well-embedded stones, also under logs, dead leaves, and the loose bark of fallen trees and logs. Adult and larval food, in captivity: mealworms. Predators: bluebirds, starlings, and thrushes. Defense-mechanism: when alarmed, the adult escapes by running and digging into the leaf litter. Dispersal power. Macropterous. Occasional

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flier (to artificial lights in captivity). Occasional in seashore drift material, which indicates flight ability. Moderate runner. Occasional climber (on plants). Strong burrower. Good swimmer. Collecting techniques. Raking the leaf litter; turning embedded stones; pitfall trapping; sweeping plants. References. Larochelle, 1971a (ecology); Kirk, 1975c (ecology). Zacotus matthewsii LeConte, 1869 Ecology. Lowlands and mountains. Coniferous forests (e.g., Abies, Thuja) and their edges. Shaded ground; moderately moist soil, often covered with thick layer of conifer needles. Nocturnal; sheltering during the day under rotten logs (often) and in needle litter. Biology. Seasonality: February, May-October. Tenerals: mid-August (ID, WA). Predators: owls. Dispersal power. Brachypterous, incapable of flight. Slow runner. Strong burrower. Collecting techniques. Pitfall trapping; baiting traps with sugar; turning logs; raking the needle litter. References. Hatch, 1944 (ecology); Lindroth, 1961 (natural history); Johnson et al. 1966 (biology); Kavanaugh, 1992 (natural history). Zuphium americanum Dejean, 1831 Ecology. Lowlands and uplands. Mostly lake shores and freshwater streams flowing into the sea. Also wet meadows and pine barrens. Moist, clayish or muddy soil. Nocturnal; sheltering during the day under flat stones and in vegetal debris. Biology. Seasonality: January-November. Adults found overwintering in wood debris and tree hollows. Defense-mechanism: when alarmed, the adult escapes by running. Dispersal power. Macropterous. Frequent flier (to artificial lights at night). Occasional climber (on trees). Fast runner. Collecting techniques. Turning stones and vegetal debris; light trapping. References. Blatchley, 1910 (ecology, biology); Malkin, 1943 (ecology); Kirk, 1969 (biology, dispersal power) and 1975c (dispersal power); Erwin, 1981 (natural history); Davidson, 1995 (dispersal power). Zuphium delectum Liebke, 1933 Ecology. Lowlands. Biology. Seasonality: December. Dispersal power. Macropterous, probably capable of flight. Zuphium longicolle LeConte, 1879 Ecology. Lowlands. A desert sand dune. Biology. Seasonality: February, April-October. Dispersal power. Macropterous, probably capable of flight. Zuphium magnum Schaeffer, 1910 Ecology. Lowlands. Biology. Seasonality: January, May-June, October. Dispersal power. Macropterous, probably capable of flight.

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Zuphium mexicanum Chaudoir, 1863 Ecology. Unknown. Biology. Seasonality: March, June. Dispersal power. Macropterous, probably capable of flight. Zuphium pseudamericanum Mateu, 1981 Ecology. Unknown. Biology. Seasonality: January, March-July, November. Dispersal power. Macropterous, probably capable of flight.

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Bibliography

(This section includes references consulted or referred to in the text .) Alcock, J. 1976. The behaviour of the seed-collecting larvae of a carabid beetle (Coleoptera). Journal of Natural History 10: 367-375. Alexander, R.D. 1959. The courtship and copulation of Pasimachus punctulatus Haldeman (Coleoptera: Carabidae). Annals of the Entomological Society of America 52: 485. Allen, R.T. 1972. A revision of the genus Loxandrus LeConte (Coleoptera: Carabidae) in North America. Entomologica Americana (New Series) 46: 1-184. Allen, R.T. 1979. The occurrence and importance of ground beetles in agricultural and surrounding habitats. pp. 485-505 in Erwin, T.L., Ball, G.E. and D.R. Whitehead (Editors), Carabid beetles, their evolution, natural history, and classification. Dr. W. Junk Publishers, The Hague, Boston and London. Allen, R.T. and G.E. Ball. 1980. Synopsis of Mexican taxa of the Loxandrus series (Coleoptera: Carabidae: Pterostichini). Transactions of the American Entomological Society 105: 481-576. Allen, R. T. and R.G. Thompson. 1977. Faunal composition and seasonal activity of Carabidae (Insecta: Coleoptera) in three different woodland communities in Arkansas. Annals of the Entomological Society of America 70: 31-34. Andersen, J. 1968. The larva of Miscodera arctica Payk. (Col. Carabidae). Norsk entomologisk Tidsskrift 15: 71-74. Anderson, J.M., Mitchell, J.C., Hall, A.A. and R.L. Hoffman. 1995. Ground beetles (Coleoptera: Carabidae) from Quantico Marine Corps Base, Virginia. Banisteria 6: 3-16. Attygalle, A.B., Meinwald, J. and T. Eisner. 1992. Defensive secretion of a carabid beetle, Helluomorphoides clairvillei. Journal of Chemical Ecology 18: 489-498. Attygalle, A.B., Meinwald, J., Liebherr, J.K. and T. Eisner. 1991. Sexual dimorphism in the defensive secretion of a carabid beetle. Experientia 47: 296-299. Bakken, H. 1985. Cold hardiness in the alpine Patrobus septentrionis and Calathus melanocephalus. Journal of Insect Physiology 31: 447-453. Ball, G.E. 1959. A taxonomic study of the North American Licinini with notes on the Old World species of the genus Diplocheila Brullé (Coleoptera). Memoirs of the American Entomological Society 16: 1-258. Ball, G.H. 1962. Descriptions of eleven species of the beetle subgenus Cryobius (Pterostichus, Col. Carabidae) from Alaska and northwestern Canada. Opuscula Entomologica 28: 1-26. Ball, G.E. 1966. A revision of the North American species of the subgenus Cryobius Chaudoir (Pterostichus, Carabidae, Coleoptera). Opuscula Entomologica Supplementum 28: 1-166.

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Index

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Index

aabaaba Brachinus (Neobrachinus) 149 abbreviatus Cymindis (Pinacodera) 223 abbreviatus Dyschiriodes (Dyschiriodes) 243 abdita Lebia (Lebia) 296 abdominalis Lebia (Lebia) 296 abjectus Euproctinus (Neoeuproctus) 268 abstinens Stenolophus (Stenolophus) 483 accelerans Loxandrus 311 acherontis Pseudanophthalmus 382 acticola Bembidion 92 actiosus Harpalus (Pseudoophonus) 273 actuosum Bembidion (Bembidion debilicolle) 92 actuosus Apristus 81 acuta acuta Nebria (Reductonebria) 325 acuta quileute Nebria (Reductonebria) 325 acuta sonorae Nebria (Reductonebria) 325 acutesculptus Elliptoleus 267 acutifrons Bembidion (Trepanedoris) 93 acutipes acutipes Pterostichus (Cylindrocharis) 405 acutipes kentuckensis Pterostichus (Cylindrocharis) 406 adductum Bembidion (Bembidion) 93 adjunctus Selenophorus (Celiamorphus) 470 adoxus Pterostichus (Hypherpes) 406 adstrictus Pterostichus (Bothriopterus) 406 aduncus aduncus Trechus (Microtrechus) 501 aduncus coweensis Trechus (Microtrechus) 501 aduncus howellae Trechus (Microtrechus) 501 aduncus Loxandrus 311 aduncus toxawayi Trechus (Microtrechus) 502 adustipennis Brachinus (Neobrachinus) 149 advena Calathus (Procalathus) 169 aenea Amara (Amara) 39

aeneicolle Bembidion (Notaphus) 93 aeneicollis Scaphinotus (Steniridia) 448 aeneipennis Paratachys 357 aeneolus Anchomenus 66 aeneolus Dyschiriodes (Dyschiriodes) (Dyschirius frigidus) 244 aeneopiceus Selenophorus (Selenophorus) 470 aeneopolita Amara (Amara) 40 aenescens Micratopus 320 aeneum Cylindronotum 223 aeneus Notiophilus 343 aenulum Bembidion (Odontium) 93 aequalis Discoderus (Discoderus) 240 aequalis Sphaeroderus stenostomus 480 aequinoctialis Galerita (Galerita) 269 aerata Coptodera 212 aeruginosum Agonum (Circinalidia) 17 aestivus Chlaenius (Chlaenius) 191 affabilis Anillodes 69 affine Agonum (Agonum) 18 affine Bembidion (Furcacampa) 93 affine Calosoma (Chrysostigma) 175 affinis Dyschiriodes (Paradyschirius) 244 affinis Harpalus (Harpalus) 273 agilis Loxandrus 312 agilis Platynus (Microplatynus) 370 agitatus Apristus 81 agonus Pterostichus (Lenapterus) 407 agricola Anisodactylus (Anisodactylus) 69 alabamae Cyclotrachelus (Evarthrus) 214 alabamae Pseudanophthalmus 382 alabamensis Cyclotrachelus (Cyclotrachelus) 214 alamedae Pterostichus (Hypherpes) 407

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André Larochelle & Marie-Claude Larivière

alaskanum Asaphidion 84 alaskense Bembidion (Bracteon) 94 alaskensis Harpalus vittatus 292 albamontana Rhadine 443 albicrus Agonum (Agonum) 18 albipes Paranchus 357 albipes Paratachys 357 alexiguus Brachinus (Neobrachinus) 150 algidus Loxandrus 312 algidus Pterostichus (Hypherpes) 407 alinae Trechus (Trechus) 502 alleni Pseudomorpha 403 alpina Amara (Curtonotus) 40 alpineanum Bembidion (Hirmoplataphus) 94 alternans Acupalpus (Philodes) 13 alternans alternans Dicaelus (Dicaelus) 229 alternans Anisodactylus (Anadaptus) 70 alternans Anisodactylus (Anadaptus) 70 alternans Brachinus (Neobrachinus) 150 alternans Cyclotrachelus (Evarthrus) 215 alternans subtropicus Dicaelus (Dicaelus) 229 alternatus Chlaenius (Agostenus) 192 alternatus Hartonymus 293 alticola Dyschiriodes (Dyschiriodes) 244 altisierrae Nebria (Nipponebria) 325 altisierrae Nipponebria (Vancouveria) 340 alutacea Pseudomorpha 403 alutaceus Oodinus 352 amandus amandus Plochionus (Menidius) 375 amandus discoideus Plochionus (Menidius) 375 amandus vittatus Plochionus (Menidius) 375 amaroides Anisodactylus (Aplocentroides) 70 amaroides Oodes 351 ambiguus Dicaelus (Paradicaelus) 230 americana Clivina 205 americanum Bembidion (Ochthedromus) 94 americanum Omophron 348 americanum Zuphium 514 americanus americanus Elaphrus (Elaphrus) 262 americanus Brachinus (Neobrachinus) 150 americanus Cymindis (Cymindis) 223 americanus Oodes 351 americanus Oodes americanus 351 americanus sylvanus Elaphrus (Elaphrus) 262 americanus Syntomus 493

amethystinus Pterostichus (Hypherpes) 407 amoena Diacheila arctica 229 amoenus Chlaenius (Anomoglossus) 192 amoenus Discoderus (Discoderus) 240 amphibius Schizogenius (Schizogenius) 465 ampliatus Cymindis (Pinacodera) 223 ampliceps Bembidion (Trepanedoris) 94 amputatus amputatus Harpalus (Harpalomerus) 274 analis Clivina 205 analis Dyschiriodes (Paradyschirius) 244 analis Lebia (Lebia) 297 anceps Elaphropus 256 anceps Stenolophus (Stenolophus) 483 anchomenoides Agonum (Europhilus) 18 andrewsii amplicollis Scaphinotus (Steniridia) 448 andrewsii andrewsiiScaphinotus (Steniridia) 449 andrewsii barksdaleiScaphinotus (Steniridia) 449 andrewsii darlingtoni Scaphinotus (Steniridia) 449 andrewsii germari Scaphinotus (Steniridia) 449 andrewsii montanusScaphinotus (Steniridia) 449 andrewsii mutabilis Scaphinotus (Steniridia) 450 andrewsii nantahalae Scaphinotus (Steniridia) 450 andrewsii parvitarsalis Scaphinotus (Steniridia) 450 andrewsii saludaeScaphinotus (Steniridia) 450 andrewsii waldensius Scaphinotus (Steniridia) 450 angulata Tachyta 496 angulatum Calosoma (Carabosoma) 176 angulatus Dicheirus dilatatus 235 angulatus Scaphinotus (Neocychrus) 450 anguliferum Bembidion (Trepanedoris) 94 angustata Amara (Zezea) 41 angustata Pseudomorpha 403 angustatoides Amara (Zezea) 41 angustatus angustatus Batenus (Platynidius) 91 angustatus Apenes (Apenes) 80 angustatus hypolithos Batenus (Platynidius) 91 angustatus Platynus 370 angusticollis angusticollis Elaphrus (Elaphroterus) 262 angusticollis Scaphinotus (Stenocantharus) 451 angustus Pterostichus (Hypherpes) 408 animosus Harpalus (Euharpalops) 275 annosus Pterostichus (Hypherpes) 408

Index anthicoides Rhadine 443 anthobia Amara (Amara) 41 anthracinum Agonum (Agonum) 19 anthracinus Anisodactylus (Gynandrotarsus) 70 anthrax Elaphropus 256 apache Trechus (Trechus) 502 apachensis Amara (Amara) 41 apicalis Trechus (Trechus) 502 appalachia Nebria (Reductonebria) 325 approximatum Bembidion (Notaphus) 95 approximatus Cyclotrachelus (Cyclotrachelus) 215 apricaria Amara (Bradytus) 41 aquaticus Notiophilus 343 aratum Bembidion (Eupetedromus) 95 aratus Dyschiriodes (Dyschiriodes) 244 arcanus Pterostichus (Hypherpes) 408 arctica Miscodera 322 arcticola Pterostichus (Cryobius) 408 arcticum Bembidion (Plataphodes) 95 ardelio Bradycellus (Stenocellus) 159 arenaria Halocoryza 273 arenaria Pseudamara 382 aridus Bradycellus (Stenocellus) 159 aridus Morion 323 arizonae Bembidion (Odontium) 95 arizonae Trechus (Trechus) 502 arizonensis Cymindis (Cymindis) 223 arizonica Lebia (Lebia) 297 arkansana arkansana Nebria (Reductonebria) 325 arkansana edwardsi Nebria (Reductonebria) 326 arkansana fragilis Nebria (Reductonebria) 326 arkansana oowah Nebria (Reductonebria) 326 assimilis Diplocheila (Isorembus) 236 assimilis Pseudanophthalmus 382 aterrimus Diplous 239 atratus Pterostichus (Abacidus) 408 atriceps Lebia (Loxopeza) 297 atriceps Philorhizus 368 atrichatus Harpalus 275 atrimedeus Bradycellus (Triliarthrus) 159 atripennis Cymindis (Pinacodera) 223 atripes Galerita (Progaleritina) 269 atriventris Lebia (Loxopeza) 297 audax Pseudanophthalmus 382 augustus Chlaenius (Chlaenius) 192

aulica Amara (Curtonotus) 42 aurata Amara (Amara) 42 aurata Blethisa multipunctata 148 auratus Carabus (Autocarabus) 185 auriga Pterostichus (Cryobius) 408 auripennis Schizogenius (Schizogenius) 465 aurocinctum Calosoma (Calodrepa) 176 austinica Rhadine 443 austinicus Paratachys 357 autumnalis Trichotichnus (Iridessus) 512 auxiliator Bembidion (Notaphus) 95 avernus Pseudanophthalmus 382 avida Amara (Bradytus) 43 avidum Bembidion (Hirmoplataphus) 95 azureipennis Brachinus (Neobrachinus) 150 azurescens Chlaenius (Chlaenius) 192 babcocki Rhadine araizai 443 badipennis Bradycellus (Triliarthrus) 159 baldwini Pterostichus (Hypherpes) 409 balesi Rhadine 443 balli Bembidion (Bracteon) 95 balli Euproctinus (Neoeuproctus) 268 balli Harpalus 275 balsamensis Trechus (Microtrechus) 502 barbarae Thalassotrechus 500 barbarinus Pterostichus (Hypherpes) 409 barberi Anillinus 67 barberi Pseudanophthalmus 383 barberi Trechus (Microtrechus) 503 barri Pseudanophthalmus 383 barryorum Pterostichus (Cryobius) 409 basale Agonum (Circinalidia) 19 basicorne Bembidion (Plataphus) 96 basillaris Amara (Amara) 43 beaklei Pseudanophthalmus 383 behrensi Pseudomorpha 403 behrensi Scaphinotus (Neocychrus) 451 belfragei Amara (Zezea) 43 belleri Agonum (Agonum) 19 bembidioides Sericoda 476 bendermani Pseudanophthalmus 383 beyeri Pterostichus (Hypherpes) 409 bicarinatus Sphaeroderus 478 bicolor Agonum (Platynomicrus) 19 bicolor Galerita (Progaleritina) 270

557

558

André Larochelle & Marie-Claude Larivière

bicolor Helluomorphoides praeustus 294 bicolor Pentagonica 366 bicolor Plochionus (Menidius) 375 bifossulatum Bembidion (Ochthedromus) 96 bifrons Amara (Amara) 43 biguttatus Notiophilus 343 bilineata Lebia (Lebia) 298 bilobus Scaphinotus (Nomaretus) 451 bimaculatum Bembidion 96 binotatus Anisodactylus (Anisodactylus) 70 binotatus Stenolophus (Agonoderus) 483 biocryus Pterostichus (Cryobius) 409 biplagiatus Axinopalpus 86 bipustulata Clivina (Paraclivina) 205 bisulcatus Porotachys 381 bitaeniata Lebia (Lebia) 298 bivittata Lebia (Lebia) 298 blanchardi Amara (Curtonotus) 44 blanchardi Gastrellarius 271 blanchardi Selenophorus (Selenophorus) 470 blandus Cymindis (Pinacodera) 224 blatchleyi Cyclotrachelus (Evarthrus) 215 bogemannii Sericoda 477 bokori Amara (Curtonotus) 44 borealis Cymindis (Cymindis) 224 borealis Notiophilus 344 borealis Pelophila 366 bowditchii Bembidion (Odontium) 96 bowlingi Trechus (Microtrechus) 503 boyeri Eucheila (Inna) 268 boyeri Inna 295 boyeri see Eucheila (Inna) 295 brachythorax Bembidion (Plataphodes) 97 bradycellinus Tachys 494 breve Bembidion (Plataphodes) (Bembidion incertum) 97 brevicolle Agonum (Agonum) 19 brevicollis Loxandrus 312 brevicollis Nebria (Nebria) 326 brevicollis Notiobia (Anisotarsus) 340 brevicornis Olisthopus 347 brevicornis Pterostichus (Cryobius) 409 brevilabris brevilabris Chlaenius (Chlaeniellus) 192 brevilabris texanellus Chlaenius (Chlaeniellus) 192 brevilobus Microlestes brevilobus 321

brevis Elaphropus 256 brevis Oodes 352 brevis Pseudanophthalmus 383 brevisetosus Schizogenius (Schizogenius) 466 brevispinus Dyschiriodes (Eudyschirius) 245 brevistriatum Bembidion (Hydriomicrus) 97 breviusculus breviusculus Pterostichus (Hypherpes) 410 breviusculus Loxandrus 312 breviusculus mimus Pterostichus (Hypherpes) 410 breviusculus Selenophorus 470 brevoorti Cyclotrachelus (Cyclotrachelus) 215 brevoorti Sphaeroderus nitidicollis 479 browni Amara (Bradytus) 44 brunnea Amara (Amara) 44 brunnea Coptodera 212 brunneomarginatus Platynus (Platynus) 370 brunneus Dicheirus 235 brunnicollis Elaphropus 256 brunnipes Amara (Amara) 44 bruxellense Bembidion (Peryphus) 97 bryanti biocryus Pterostichus (Cryobius) 410 bryanti bryanti Pterostichus (Cryobius) 410 bryanti bryantoides Pterostichus (Cryobius) 410 bryanti cacumenis Pterostichus (Cryobius) 410 bryanti cacumenis see Pterostichus (Cryobius) 411 bryanti stantonensis Pterostichus (Cryobius) 410 bryanti Tachys 494 bryanti tiliaceoradix Pterostichus (Cryobius) 410 buchannani Colpodes 212 buchannani Metacolpodes 319 bucolicus Pterostichus (Hypherpes) 411 bullatus Scaphinotus (Brennus) 452 bumeliae Lebia (Lebia) 298 cacumenis Pterostichus (Cryobius) 411 caenus Anisodactylus (Aplocentroides) 71 caerulea Calleida (Philophuga) 172 caeruleicollis Chlaenius (Agostenus) 193 calathinus Loxandrus 312 calcareus Pseudanophthalmus 383 calceus Calathus (Neocalathus) 169 calidum Calosoma (Chrysostigma) 176 californica californica Amara (Amara) 44 californica Clivina 206 californicum Bembidion (Hydriomicrus) 97

Index californicus Anisodactylus (Anisodactylus) 71 californicus Axinopalpus 86 californicus Bradycellus (Stenocellus) 160 californicus Cymindis (Cymindis) 224 californicus Diplous 239 californicus Elaphrus (Elaphrus) 262 californicus Pasimachus (Pasimachus) 360 californicus Pterostichus (Hypherpes) 411 californicus Stenomorphus californicus 491 caligans Pterostichus (Hypherpes) 411 caliginis Trechus (Trechus) 503 caliginosus Harpalus (Megapangus) 275 callens Bembidion 97 calliope Lebia (Lebia) 298 calva Nebria (Catonebria) 326 campbelli Nebria (Nipponebria) 326 campbelli Nipponebria (Vancouveria) 340 campbelli Pterostichus (Pseudoferonina) 411 campicola Dyschiriodes (Dyschiriodes) 245 canadense Agonum (Europhilus) 20 canadensis Acupalpus (Acupalpus) 13 canadensis Sphaeroderus canadensis 478 canadianum Bembidion (Trepanedoris) 98 canallatus Pterostichus (Hypherpes) 411 cancellatum Calosoma (Tapinosthenes) 177 cancellatus Carabus (Autocarabus) 186 capax Elaphropus 257 capitata Polpochila (Phymatocephalus) 380 capnicus Brachinus (Neobrachinus) 150 carbo Stenolophus (Stenolophus) (Stenolophus carbonarius) 483 carbonarius Anisodactylus (Anisodactylus) 71 caribou Pterostichus (Cryobius) 411 carinata Amara (Curtonotus) 45 carinatum Bembidion (Odontium) 98 carinula Bembidion (Bracteon) 98 carolinae Anillinus 67 carolinae Trechus (Trechus) 503 carolinense Bembidion (Plataphus) 98 carolinensis Bradycellus (Stenocellus) 160 carolinus carolinus Pterostichus (Monoferonia) 411 carolinus fumorum Pterostichus (Monoferonia) 412 carri Dyschiriodes 245 carri Nebria (Catonebria) 327 carus Acupalpus (Acupalpus) 14

559

caseyi Anillaspis 67 castanea Pseudomorpha 403 castaneus Infernophilus 295 castaneus Pterostichus (Hypherpes) 412 castanipes Pterostichus (Hypherpes) 412 castor Bembidion (Notaphus) 99 castor Pterostichus (Melanius) 412 castum Bembidion (Lionepha) 99 catenaria Blethisa 148 catenata Nebria (Catonebria) 327 catherinae Pseudanophthalmus 383 catoryctos Pseudanophthalmus 383 caudata Rhadine 443 caudicalis Pterostichus (Lamenius) 412 caurinus Chlaenius (Brachylobus) 193 cautus Harpalus 276 cavicollis Scaphinotus (Nomaretus) 452 cazieri Platynus 370 celeris Loxandrus 312 celiana Amara (Bradytus) 45 cephalotes cephalotes Broscus 168 cephalus Apristus 81 cephalus Notiobia (Anisotarsus) 341 cerberus cerberus Pseudanophthalmus 384 cerberus completus Pseudanophthalmus 384 cerberus Dyschiriodes (Dyschiriodes) 245 cervicalis Loxandrus 312 chalcea Amara (Amara) 45 chalceum Bembidion (Pseudoperyphus) 99 chalcites Poecilus (Poecilus) 376 chalybeus Trechus (Trechus) 503 chamissonis Carabus (Tomocarabus) 186 champlaini Pseudomorpha 403 chaparralus Selenophorus 470 charlottae Nebria (Reductonebria) 327 chaudoiri Chlaenius (Chlaenius) 193 chermocki Dicaelus (Liodicaelus) 230 chintimini Bembidion (Lionepha) 100 chipewyan Pterostichus (Cryobius) 413 chiricahuanus Schizogenius (Schizogenius) 466 chthonius Pseudanophthalmus 384 cibolensis Brachinus (Neobrachinus) 150 cicatricosus Elaphrus (Neoelaphrus) 263 ciliaris ciliaris Pseudanophthalmus 384 ciliaris colemanensis Pseudanophthalmus 384

560

André Larochelle & Marie-Claude Larivière

ciliaris orlindae Pseudanophthalmus 384 cincinnati Loxandrus 313 cincticollis Batenus (Batenus) 91 cincticollis Platynus 370 cincticollis Stenolophus (Stenolophus) 483 circulosus Pterostichus (Eosteropus) 413 circulus Loxandrus 313 circumcincta Calleida (Calleida) 172 circumcinctus Chlaenius (Chlaeniellus) (Chlaenius perplexus) 193 ciudadensis Amara (Amara) 45 clairvillei Elaphrus (Neoelaphrus) 263 clairvillei Helluomorphoides 293 clemens Bembidion (Trepanedoris) 100 cnephosus Pseudanophthalmus 384 cochlearis Pterostichus (Hypherpes) 413 cockerelli Elaphropus 257 coelebs Amara (Amara) 45 cognatus Dicheirotrichus (Trichocellus) 234 cognatus Trichocellus (Trichocellus) 511 cohni Harpalus (Glanodes) 276 cohni Platynus 371 collare Agonum 20 collaris Clivina (Clivina) 206 collaris Lebia (Lebia) 299 collucens Loxandrus 313 coloradense Bembidion (Notaphus) 100 coloradensis Axinopalpus 86 coloradensis Nebria (Catonebria) 327 coloradensis Poecilus (Poecilus) 376 coloradensis Trechus (Trechus) 504 columbiensis Paratachys 357 colvillense Bembidion (Bracteon) 100 colvillensis Amara (Bradytus) 46 comatus Dyschiriodes (Eudyschirius) 245 comma Stenolophus (Agonoderus) 484 commotum Bembidion 100 communis Amara (Amara) 46 commutabilis Pterostichus (Argutor) (Pterostichus leconteianus) 413 compactus Dyschiriodes (Dyschiriodes) 245 compar Harpalus (Pseudoophonus) (Harpalus bicolor) 276 complanatus Cymindis (Pinacodera) 224 complanatus Laemostenus (Laemostenus) 296

complanulum Bembidion (Plataphodes) 101 compressum Bembidion (Plataphodes) 101 comptus Loxandrus 313 concinnus Loxandrus 313 concinnus Selenophorus (Selenophorus) 470 concolor Bembidion (Hirmoplataphus) 101 concreta Amara (Curtonotus) 46 concretum Bembidion (Trepanedoris) 101 condei Xenotrechus 513 conditus Pseudanophthalmus 384 conflata Amara (Amara) 46 conformis Brachinus (Neobrachinus) 151 conformis Bradycellus (Triliarthrus) 160 conformis Trechus (Trechus) 504 confusa Amara (Amara) 46 confusum Bembidion (Odontium) 102 congener Bradycellus (Stenocellus) 160 congener Elaphropus 257 congesta Loricera pilicornis 311 congestus Pterostichus (Hypherpes) 414 congruens Discoderus (Discoderus) 240 conjugens Elaphropus 257 conjunctus Stenolophus (Agonoleptus) 484 connivens Bembidion (Trepanedoris) 102 consanguinea Pseudomorpha 404 consanguineum Bembidion 102 consimile Agonum (Europhilus) 20 consimile Bembidion 102 consobrinus Anisodactylus (Anisodactylus) 72 consobrinus Dyschiriodes (Dyschiriodes) 246 conspersum Bembidion (Notaphus) 103 constricta Rhadine 444 constricticolle Bembidion 103 constrictum Bembidion 103 constrictus Apristus 82 constrictus Cyclotrachelus (Evarthrus) 215 consuetum Bembidion (Notaphus) 103 contractum Bembidion 103 contractus Metrius contractus 319 contractus Selenophorus (Celiamorphus) 470 contumax Loxandrus 313 convexa Amara (Amara) 47 convexa Clivina (Paraclivina) 206 convexior Stenomorphus 491 convexulum Bembidion (Semicampa) 103

Index convivus Cyclotrachelus (Evarthrus) 216 coracinus Myas 323 coracinus Pterostichus (Euferonia) 414 corax Tachys 494 cordatum Bembidion (Notaphus) 104 cordatus Scaphinotus (Brennus) 452 cordicollis Brachinus (Neobrachinus) 151 cordicollis Chlaenius (Chlaenius) 193 cordicollis Discoderus (Selenalius) 241 cordicollis Pseudanophthalmus 385 cordifer Harpalus (Opadius) 277 corpulentus Harpalus (Glanodes) 277 corrusculus Pterostichus (Phonias) 414 corvinus Pterostichus (Melanius) 415 corvus Agonum (Agonum) 20 corvus Poecilus (Poecilus) 377 costatus Dicaelus (Dicaelus) 230 costatus Pterostichus (Lenapterus) 415 costipennis Brachinus (Neobrachinus) 151 coxendix Bembidion (Odontium) 104 crassicollis Discoderus (Discoderus) 241 crassicornis crassicornis Nebria (Boreonebria) 327 crassicornis intermedia Nebria (Boreonebria) 327 crassiscapus Trechus (Trechus) 504 crassispina Amara (Amara) 47 crassus Promecognathus 381 craterensis Pterostichus (Hypherpes) 415 crenatus Dicaelus (Dicaelus) 230 crenatus Loxandrus 313 crenatus Scaphinotus (Brennus) 452 crenicollis Pterostichus (Hypherpes) 415 crenistriatum Agonum (Olisares) 20 crenulatum Agonum (Circinalidia) 21 crenulatus crenulatus Schizogenius (Genioschizus) 466 crenulatus Loxandrus 313 cribricollis Cymindis (Cymindis) 224 criddlei Dyschiriodes (Eudyschirius) 246 cristatus Scaphinotus (Brennus) 452 croceicollis Technophilus croceicollis 498 cronkhitei Pseudomorpha 404 crossi Diplocheila (Isorembus) 237 crucigerus Panagaeus 356 cubanus Thalpius 500 cumatilis Chlaenius (Chlaenius) 193

561

cumberlandus Pseudanophthalmus 385 cumberlandus Trechus (Trechus) 504 cunctipeps Harpalus (Glanodes) 277 cuneatulus Pterostichus (Hypherpes) 416 cuprea Stenocrepis (Stenous) 481 cupreolata Amara (Amara) 47 cupreolus Selenophorus (Selenophorus) 470 cupreum Agonum (Agonum) 21 cupripenne Agonum (Agonum) 21 cursitans Loxandrus 314 cursitor Poecilus (Poecilus) 377 cursor Chlaenius (Chlaenius) 194 curticollis Bradycellus (Liocellus) 161 curtipennis Microlestes 321 curtulatum Bembidion (Plataphus) (Bembidion flebile) 104 curvispinus Dyschiriodes (Dyschiriodes) 246 cuyama Tanystoma 497 cyanescens Myas 324 cyanicolor Poecilus (Poecilus) 377 cyanipennis Brachinus (Neobrachinus) 151 cyanipennis Lebia (Lebia) 299 cyanochroaticus Brachinus (Neobrachinus) 152 cyanope Agonum (Stictanchus) 22 cyclifer Agonum (Agonum) 22 cylindrica Pseudomorpha 404 cyrtonotoides Harpalus (Pharalus) 277 dallasensis Discoderus (Discoderus) 241 danmanni Nebria (Reductonebria) 328 darlingtoni Agonum (Europhilus) 22 darlingtoni darlingtoni Pseudanophthalmus 385 darlingtoni Nebria (Reductonebria) 328 darlingtoni persimilis Pseudanophthalmus 385 daurica Amara (Curtonotus) 48 dauricum Bembidion (Peryphanes) 104 debiliceps Bembidion (Notaphus) 105 debilis alpinus Scaphinotus (Maronetus) 452 debilis Anillodes 69 debilis debilis Scaphinotus (Maronetus) 452 decempunctata Loricera 310 decentis Platynus (Platynus) 371 deceptivum Agonum (Agonum) 22 deceptivus Pseudanophthalmus 385 deceptor Thalpius 500 deceptrix Lebia (Loxopeza) 299

562

André Larochelle & Marie-Claude Larivière

decora Calleida (Calleida) 172 decorum Agonum (Stictanchus) 23 decorus Bradycellus (Stenocellus) 161 dejeanellus Cyclotrachelus (Cyclotrachelus) (Cyclotrachelus morio) 216 dejeanii Dyschiriodes 246 dejectum Bembidion 105 delectum Zuphium 514 delicatus Pseudanophthalmus 385 demissus Axinopalpus 86 dentatum Omophron 348 denticollis Xenotrechus 513 denticulatus Axinopalpus 86 dentipes Clivina (Semiclivina) 206 deparca Amara (Curtonotus) 48 depressicollis Anisodactylus (Anadaptus) 72 depressulus Selenophorus (Selenophorus) 471 depressus Leptotrachelus 308 depressus Micranillodes 320 depressus Pasimachus (Pasimachus) 360 depressus Schizogenius (Schizogenius) 466 desertus desertus Pseudanophthalmus 385 desertus Diplochaetus 235 desertus Harpalus (Pharalus) 277 desertus major Pseudanophthalmus 385 desolata Nebria (Reductonebria) 328 diabolus Pterostichus (Hypherpes) 416 diaphana Polyderis 380 dichrous Trichotichnus (Trichochichnus) 512 dietriechi Trechus (Trechus) 504 dietzii Calosoma (Microcallisthenes) 177 digitus Pseudanophthalmus 386 dilatatus Dicheirus dilatatus 235 dilatatus dilatatus Dicaelus (Paradicaelus) 230 dilatatus sinuatus Dicaelus (Paradicaelus) 230 diligendus Pterostichus (Monoferonia) 416 diligens Bembidion (Emphanes) 105 dilutior Hyboptera 295 diplophryus Poecilus (Poecilus) 377 directus Notiophilus 344 discipulus Bradycellus (Stenocellus) 161 discoderoides Selenophorus (Selenophorus) 471 discoideus Anisodactylus (Anadaptus) 72 discopunctatus Selenophorus (Celiamorphus) 471 discors Amara (Amara) 48

discors Calosoma (Microcallisthenes) 177 discus Blemus 147 disjunctum Bembidion (Lionepha) 105 dissecta Rhadine 444 dissimilis Stenolophus (Stenolophus) 485 distincta Micrixys 320 diversa Nebria (Reductonebria) 328 divisa Lebia (Lamprias) 299 dohrni Anillinus 67 dolosus Elaphropus 257 dorsale Bembidion (Notaphus) 105 dorsalis Leptotrachelus 308 dorsalis Thalpius 500 dubius Cratacanthus 213 dubius Synuchus (Pristodactyla) 493 dulcicollis Anisodactylus (Gynandrotarsus) 72 dunavani Anillinus 68 duodecimstriata Stenocrepis (Stenous) 481 duplicatus Pasimachus (Pasimachus) 361 durangoense Bembidion (Odontium) 106 duryi Loxandrus 314 dyschirinum Bembidion (Metallina) 106 Dyschirius 256 ebeninus Pterostichus (Melanius) 416 ecarinatus Pterostichus (Hypherpes) 417 edax Paratachys 357 edentulus Dyschiriodes (Eudyschirius) 246 egberti Pseudanophthalmus 386 egens Bembidion 106 elegans Badister (Badister) 87 elegans elegans Cymindis (Cymindis) 225 elegans mobilensis Cymindis (Cymindis) 225 elegans Nemotarsus 339 elegans Stenocrepis (Stenous) 482 elegantulus Lachnophorus 296 elevatus coloradensis Scaphinotus (Scaphinotus) 453 elevatus elevatus Scaphinotus (Scaphinotus) 453 elevatus flammeus Scaphinotus (Scaphinotus) 453 elevatus lengi Scaphinotus (Scaphinotus) 453 elevatus neomexicanus Scaphinotus (Scaphinotus) 453 elevatus Pseudanophthalmus 386 elevatus tenebricosus Scaphinotus (Scaphinotus) 454

Index ellipsis Amara (Amara) 48 ellipsis Harpalus 278 ellipticus Selenophorus (Celiamorphus) 471 elongatulum Agonum (Stictanchus) 23 elongatulus Brachinus (Neobrachinus) 152 elongatus Anillinus 68 elongatus Dicaelus (Paradicaelus) 231 elongatus Pasimachus (Pasimachus) 361 elongatus Pseudanophthalmus 386 elongatus Selenophorus (Selenophorus) 471 emaciatus Diplochaetus (Diplochaetus desertus) 236 emancipata Amara (Amara) 49 emarginatus Chlaenius (Anomoglossus) 194 emersoni Pseudanophthalmus 386 empetricola Pterostichus (Cryobius) 417 engelhardti Pseudanophthalmus 386 engelmani Cyclotrachelus (Evarthrus) 216 enodis Pterostichus (Gastrosticta) 417 ephippiatus Pericompsus (Pericompsus) 367 ephippigerum Bembidion (Peryphodes) 106 erasum Bembidion (Lionepha) 106 eraticus Harpalus (Plectralidus) 278 eremicola Calosoma (Carabosoma) 177 eremita Pseudanophthalmus 386 errans Agonum (Stereagonum) 23 erratica Amara (Amara) 49 erraticus Loxandrus 314 erro Polpochila (Phymatocephalus) 380 erythrocerus Dyschiriodes (Dyschiriodes) 246 erythropus Chlaenius (Chlaenius) 194 erythropus Harpalus (Pseudoophonus) 279 eschscholtzii Nebria (Reductonebria) 328 esurialis Lebia (Lebia) 299 esuriens Pterostichus (Hypherpes) 417 evanescens Cymindis (Cymindis) 225 evansi Amblygnathus 63 evidens Bembidion (Notaphus) 106 exarata Amara (Bradytus) 49 exarata Evolenes 269 exaratum Agonum (Europhilus) 23 excavatum Agonum (Agonum) 24 excrucians Pseudomorpha 404 exiguus exiguus Pseudanophthalmus 386 exiguus furtivus Pseudanophthalmus 387

563

exilis Rhadine 444 exlineae Amara (Amara) 49 exochus Dyschiriodes (Dyschiriodes) 247 exoticus Pseudanophthalmus 387 explanatus Anillaspis 67 explodens Metrius 320 exstans Bradycellus (Stenocellus) 161 extendus Loxandrus 314 extensicolle Agonum (Stictanchus) 24 extensicollis Stenolophus (Stenolophus) 485 externum Calosoma (Callitropa) 177 extimum Agonum (Stictanchus) 24 eyrinota Amara (Amara) 49 faber Cyclotrachelus (Cyclotrachelus) 216 fallax Pterostichus (Abacidus) 417 falli Platynus 371 falli Pseudomorpha 404 falli Pterostichus (Hypherpes) 417 falli Schizogenius (Schizogenius) 466 falli Tachyta 496 fallianum Agonum (Agonum) 25 falsum Bembidion (Plataphus) 106 familiaris Amara (Amara) 50 famulus Selenophorus (Selenophorus) 471 farcta Amara (Amara) 50 farrarae Bembidion (Plataphodes) 107 farrelli Pseudanophthalmus 387 fasciata Clivina (Paraclivina) 206 fasciatus Panagaeus 356 fasciatus Tetragonoderus (Tetragonoderus) 498 fastigatus Pseudanophthalmus 387 fatuus Elaphropus 257 fatuus Selenophorus (Selenophorus) 471 faunus Harpalus (Pseudoophonus) 279 favicollis Brachinus (Neobrachinus) 152 felix Pentagonica 366 femoralis Pterostichus (Phonias) 418 femoratum Bembidion (Peryphus) 107 fenderi Bradycellus (Bradycellus) 161 fenestratus Dromius (Dromius) 243 fenyesi fenderi Pterostichus (Hypherpes) 418 fenyesi fenyesi Pterostichus (Hypherpes) 418 ferrea Clivina (Paraclivina) 207 ferreum Agonum (Stereagonum) 25 ferrugineus Badister (Badister) 87

564

André Larochelle & Marie-Claude Larivière

ferrugineus Dyschiriodes (Eudyschirius) 247 ferrugineus Elaphropus 257 ferrugineus Helluomorphoides 293 ferrugineus Leistus (Leistus) 307 ferrugineus Schizogenius (Schizogenius) 467 ferruginosum Agonum (Platynomicrus) 25 ferruginosus Leistus (Neoleistus) 307 festinans Bradycellus (Stenocellus) 161 festiva Coptodera 213 festivum Bembidion (Liocosmius) 107 fidele Agonum (Agonum) 25 filicornis Diplous 240 filicornis Olisthopus 347 filiformis Dyschiriodes (Dyschiriodes) 247 fimbriata Calleida (Calleida) 173 finitimus Carabus (Tanaocarabus) 186 finitimus Elaphrus (Elaphrus) 264 fissicollis Scaphinotus (Nomaretus) 454 flaccidus Chlaenius (Chlaeniellus) 195 flavicauda Mioptachys 322 flavilimbus Acupalpus (Philodes) 14 flavipes Asaphidion 84 flavipes flavipes Badister (Badister) 87 flavipes laticeps Badister (Badister) 88 flavipes Pentagonica flavipes 366 flavipes Stenolophus (Stenolophus) 485 flavopictum Bembidion 107 flebilis Amara (Zezea) 51 flohri Bembidion (Bembidion henshawi) 107 flohri Ophryogaster 354 floridae Clivina (Paraclivina) 207 floridana Onota 351 floridanus Chlaenius (Chlaeniellus) 195 floridanus Helluomorphoides praeustus 294 floridanus Loxandrus 314 floridensis Cyclotrachelus (Evarthrus) 216 fluvialis Oodes 352 fluvialis Oodes americanus 351 forreri Carabus (Tanaocarabus) 186 forreri Galerita (Progaleritina) 270 fortestriatum Bembidion (Trepanedoris) 107 fortis Amara (Neopercosia) 51 fortis Anillinus 68 fossifrons Patrobus (Patrobus) 363 fossigerum Agonum (Agonum) 26

fossor Clivina (Clivina) 207 fossulatus Selenophorus (Celiamorphus) 472 foveata Loricera 310 foveocollis Patrobus (Patrobus) 363 foveum Bembidion (Bracteon) 108 fowlerae Pseudanophthalmus 387 fraternus Harpalus (Euharpalops) 279 freitagi Cyclotrachelus (Cyclotrachelus) 217 frigida Nebria (Boreonebria) 328 frigidum Calosoma (Calosoma) 178 frigidus Pseudanophthalmus 387 frontale Bembidion (Trepanedoris) 108 frostii Chlaenius purpuricollis 201 fucatus Cyclotrachelus (Cyclotrachelus) 217 fuchsi Pterostichus (Hypherpes) 418 fuchsii Bembidion 108 fugax Bembidion (Trichoplataphus) 109 fulgens Harpalus 280 fulgens Loxandrus 314 fulgens Trichotichnus (Iridessus) 512 fulgida Calleida (Calleida) 173 fuliginosus Elaphrus (Neoelaphrus) 264 fuliginosus Stenolophus (Stenolophus) 485 fulleri Pseudanophthalmus 387 fulminatus Brachinus (Neobrachinus) 152 fulva Amara (Bradytus) 51 fulvilabris Harpalus (Euharpalops) 280 fumans Brachinus (Neobrachinus) 152 funebris Anchomenus 66 furtivus Cyclotrachelus (Evarthrus) 217 furvus Anisodactylus (Anisodactylus) 73 furvus carinatus Dicaelus (Paradicaelus) 231 furvus furvus Dicaelus (Paradicaelus) 231 fuscata Lebia (Lebia) 299 fuscatus Stenolophus (Stenolophus) 486 fusciceps Axinopalpus 86 fuscicornis Chlaenius (Chlaenius) 195 fuscicornis Elaphropus 258 fuscipalpis Harpalus (Harpalobius) (Harpalellus basilaris) 280 fuscipes Calathus (Calathus) 169 fuscus constrictus Pseudanophthalmus 387 fuscus fuscus Pseudanophthalmus 388 gagatinus Selenophorus (Selenophorus) 472 galvestonicum Agonum (Europhilus) 26

Index gebhardis Brachinus (Neobrachinus) 153 gebleri albimontis Nebria (Catonebria) 329 gebleri cascadensis Nebria (Catonebria) 329 gebleri fragariae Nebria (Catonebria) 329 gebleri gebleri Nebria (Catonebria) 329 gebleri rathvoni Nebria (Catonebria) 329 gebleri siskiyouensis Nebria (Catonebria) 329 gebleri turbatum Bembidion (Plataphus) 109 geniculatus Brachinus (Neobrachinus) 153 georgei Bradycellus (Triliarthrus) 161 georgiae Pseudanophthalmus 388 gerstlensis Pterostichus (Cryobius) 418 gibba Amara (Amara) 51 gibbipennis Dyschiriodes (Dyschiriodes) 247 gibbus Loxandrus 314 gigas Cyclotrachelus (Evarthrus) 217 gilae Omophron 348 glacialis Amara (Bradytus) 51 glaucus Chlaenius (Chlaeniellus) 195 gliscans Pterostichus (Hypherpes) 419 globiceps Pseudanophthalmus 388 globosus Dyschiriodes (Eudyschirius) 247 globulosus Dyschiriodes (Dyschiriodes) 248 glomerosus Dyschiriodes (Eudyschirius) 248 gordoni Bembidion (Plataphus) 109 goryi Carabus (Carabus) (Carabus limbatus) 186 gouleti Nebria (Boreonebria) 330 glacialis Amara (Bradytus) 51 gracilenta Rhadine 444 graciliforme Bembidion (Eupetedromus) 109 gracilior Pterostichus (Hypherpes) 419 gracilis Pseudanophthalmus 388 granarius Elaphropus 258 granarius Selenophorus (Celiamorphus) 472 grandiceps Badister (Baudia) 88 grandiceps Bembidion (Trichoplataphus) 109 grandis Lebia (Loxopeza) 300 grandis Pseudanophthalmus 388 granulatus granulatus Carabus (Carabus) 187 granulatus hibernicus Carabus (Carabus) 187 graphicum Bembidion (Notaphus) 110 grapii Bembidion (Peryphanes) 110 gratiosum Agonum (Europhilus) 26 gratiosum Bembidion (Plataphus) 110 gravesi Cyclotrachelus (Evarthrus) 217

565

gravidus Cyclotrachelus (Evarthrus) 217 gravis Harpalus (Pharalus) 281 gravis Lophoglossus 309 gregalis Pterostichus (Hypherpes) 419 gregaria Nebria (Reductonebria) 330 gregarius Calathus (Neocalathus) 170 grossum Omophron 349 grossus Euryderus 269 guttula Lebia (Lebia) 300 guyotii Scaphinotus (Steniridia) 454 gyllenhali castanipes Nebria (Boreonebria) 330 gyllenhali lassenensis Nebria (Boreonebria) 330 gyllenhali lindrothi Nebria (Boreonebria) 330 gyllenhalii Pachyteles (Pachyteles) 355 habilis Axinopalpus 86 hadenoecus Pseudanophthalmus 388 haematopus Stereocerus 492 haemorrhoidalis Dyschiriodes (Paradyschirius) 248 haftorni Pterostichus (Cryobius) 419 hageni Bembidion 110 haida Nebria (Reductonebria) 330 haldemanni Lophoglossus 309 halophilus Tachys 494 hamiltoni Pterostichus (Abacidus) 419 haoe Trechus (Microtrechus) 504 haplomus Anisodactylus (Gynandrotarsus) 73 harpalina Amara (Amara) 52 harpalinus Bradycellus (Bradycellus) 161 harpalinus Chlaenius (Agostenus) 195 harpaloides Anisodactylus (Gynandrotarsus) 73 harrisii Agonum (Agonum) 27 harrisii Anisodactylus (Anisodactylus) 73 haruspex Bembidion (Plataphodes) 110 hastii Bembidion (Blepharoplataphus) 111 hatchi Harpalus (Pseudoophonus) 281 hatchi Pterostichus (Hypherpes) 419 hatchii Scaphinotus (Stenocantharus) 454 haydeni haydeni Calosoma (Blaptosoma) 178 haydeni punctulicolle Calosoma (Blaptosoma) 178 hemphillii Cychrus hemphillii 214 henroti Neaphaenops tellkampfii 324 henroti Pseudanophthalmus 388 herbivagus Harpalus 281 herculaneus Pterostichus (Hypherpes) 419 hernandensis Cyclotrachelus (Cyclotrachelus) 217

566

André Larochelle & Marie-Claude Larivière

heros Cyclotrachelus (Evarthrus) 218 hesperium Bembidion (Bracteon) 111 hesperus Pseudanophthalmus 388 hicksi Amara (Amara) 52 hiemalis Dyschiriodes (Dyschiriodes) 248 higginbothami Pseudanophthalmus 389 hirsutus Brachinus (Neobrachinus) 153 hirsutus Pseudanophthalmus 389 hirta Physea 369 histrionica Lebia (Lebia) 300 hoegei Thalpius 500 hoffmani Pseudanophthalmus 389 holsingeri Pseudanophthalmus 389 honestum Bembidion (Pseudoperyphus) 111 honestus Gastrellarius 272 hoodi Hartonymus 293 horni abditus Pseudanophthalmus 389 horni Bembidion (Liocosmius) 111 horni caecus Pseudanophthalmus 389 horni horni Pseudanophthalmus 389 horni Thalpius 500 hornii Pterostichus (Hypherpes) 420 hortulanus Pseudanophthalmus 389 houstoni Selenophorus (Selenophorus) 473 howdeni Rhadine 444 huachuca Harpalus (Glanodes) 282 hubbardi Pseudanophthalmus 390 hubbardi Pseudomorpha 404 hubbardi Scaphinotus (Maronetus) 454 hubrichti Pseudanophthalmus 390 hudsonica Nebria (Boreonebria) 331 hudsonicus Pterostichus (Cryobius) 420 humboldtense Bembidion (Hirmoplataphus) 111 humboldti Pterostichus (Hypherpes) 420 humboldti Trechus (Trechus) 505 humboldtianus Bradycellus (Stenocellus) 162 humeralis Pseudanophthalmus 390 humidulus Pterostichus (Pseudoferonina) 420 humidus Stenolophus (Stenolophus) 486 humilis Pterostichus (Hypherpes) 420 hyalinus Paratachys 357 hydropicus Acupalpus (Acupalpus) 14 hydropicus avus Trechus (Trechus) 505 hydropicus beutenmuelleri Trechus (Trechus) 505 hydropicus canus Trechus (Trechus) 505

hydropicus hydropicus Trechus (Trechus) 505 hylacis Selenophorus (Selenophorus) 473 hyperboraeorum Bembidion (Plataphus) 111 hyperborea Amara (Curtonotus) 52 hypertrichosis Pseudanophthalmus 390 hypherpiformis Cyclotrachelus (Evarthrus) 218 hypogeus Pterostichus (Cylindrocharis) 420 hypolithos Platynus 371 hypolithos Pseudanophthalmus 390 icarus Loxandrus 314 ichabodopsis Brachinus (Neobrachinus) 153 idahoae Pterostichus (Hypherpes) 420 idahoana Amara (Amara) 52 idoneum Bembidion (Notaphus) 112 illectus Axinopalpus 87 illinoiensis Pseudanophthalmus 390 illustris Pterostichus (Hypherpes) 421 imitans Agonum (Agonum) 27 immaturum Bembidion (Eupetedromus) 112 immunda Amara (Triaena) 52 imperfectus Scaphinotus (Maronetus) 455 imperialensis Brachinus (Neobrachinus) 153 implicans Selenophorus (Selenophorus) 473 imporcitis Brachinus (Neobrachinus) 154 impotens Bembidion 112 impotens Discoderus (Discoderus) 241 impressefrons Clivina (Clivina) 208 impressicollis Diplocheila (Isorembus) 237 impressus Dercylinus 229 improvidens Bembidion (Plataphodes) 112 impunctatus Synuchus (Pristodactyla) 493 impuncticollis Amara (Amara) 52 impunctifrons Chlaenius (Chlaeniellus) 195 inaequale Bembidion (Bracteon) 113 inanis Pterostichus (Hypherpes) 421 incisus Cyclotrachelus (Evarthrus) 218 incompletus Scaphinotus (Maronetus) 455 incrassatus Geopinus 273 incrematum Bembidion (Eupetedromus) 113 incultus Stenolophus (Stenolophus) 487 incurvus Elaphropus 258 indecentis Platynus 372 indianae Anillinus 68 indianae Sphaeroderus 479 indianus Harpalus (Pharalus) 282

Index indigens Harpalus (Euharpalops) 282 indistinctum Bembidion (Notaphus) 113 indistinctus Acupalpus (Tachistodes) 14 inermis Pterostichus (Hypherpes) 421 inexpectatus inexpectatus Pseudanophthalmus 390 inexpectatus orientalis Pseudanophthalmus 390 inexpectatus Trechus (Microtrechus) 505 inexspectata Amara (Zezea) 53 inexspectata Blethisa 148 infernalis ewersi Rhadine 444 infernalis infernalis Rhadine 444 infernalis Pterostichus (Hypherpes) 421 inferus Loxandrus 315 infimus Loxandrus 315 infletus Scaphinotus (Nomaretus) 455 infuscatus Stenolophus (Agonoderus) 487 ingens ingens Nebria (Catonebria) 331 ingens riversi Nebria (Catonebria) 331 ingratus Calathus (Neocalathus) 170 innocuus Harpalus 282 innuens Olisthopus 347 inopinus Pterostichus (Hypherpes) 421 inquisitor Pseudanophthalmus 391 insignis Amara (Bradytus) 53 insignis Broscodera 168 insolita Rhadine 444 insulana see Stenocrepis (Stenocrepis) 482 insulana Stenocrepis (Stenocrepis) (Oodes texanus) 482 insularis Amara (Bradytus) 53 insularis Pseudanophthalmus 391 insulata Lebia (Lebia) 301 insulatum Bembidion (Notaphus) 114 insulsus Bradycellus (Stenocellus) 162 intectus Pterostichus (Hypherpes) 421 integer Dyschiriodes (Dyschiriodes) 249 integrum Bembidion (Pseudoperyphus) 114 intercepta Oxycrepis (Stolonis) 354 interior Cymindis (Cymindis) 225 interior Dyschiriodes (Eudyschirius) 249 intermedium Bembidion (Notaphus) 114 intermedius Bradycellus (Liocellus) 162 intermedius Notiophilus 344 intermedius Pseudanophthalmus 391 interruptus Chlaenius (Agostenus) 196

567

interruptus Scaphinotus (Brennus) 455 intersectus Pseudanophthalmus 391 intersectus Tetragonoderus (Tetragonoderus) 499 interstitialis Amara (Amara) 53 interstitialis Amphasia (Amphasia) 65 interventor Bembidion (Eurytrachelus) 114 iowensis Cyclotrachelus (Evarthrus) 218 iridescens Bembidion (Trechonepha) 115 iripennis Amblygnathus 64 iripennis Pterostichus (Euferonia) 421 isabellae Pterostichus (Hypherpes) 421 iterans Olisthopus 347 iuvenis Cyclotrachelus (Cyclotrachelus) 218 jacobianum Bembidion (Notaphus) 115 jacobina Amara (Curtonotus) 54 jacobinus Pterostichus (Hypherpes) 422 janthinipennis Brachinus (Neobrachinus) 154 janus Galerita (Progaleritina) 270 javalinopsis Brachinus (Neobrachinus) 154 jeanneli jeanneli Ameroduvalius 64 jeanneli rockcastlei Ameroduvalius 65 jeannelli Serranillus 478 jeffreyi Nebria (Reductonebria) 331 jejuna Rhadine 444 johnsoni Pterostichus (Orsonjohnsonus) 422 johnsoni Scaphinotus (Brennus) 455 jonesi Nelsonites 339 jonesi Pseudanophthalmus 391 jucunda Amara (Amara) 54 jucundum Bembidion (Bembidion) 115 julii Blethisa 148 kalumae Bembidion (Plataphus) 115 kansanus Brachinus (Neobrachinus) 154 kansanus Pterostichus (Hypherpes) 422 katiae Harpalus (Megapangus) 282 kavanaughi Amara (Zezea) 54 kavanaughi Brachinus (Neobrachinus) 154 kelloggi Scaphinotus (Scaphinotus) 456 kentuckensis Darlingtonea kentuckensis 228 kincaidi balli Nebria (Catonebria) 331 kincaidi kincaidi Nebria (Catonebria) 331 kirbyi Anisodactylus (Anisodactylus) 74 kirbyi Bradycellus (Triliarthrus) 162 koepkei koepkei Rhadine 445 koepkei privata Rhadine 445

568

André Larochelle & Marie-Claude Larivière

kotzebuei Pterostichus (Cryobius) 422 krameri Pseudanophthalmus 391 krekeleri Pseudanophthalmus 391 kuntzeni Pachyteles (Goniotropis) 355 kuprianovi Bembidion 115 kurnakowi Amara (Curtonotus) 54 labiatum Omophron 349 labontei Nebria (Catonebria) 332 laborans Pterostichus (Hypherpes) 422 lacertus Pterostichus (Hypherpes) 422 lachnophoroides Bembidion (Actedium) 115 lachrymosus Pterostichus (Euferonia) 422 lacunarium Bembidion (Peryphanes) 116 lacustris Amara (Curtonotus) 54 lacustris bellorum Nebria (Boreonebria) 332 lacustris lacustris Nebria (Boreonebria) 332 lacustris Platypatrobus 374 laesus Selenophorus (Selenophorus) 473 laetula Calybe (Ega) 185 laetulus Pericompsus (Pericompsus) 367 laetulus Poecilus (Poecilus) 377 laetus Anisodactylus (Pseudaplocentrus) 74 laevifasciatus Dyschiriodes (Antidyschiriodes) 249 laevigatus Elaphrus (Neoelaphrus) 264 laevipennis Amara (Amara) 54 laevipennis Cyclotrachelus (Cyclotrachelus) 218 laevipennis laevipennis Dicaelus (Liodicaelus) 231 laevipes Harpalus (Euharpalops) (Harpalus quadripunctatus; Harpalus egregius) 283 laevis Polyderis 380 laevissimus Promecognathus 381 lallemanti Pseudanophthalmus 391 lama Pterostichus (Hypherpes) 423 lampros Bembidion (Metallina) 116 lanei Pterostichus (Hypherpes) 423 lanei Pterostichus (Pseudoferonina) 423 lanei Rhadine 445 lapponicum Bembidion (Bracteon) 116 lapponicus lapponicus Elaphrus (Arctelaphrus) 264 lapponicus obliteratus Elaphrus (Arctelaphrus) 265 lariversi Calosoma (Microcallisthenes) 179 larochellei Dyschiriodes (Dyschiriodes) 249 larvalis Rhadine 445 larvatus Bradycellus (Stenocellus) 162 lassulus Pterostichus (Hypherpes) 423

latebricola Bembidion (Notaphus) 117 latens Apristus 82 lateralis Brachinus (Neobrachinus) 155 lateralis Loxandrus 315 laticeps Bembidion (Lymneops) 117 laticeps Harpalus (Euharpalops) 283 laticollis Apristus 82 laticollis Bradycellus (Liocellus) 163 laticollis Chlaenius (Chlaenius) 196 laticollis Cymindis (Cymindis) 225 latior Amara (Bradytus) 55 latipenne Calosoma (Microcallisthenes) 179 latipennis Tetragonoderus (Peronoscelis) 499 latitarsis Helluomorphoides 293 laxatum Bembidion (Plataphodes) 117 lecontei Bembidion 117 lecontei Bradycellus (Catharellus) 163 lecontei Diplochaetus 236 lecontei Elaphrus (Elaphrus) 265 lecontei Lebia (Polycheloma) 301 lecontei lecontei Galerita (Progaleritina) 271 lecontei Patrobus (Patrobus) 364 lecontei Pseudaptinus 402 lecontei Sphaeroderus stenostomus 480 lecontei Stenolophus (Agonoderus) 487 lecta Lebia (Lebia) 301 leiroides Harpalus (Harpalobrachys) 283 lemoulti Ozaena 355 lenae Bembidion 117 lengi Colliuris (Cosnania) 211 lengi Sphaeroderus canadensis 479 leonae Pseudanophthalmus 392 lepidum Calosoma (Chrysostigma) 179 leucoscelis monachus Chlaenius (Chlaenius) 196 leucoscelis sanantonialis Chlaenius (Chlaenius) 196 leucoscelis sonomae Chlaenius (Chlaenius) 197 levettei Bembidion (Bracteon) 117 levifaber Cyclotrachelus (Cyclotrachelus) 219 levigatum Bembidion (Hydrium) 118 levipes Elaphropus 259 lewisii Harpalus (Euharpalops) 284 lexingtoni Darlingtonea kentuckensis 229 liebecki Elaphropus 259 liebecki Scaphinotus (Nomaretus) 456 limatus Loxandrus 315

Index limbalis Stenolophus (Stenolophus) 488 limbatum Agonum (Circinalidia) 27 limbatus Cymindis (Pinacodera) 225 limicola Pseudanophthalmus 392 lindrothellus Bembidion (Lionepha) 118 lindrothi Amara (Bradytus) 55 lindrothi Elaphrus (Neoelaphrus) 265 lindrothi Microlestes 321 lindrothi Myas 324 lindrothi Schizogenius (Schizogenius) 467 lindrothi Rhadine 445 linearis Amerinus 64 linearis Microlestes 321 lineatus Bradycellus (Stenocellus) 163 lineola Stenolophus (Agonoderus) 488 lineolatus Schizogenius (Schizogenius) 467 liobasis Harpalus (Pseudoophonus) 284 lioptera Colliuris (Cosnania) 211 liratus Apristus 82 lissopterus Scarites (Scarites) 463 lithophilus indigaceus Chlaenius (Brachylobus) 197 lithophilus lithophilus Chlaenius (Brachylobus) 197 litigiosus Schizogenius (Schizogenius) 467 litoralis Tachys 495 littoralis Amara (Amara) 55 lituyae Nebria (Reductonebria) 332 lobatus Pterostichus (Hypherpes) 424 lobulata Lebia (Lebia) 301 lodingi Anisodactylus (Anisodactylus) 75 lodingi lodingi Scaphinotus (Steniridia) 456 lodingi obscurus Scaphinotus (Steniridia) 456 loedingi aladdini Pseudanophthalmus 392 loedingi aquaticus Pseudanophthalmus 392 loedingi distinguens Pseudanophthalmus 392 loedingi fluviatilis Pseudanophthalmus 392 loedingi loedingi Pseudanophthalmus 392 loedingi meridionalis Pseudanophthalmus 392 loedingi profundus Pseudanophthalmus 392 loganensis Pseudanophthalmus 393 longiceps Anillinus 68 longiceps Pseudanophthalmus 393 longiceps Rhadine 445 longiceps Scaphinotus (Neocychrus) 456 longicolle Zuphium 514 longicollis Discoderus (Discoderus) 241

569

longicollis Harpalus (Pseudoophonus) 284 longicollis Rhadine 446 longicornis Patrobus (Neopatrobus) 364 longipennis Leistus (Neoleistus) 308 longipennis Schizogenius (Schizogenius) 467 longipes Rhadine 446 longula Amara (Zezea) 56 longulus Acupalpus (Philodes) 15 longulus Dyschiriodes (Dyschiriodes) 250 lorquinii Bembidion (Bracteon) 118 louiseae Nebria (Reductonebria) 332 lubricus Pterostichus (Paraferonia) 424 lucens Acupalpus (Philodes) 15 lucens Loxandrus 315 lucidulus Apenes (Apenes) 80 lucidus Microlestes lucidus 321 luctuosus Pterostichus (Pseudomaseus) 424 lucublandus acomanus Poecilus (Poecilus) 377 lucublandus louisinus Poecilus (Poecilus) 377 lucublandus lucublandus Poecilus (Poecilus) 377 luculentum Bembidion 118 luculentus luculentus Trechus (Microtrechus) 505 luculentus unicoi Trechus (Microtrechus) 506 luculentus wayahensis Trechus (Microtrechus) 506 ludoviciana Colliuris (Calocolliuris) 211 ludovicianus Loxandrus 315 lugubris Bradycellus (Triliarthrus) 163 lugubris Xestonotus 513 lummi Bembidion (Lionepha) 118 lunicollis Amara (Amara) 56 lustrans Pterostichus (Bothriopterus) 424 lustrellus Bradycellus (Stenocellus) 164 lutulentum Agonum (Europhilus) 27 luxatum Calosoma (Microcallisthenes) 179 lyelli Nebria (Reductonebria) 333 lyratus Platynus (Platynus) 372 macradei Pseudanophthalmus 393 macrovulum Cyclotrachelus (Cyclotrachelus) 219 macrum Calosoma (Callitropa) 179 maculatus Badister (Badister) 88 maculatus Stenolophus (Agonoderus) 488 maculicolle Tanystoma 497 maculicornis Notiobia (Anisotarsus) 341 madmeridianus Leistus (Neoleistus) 308 maeander Carabus (Homoeocarabus) 188

570

André Larochelle & Marie-Claude Larivière

magnum Zuphium 514 major major Diplocheila (Isorembus) 237 major melissisa Diplocheila (Isorembus) 238 major Microlestes 322 malkini Pterostichus (Hypherpes) 425 mancus plethorus Pterostichus (Monoferonia) 425 mancus Pterostichus (Monoferonia) 425 mandibularoides Pterostichus (Cryobius) 425 mannerheimii Batenus (Batenus) 92 mannerheimii Dicheirotrichus (Oreoxenus) 234 mannerheimii Nebria (Reductonebria) 333 mannerheimii Platynus 372 mannerheimii Trichocellus (Oreoxenus) 512 mannii Scaphinotus (Pseudonomaretus) 456 manningense Bembidion (Plataphodes) 119 marginale Calosoma (Camegonia) 179 marginatus Pasimachus (Pasimachus) 362 marginatus Scaphinotus (Brennus) 457 marginicollis Elaphrus (Elaphrus) 265 marginicollis Lebia (Lebia) 301 marginipennis Clivina (Paraclivina) 208 marinensis Pterostichus (Hypherpes) 425 marinianum Bembidion (Hydriomicrus) 119 marinus Akephorus 39 marinus Scarites (Scarites) 464 maritimus Selenophorus (Selenophorus) 473 marshalli Pentagonica 367 martini Harpalus 284 matthewsii Zacotus 514 mattolensis Pterostichus (Hypherpes) 425 maxillosus Chlaenius (Pseudanomoglossus) 197 maxwelli Amara (Amara) 56 mckinleyi carneum Bembidion (Terminophanes) 119 mckinleyi mckinleyi Bembidion (Terminophanes) 119 meanyi giulianii Nebria (Catonebria) 333 meanyi lamarckensis Nebria (Catonebria) 333 meanyi meanyi Nebria (Catonebria) 333 meanyi sylvatica Nebria (Catonebria) 333 medius Brachinus (Neobrachinus) 155 megacephalus Diplochaetus 236 megacephalus Harpalus 285 megacephalus Stenolophus (Stenolophus) 488 megalops Platynus 372 melanarium Agonum (Agonum) 28

melanarius Pterostichus (Morphnosoma) 425 melancholicus hiemalis Dyschiriodes (Dyschiriodes) 250 melancholicus melancholicus Dyschiriodes (Dyschiriodes) 250 melanocephalus Calathus (Neocalathus) 171 melanopus Anisodactylus (Anisodactylus) 75 menetriesii campestris Pseudanophthalmus 393 menetriesii menetriesii Pseudanophthalmus 393 menetriesii Pterostichus (Hypherpes) 427 mercedianus Pterostichus (Hypherpes) 427 meridianus Acupalpus (Acupalpus) 15 meridionalis Neaphaenops tellkampfii 324 merkelii Scaphinotus (Pseudonomaretus) 457 merula Amara (Amara) 56 merula Anisodactylus (Gynandrotarsus) 75 metallescens Agonum (Agonum) 28 metallica Nebria (Catonebria) 334 mexicana Galerita (Progaleritina) 271 mexicana Notiobia (Anisotarsus) 341 mexicana see Stenocrepis (Stenous) 482 mexicana Stenocrepis (Stenous) (Stenocrepis (Crossocrepis) quatuordecimstriata) 482 mexicanum Bembidion 119 mexicanum Zuphium 515 mexicanus Amblygnathus 64 mexicanus Brachinus (Neobrachinus) 155 mexicanus Poecilus (Poecilus) 378 mexicanus Pogonodaptus 379 micans Loxandrus 315 micans Olisthopus 347 microamericanus Brachinus (Neobrachinus) 156 microcephalus Thalpius 501 mimus Bembidion 119 mimus Elaphrus (Elaphrus) 265 minor Loxandrus 315 minuta Anatrichis 65 minutus Anillodes 69 minutus Loxandrus 316 miranda Lebia (Lebia) 301 miscellus Pterostichus (Hypherpes) 427 misellus Tachys 495 mitchellensis Trechus (Trechus) 506 mobilis Brachinus (Neobrachinus) 156 modocianum Bembidion 120

Index moerens Agonum (Agonum) 29 moesta Lebia (Lebia) 302 moestus Pterostichus (Eosteropus) 427 moniliatum Calosoma (Microcallisthenes) 180 monilicornis Morion 323 montanus Bradycellus (Stenocellus) 164 montanus Dyschiriodes (Dyschiriodes) 250 montanus Pseudanophthalmus 393 monticola Calosoma (Microcallisthenes) 180 monticola Elaphropus 259 mordax Tachys 495 morio Ardistomis 83 morionides Pterostichus (Hypherpes) 427 mormon Bembidion 120 morrisonii Calosoma (Chrysostigma) 180 morula Clivina 208 morulum Bembidion (Semicampa) 120 muelleri Agonum (Agonum) 29 muiri Agonum (Agonum) 29 mundum Bembidion (Liocosmius) 120 mundus Loxandrus 316 municeps Selenophorus (Celiamorphus) 473 muscicola Bembidion (Semicampa) 120 musculis Amara (Amara) 56 mustus Selenophorus (Selenophorus) 473 mutatum Agonum 30 mutatum Bembidion (Bembidion) 121 mutoides Pterostichus (Gastrosticta) 427 mutus Pterostichus (Bothriopterus) 427 myops Clivina (Clivina) 209 myrmecodes Rhadine 446 nana inornata Tachyta (Tachyta) 496 nana kirbyi Tachyta (Tachyta) 496 nanellus Acupalpus (Acupalpus) 15 nantahalae Trechus (Microtrechus) 506 nanulus Selenophorus (Celiamorphus) 473 navajo Nebria (Reductonebria) 334 nearcticus Poecilus (Derus) 379 nebraskense Bembidion 121 nebraskensis Chlaenius (Chlaeniellus) 197 nebulosus Apenes (Apenes) 81 nebulosus Bradycellus (Stenocellus) 164 nebulosus Elaphropus 259 nebulosus Trechus (Microtrechus) 506 neglectus Brachinus (Neobrachinus) 156

571

neglectus Bradycellus (Stenocellus) 164 neglectus Cymindis (Cymindis) 226 nelsoni Pseudanophthalmus 393 nemoralis Carabus (Archicarabus) 188 nemoralis Chlaenius (Chlaeniellus) 198 nemoralis Notiophilus 345 neobrunneus Pterostichus (Hypherpes) 428 neocoerulescens Bembidion 121 neomexicana Amara (Bradytus) 57 neopulchellus Badister (Badister) 88 neovalidus Schizogenius (Schizogenius) 468 nevadense Bembidion 122 nevadensis Apristus 82 nexa Amara (Amara) 57 nickajackensis Pseudanophthalmus 394 niger Chlaenius (Agostenus) 198 nigerrimus Anisodactylus (Anisodactylus) 76 nigerrimus Bradycellus (Stenocellus) 164 nigricapitata Lebia (Lebia) 302 nigriceps Agonum (Platynomicrus) 30 nigriceps Bradycellus (Stenocellus) 164 nigriceps Perigona 368 nigricornis Pentagonica 367 nigrinus Bradycellus (Lipalocellus) 165 nigrinus Microlestes 322 nigripennis Helluomorphoides 294 nigripes Bembidion (Notaphus) 122 nigrita Anisodactylus (Anisodactylus) 76 nigritarsis Harpalus 285 nigrocaeruleus Pterostichus (Hypherpes) 428 nigrocoeruleum Bembidion 122 nigropiceum Bembidion (Lymneops) 122 nigrum Bembidion (Hirmoplataphus) 122 nitens Notiophilus 345 nitidicollis Sphaeroderus nitidicollis 480 nitidipennis Notiobia (Anisotarsus) 341 nitidula Coptodera 213 nitidulus Loxandrus 316 nitidum Bembidion (Eurytrachelus) 123 nitidum Omophron 349 nitidus Bradycellus (Liocellus) 165 nivalis Anisodactylus (Anadaptus) 77 nivalis gaspesiana Nebria (Boreonebria) 334 nivalis nivalis Nebria (Boreonebria) 334 nivalis Pterostichus (Cryobius) 428

572

André Larochelle & Marie-Claude Larivière

nivalis Rhadine446 noctivagaRhadine446 nogalesium Bembidion 123 nonnitens Cyclotrachelus (Evarthrus) 219 nortoni Pseudanophthalmus 394 notatus Badister (Badister) 89 novaculosus Trechus (Microtrechus) 506 novella Amara (Bradytus) 57 novemstriatus Notiophilus 345 novus Pterostichus (Euferonia) 428 nubiculosum Bembidion (Notaphus) 123 nubifer Bradycellus (Stenocellus) 166 nudipenne Bembidion 123 nupera Amara (Amara) 57 nupera Diplocheila (Isorembus) 238 nutans Agonum (Circinalidia) 30 oberthueri Bembidion 123 obesa Amara (Percosia) 57 obesulus Elaphropus 259 obesulus Pterostichus (Gastrosticta) 429 obesus Akephorus 39 obliqua chuskae Nebria (Reductonebria) 334 obliqua obliqua Nebria (Reductonebria) 334 obliquatus Ardistomis 83 obliquulum Bembidion (Eurytrachelus) 124 obliquus Harpalus (Glanodes) 285 obliquus Scaphinotus (Brennus) 457 obliteratum Omophron 349 oblitus Paratachys (Tachys obliquus) 358 oblonga Anatrichis 66 oblongopunctata Agra oblongopunctata 39 oblonguloides Bembidion (Amerizus) 124 oblongulum Bembidion (Amerizus) 124 obnixus Harpalus 285 obrieni Calleida (Calleida) 173 obscurellum Bembidion (Peryphus) 124 obscuripenne Bembidion 125 obscuromaculatum Bembidion (Notaphus) 125 obscurus Cymindis (Pinacodera) 226 obsidianus Discoderus (Discoderus) 241 obsidianus Pterostichus (Hypherpes) 429 obsoleta Sericoda 477 obsoletum Calosoma (Chrysostigma) 180 obsoletus Chlaenius (Chlaeniellus) 198 obsoletus Pasimachus (Pasimachus) 362

obtusa Diplocheila (Isorembus) 238 obtusangulum Bembidion 125 obtusidens Bembidion 125 obtusum Bembidion (Phyla) 125 obtusus Badister (Badister) 89 obtusus Bradycellus (Liocellus) 166 obtusus Dicheirus 235 obtusus Pelmatellus (Pelmatellus) 365 obtusus Trechus (Trechus) 507 ocalensis Scarites (Scarites) 464 occidentalis Euphorticus 268 occidentalis Poecilus (Poecilus) 379 occidentalis Pseudanophthalmus 394 occultator Bembidion (Plataphodes) 126 occultus Elaphropus 259 occultus Pterostichus (Hypherpes) 429 ochropezus Stenolophus (Stenolophus) 489 ochropus Harpalus 285 ochthocephalus Schizogenius (Schizogenius) 468 octopunctatum Agonum (Agonum) 31 ocularis Badister (Baudia) 89 ohioensis Pseudanophthalmus 394 olivaceus Elaphrus (Neoelaphrus) 266 olympica Gehringia 272 opacipennis Harpalus 286 opaculus Anisodactylus (Gynandrotarsus) 77 opaculus Calathus (Neocalathus) 171 opaculus Platynus (Platynus) 372 opacus Apenes (Apenes) 81 opalinus Selenophorus (Selenophorus) 474 operosum Bembidion (Notaphus) 126 ophryoderus Pterostichus (Gastrosticta) 429 oppressum Bembidion (Plataphus) 126 orbus Chlaenius (Chlaenius) 199 ordinarius Pterostichus (Hypherpes) 429 oregona Clivina (Clivina) 209 oregona Diplocheila (Isorembus) 238 oregonense Bembidion (Bembidion) 126 oregonensis Blethisa 149 oregonensis Trechus (Trechus) 507 oregonum Calosoma (Microcallisthenes) 180 oregonus Pterostichus (Bothriopterus) 429 oreophilus Scaphinotus (Brennus) 457 ornata Lebia (Lebia) 302 orthosulcatus Pseudanophthalmus 394

Index osculans Bembidion (Lionepha) 126 otiosa Amara (Amara) 58 otiosus Selenophorus (Selenophorus) 474 ovale Omophron 350 ovalipennis Pterostichus (Hypherpes) 429 ovata Amara (Amara) 58 ovatulus Platynus (Glyptolenopsis) 373 oviceps Pseudaptinus 403 ovipennis Brachinus (Neobrachinus) 156 ovipennis Nebria (Catonebria) 335 ovipennis Platynus (Platynus) 373 ovipennis Trechus (Trechus) 507 ovularis Anisodactylus (Gynandrotarsus) 77 ovulum Cyclotrachelus (Cyclotrachelus) 219 oxygonus Brachinus (Neobrachinus) 156 oxygonus Chlaenius (Chlaeniellus) 199 ozarkensis Rhadine 446 ozarkensis Schizogenius (Schizogenius) 468 pacificum Agonum (Agonum) 31 pacificus Dyschiriodes (Dyschiriodes) 250 packardi Pseudanophthalmus 394 pactinullus Loxandrus 316 pallens Plochionus (Plochionus) 375 palliatus Selenophorus (Selenophorus) 474 pallida Clivina (Clivina) 209 pallidipes Apenes (Apenes) 81 pallidulus Leptotrachelus 309 pallidus Brachinus (Neobrachinus) 156 pallidus Pseudanophthalmus 394 pallidus Tachys 495 pallidus Tetragonoderus 499 pallipennis Dyschiriodes (Dyschiriodes) 251 pallipennis Perigona 368 pallipes Agonum (Olisares) 31 pallipes Amara (Zezea) 58 pallipes Nebria (Reductonebria) 335 palmi Pterostichus (Feronina) 429 palosverdes Bembidion (Cillenus) 127 palustre Agonum (Europhilus) 32 palustris Notiophilus 345 panticulatus Pterostichus (Hypherpes) 430 papago Helluomorphoides 294 papagonis Discoderus (Discoderus) 241 paradisi Nebria (Boreonebria) 335 paradoxus Pseudanophthalmus 394

573

parafaber Cyclotrachelus (Cyclotrachelus) 219 parallela Lachnocrepis 295 parallela Pseudomorpha 404 parallelepipedus Abax (ater Abax) 13 parallelus Discoderus (Discoderus) 241 parallelus Loxandrus 316 parallelus Pogonus 379 parasimilis Pterostichus (Cryobius) 430 parasodalis Cyclotrachelus (Evarthrus) 219 paratus Harpalus (Pseudoophonus) 286 parca Pachyteles (Goniotropis) 356 parens Pterostichus (Hypherpes) 430 parextimum Agonum (Stictanchus) 32 parilis Discoderus (Selenalius) 242 parisiana Scaphinotus (Scaphinotus) 457 parmarginatus Platynus (Platynus) 373 parmatus Olisthopus 347 partiarius Acupalpus (Tachistodes) 15 parviceps Badister (Baudia) 90 parviceps Elaphrus (Elaphrus) 266 parviceps Stenolophus (Agonoleptus) 489 parvicolle Calosoma (Camegonia) 181 parvicollis Loxandrus 316 parvicollis Pseudanophthalmus 395 parvicornis Tachyta (Tachyta) 497 parvulus Elaphropus 259 parvulus Loxandrus 316 parvus Pseudanophthalmus 395 patruele Bembidion (Notaphus) 127 patruelis Amara (Amara) 59 patruelis Brachinus (Neobrachinus) 157 patruelis Chlaenius (Chlaenius) 199 patruelis Dyschiriodes (Eudyschirius) 251 patruelis Pterostichus (Phonias) 430 patruelis Scarites (Scarites) 464 paulus Pseudanophthalmus 395 pauperculus Acupalpus (Tachistodes) 16 paynei Pseudanophthalmus 395 pecki Platynus (Microplatynus) 373 pectita Lebia (Lebia) 302 pedicellatum Bembidion (Bembidion) 127 pedicularius Selenophorus (Selenophorus) 474 peigani Technophilus croceicollis 498 pennsylvanica Amara (Curtonotus) 59

574

André Larochelle & Marie-Claude Larivière

pennsylvanicus blanditus Chlaenius (Chlaeniellus) 199 pennsylvanicus pennsylvanicus Chlaenius (Chlaeniellus) 199 pensylvanica Colliuris (Cosnania) 211 pensylvanicus Harpalus (Pseudoophonus) 286 pensylvanicus Pterostichus (Bothriopterus) 431 perbrevicolle Bembidion 127 peregrinator Calosoma (Camedula) 181 peregrinus Discoderus (Discoderus) 242 peregrinus Stenolophus (Stenolophus) 489 pergracilis Pterostichus (Hypherpes) 431 perita Lebia (Lebia) 303 perlevis Rhadine 446 permundus Pterostichus (Abacidus) 431 pernotum Bembidion 128 peropacus Calathus (Neocalathus) 172 perpallida Lebia (Lebia) 303 perplexus Brachinus (Neobrachinus) 157 perpolitus Selenophorus (Selenophorus) 475 persephone Rhadine 446 pertenuis Rhadine 446 pertinax Chlaenius (Chlaeniellus) 200 perversus Dyschiriodes (Dyschiriodes) 251 petersi biedermani Scaphinotus (Scaphinotus) 458 petersi catalinae Scaphinotus (Scaphinotus) 458 petersi corvus Scaphinotus (Scaphinotus) 458 petersi grahami Scaphinotus (Scaphinotus) 458 petersi kathleenae Scaphinotus (Scaphinotus) 458 petersi petersi Scaphinotus (Scaphinotus) 458 petrosum attuense Bembidion (Peryphus) 128 petrosum petrosum Bembidion (Peryphus) 128 petrunkevitchi Pseudanophthalmus 395 phaeocerus Brachinus (Neobrachinus) 157 pholeter Pseudanophthalmus 395 picea Anatrichis 66 picea Coptodera 213 piceolum Agonum (Platynomicrus) 32 piceolus Loxandrus 316 piceus Dicheirus 235 piceus Dromius (Dromius) 243 piceus Oodinus 353 piceus Psydrus 405 picicornoides Agonum (Europhilus) 32 picipes Bradycellus (Stenocellus) 166

piciventris Loxandrus 317 picta picta Colliuris (Cosnania) 212 picticornis Pentagonica 367 picticornis Tetraleucus 499 pilatei Bembidion 128 pilatei Technophilus 498 pilicornis Loricera pilicornis 311 pilosus Cymindis (Cymindis) 226 pilosus Dyschiriodes (Dyschiriodes) 251 pilosus Pseudanophthalmus 395 pimalicus Chlaenius (Eurydactylus) 200 pimalis Lebia (Loxopeza) 303 pimanum Bembidion (Notaphus) 128 pinguedineus Pterostichus (Cryobius) 432 pinguis Discoderus (Discoderus) 242 piperi Nebria (Catonebria) 335 pisgahensis Trechus (Microtrechus) 507 pitychrous Anisodactylus (Anadaptus) 77 piute piute Nebria (Catonebria) 335 piute sevieri Nebria (Catonebria) 336 piute utahensis Nebria (Catonebria) 336 placeranum Bembidion (Plataphodes) 128 placerensis Pterostichus (Hypherpes) 432 placerum Calosoma (Microcallisthenes) 181 placidum Agonum (Agonum) 33 plagiatum Bembidion 128 planatum Bembidion (Plataphus) 129 planatus Diplochaetus 236 planatus Dyschiriodes (Dyschiriodes) 252 planatus Metrius contractus 320 planatus Pogonistes 379 planctus Pterostichus (Hypherpes) 432 planicollis Clivina (Clivina) 209 planipennis Cymindis (Cymindis) 227 planipennis Selenophorus (Selenophorus) 475 planiusculum Bembidion (Plataphus) 129 planulata Calleida (Calleida) 173 planulatus Schizogenius (Schizogenius) 468 planuloides Schizogenius (Schizogenius) 468 planum Bembidion (Trichoplataphus) 129 planus Pterostichus (Cryobius) 432 platicollis Cymindis (Pinacodera) 227 platyderus Chlaenius (Chlaenius) 200 platynoides Bembidion (Peryphanes) 129 platynoides Calleida (Calleida) 173

Index plebejus Stenolophus (Stenolophus) 489 plenalis Harpalus 287 pleuritica Lebia (Lebia) 303 plumasensis Rhadine447 pluripunctatus Schizogenius (Schizogenius) 468 plutonicus Pterostichus (Hypherpes) 432 poculare Bembidion (Cyclolopha) 130 polita Diacheila 229 politum Bembidion (Trepanedoris) 130 politus Bradycellus (Liocellus) 166 politus Dicaelus (Paradicaelus) 231 politus politus Dyschiriodes (Dyschiriodes) 252 pomona Amara (Amara) 59 pomonae Trechus (Trechus) 507 poncei Harpalus (Pseudoophonus) 288 pontis Pseudanophthalmus 395 poppii schalleri Bembidion (Peryphus) 130 porosus Anisodactylus (Anadaptus) (Anisodactylus viridescens) 77 postica Clivina (Paraclivina) 209 postremum Bembidion 130 potomaca Paratachys 358 potomacus potomacus Pseudanophthalmus 396 potomacus senecae Pseudanophthalmus 396 praecinctum Bembidion (Bembidion) 130 praetermissus Pseudanophthalmus 396 praeustus Helluomorphoides praeustus 294 prasinus Chlaenius (Chlaenius) 200 praticola Bembidion (Semicampa) 130 pratti Axinopalpus 87 pravitubus Loxandrus 317 primus Pterostichus (Monoferonia) 433 princeps Pseudanophthalmus 396 productus Pseudanophthalmus 396 prognathus Platynus 373 prominens Calosoma (Camegonia) 181 properans Bembidion (Metallina) 131 propinquum Agonum (Agonum) 33 protensiformis Pterostichus (Hypherpes) 433 protensipennis Pterostichus (Hypherpes) 433 protractum Calosoma (Callitropa) 181 protractus Harpalus (Pseudoophonus) 288 protractus Pterostichus (Hypherpes) 433 providens Harpalus (Euharpalops) (Harpalus viduus) 288

575

provoensis Bradycellus (Stenocellus) 166 proximus Loxandrus 317 proximus Paratachys 358 pseudagricola Anisodactylus (Anisodactylus) 78 pseudamericanum Zuphium 515 pseudobrunnea Amara (Amara) 59 pseudocautum Bembidion (Trepanedoris) 131 pseudoerasum Bembidion (Lionepha) 131 pseudopiceus Oodinus 353 pterostichina Amara (Curtonotus) 60 puberulus Brachinus (Neobrachinus) 157 pubescens Atranus 85 pubescens Euphorticus 268 pubescens intrepidus Pseudanophthalmus 396 pubescens pubescens Pseudanophthalmus 396 pugetanus Apristus 82 pulchella Lebia (Lebia) 303 pulchellus Badister (Badister) 90 pulchellus Tachys 495 pumicatus Stomis (Stomis) 492 pumila Lebia (Lebia) 304 pumilus Acupalpus (Acupalpus) 16 pumilus Dyschiriodes (Dyschiriodes) 252 pumilus Paratachys 358 pumilus pumilus Pterostichus (Hypherpes) 433 pumilus willamettensis Pterostichus (Hypherpes) 433 punctata Calleida (Calleida) 173 punctatissimus Pterostichus (Lenapterus) 434 punctatostriatum Bembidion (Bracteon) 131 punctatulus Athrostictus 85 punctatus Pseudanophthalmus 396 punctatus Scaphinotus (Brennus) 458 puncticeps Harpalus (Ophonus) 288 puncticeps Ophonus 353 puncticollis Bradycellus (Stenocellus) 166 puncticollis Semiardistomis 476 punctifer Cymindis (Pinacodera) 227 punctiforme Agonum (Olisares) 34 punctiger Cymindis (Pinacodera) 228 punctigera Clivina (Clivina) 209 punctiventris Pterostichus (Gastrosticta) 434 punctulata Calleida (Calleida) 174 punctulata Clivina (Clivina) 210 punctulatus Acupalpus (Anthracus) 16

576

André Larochelle & Marie-Claude Larivière

punctulatus Pasimachus (Pasimachus) 362 purgatus Bradycellus (Stenocellus) 166 purpurans Elaphrus (Elaphroterus) (Elaphrus pallipes) 266 purpurascens Notiobia (Anisotarsus) 341 purpurata Nebria (Catonebria) 336 purpuratus darlingtoni Dicaelus (Dicaelus) 232 purpuratus purpuratus Dicaelus (Dicaelus) 232 purpuratus quadratus Dicaelus (Dicaelus) 232 purpuratus splendidus Dicaelus (Dicaelus) 233 purpurea Calleida (Calleida) 174 purpureus Chlaenius (Chlaenius) 200 purpuricollis Chlaenius purpuricollis 201 pusillus Chlaenius (Anomoglossus) 201 pusillus Oxypselaphus (Agonum puncticeps) 355 pusillus Pseudanophthalmus 397 pusio Microlestes 322 pusio Olisthopus 348 pusio Pseudanophthalmus 397 puteanus Pseudanophthalmus 397 putus Pterostichus (Gastrosticta) 434 pygmaeus Nomius 340 pygmaeus Schizogenius (Schizogenius) 469 pygmaeus Thalpius 501 quadratulum Bembidion (Hydriomicrus) 132 quadratus Anchomenus 66 quadratus Pseudanophthalmus 397 quadriceps Scarites (Scarites) (Scarites substriatus) 464 quadricollis Blethisa 149 quadrifoveolatum Bembidion (Plataphodes) 132 quadrimaculatum Agonum (Deratanchus) 34 quadrimaculatum dubitans Bembidion (Bembidion) 132 quadrimaculatum oppositum Bembidion (Bembidion) 132 quadrimaculatus Dyschiriodes (Paradyschirius) 252 quadrinotatus Anchonoderus 67 quadripennis Brachinus (Neobrachinus) 157 quadripunctata Sericoda 477 quadristriatus Trechus (Trechus) 508 quadrulum Bembidion (Hirmoplataphus) 133 quatuordecimstriata Stenocrepis (Crossocrepis) 482 quenseli Amara (Amara) 60

quinquepunctatum Agonum (Agonum) 34 rapidum Bembidion (Notaphus) 133 rectangula Amara (Amara) 60 rectangulus Acupalpus (Philodes) 17 rectangulus Loxandrus 317 rectangulus Paratachys 359 recticolle Bembidion (Hirmoplataphus) 134 rectus Loxandrus 317 reflexus Badister (Baudia) 90 reflexus Loxandrus 317 regressus Harpalus (Glanodes) 288 regularis Scaphinotus (Pseudonomaretus) 459 reichardti Galerita (Progaleritina) 271 relictus Pterostichus (Euferonia) 434 relictus Scaphinotus (Pseudonomaretus) 459 remissus Stenolophus (Stenolophus) 490 renoanum Bembidion 134 renoicus Elaphropus 260 restrictus Pterostichus (Hypherpes) 434 retractum Agonum (Europhilus) 35 retractus Harpalus (Plectralidus) 289 reversus Harpalus (Euharpalops) (Harpalus funerarius) 289 rewolinskii Harpalus 289 rhodeanus Paratachys 359 richardsoni Opisthius 354 rickseckeri Cychrus hemphillii 214 rickseckeri Phrypeus 369 ridingsii intermedius Scaphinotus (Steniridia) 459 ridingsii monongahelae Scaphinotus (Steniridia) 459 ridingsii ridingsii Scaphinotus (Steniridia) 459 rigidulum Agonum (Olisares) 35 riparius Pterostichus (Cryobius) 435 riparius Selenophorus (Selenophorus) 475 rittmani Pseudanophthalmus 397 rivalis Bradycellus (Stenocellus) 166 riversi Scaphinotus (Brennus) 460 roanicus Trechus (Trechus) 508 robusticolle Bembidion (Odontium) 134 robustum Omophron 350 robustus Loxandrus 317 robustus piceus Discoderus (Discoderus) 242 robustus Pseudanophthalmus 397 robustus robustus Discoderus (Discoderus) 242

Index rogersae Pseudanophthalmus 397 rolandi Bembidion (Trichoplataphus) 134 roosevelti Bembidion (Semicampa) 134 rosenbergi Trechus (Microtrechus) 508 rossi Loxandrus 317 rossi Rhadine 447 rosslandicum Bembidion (Plataphodes) 135 rostratus Pterostichus (Cylindrocharis) 435 rothi Pterostichus (Hypherpes) 435 rotundatus Pseudanophthalmus 397 rotundatus Stenolophus (Agonoleptus) 490 rotundicollis Polpochila (Polpochila) 380 rotundicollis Stenolophus (Agonoleptus) 490 rubens Trechus (Trechus) 508 rubicunda Clivina (Antroforceps) 210 rubiginosum Bembidion (Semicampa) 135 rubra Rhadine 447 rubrica Amara (Amara) 60 rubricauda Elaphropus 260 rubripes Harpalus (Harpalus) 289 rubripes Stereocerus 492 rudis Anisodactylus (Anadaptus) 78 rudis Pelophila 366 rufa Clivina 210 rufibarbis Harpalus (Ophonus) 290 rufibarbis Ophonus 353 ruficauda Chlaenius (Callistometus) 201 ruficollis grandicollis Calathus (Neocalathus) 172 ruficollis ignicollis Calathus (Neocalathus) 172 ruficollis Pseudomorpha 404 ruficollis ruficollis Calathus (Neocalathus) 172 rufinum Bembidion (Plataphus) 135 rufipes Agonum (Olisares) 35 rufipes Harpalus (Pseudoophonus) 290 rufipes Stenomorphus californicus 491 rufiventris Platynus 374 rufopleura Lebia (Lebia) 304 rufotestacea Polyderis 381 rufotinctum Bembidion (Pseudoperyphus) 135 rufulus Thalpius 501 rufus Oxydrepanus 354 rugiceps incipiens Scaphinotus (Brennus) 460 rugiceps rugiceps Scaphinotus (Brennus) 460 rugicollis Diplous 240 rugicollis Stenolophus (Agonoderus) 490

577

rugipennis Brachinus (Neobrachinus) 157 rupestris Bradycellus (Stenocellus) 166 rupicola Bembidion (Peryphus) 135 ruscarius Elaphrus (Elaphrus) 266 russelli Rhadine 447 rusticum lenensoides Bembidion (Plataphus) 136 rusticum rusticum Bembidion (Plataphus) 136 rusticus Anisodactylus (Gynandrotarsus) 78 rutilus Diplochaetus (Diplochaetus lecontei) 236 saccisecundaris Loxandrus 318 sagax Paratachys 359 sahlbergii modoc Nebria (Reductonebria) 336 sahlbergii sahlbergii Nebria (Reductonebria) 336 sahlbergii triad Nebria (Reductonebria) 336 salebratum Bembidion (Hirmoplataphus) 136 salinarium Bembidion (Peryphodes) 136 salivagans Dyschiriodes (Dyschiriodes) 253 sallei Calybe (Ega) 185 sallei Cyclotrachelus (Evarthrus) 220 sallei Panagaeus 356 sallei Schizogenius (Schizogenius) 469 sanctaecrucis Anisodactylus (Anadaptus) 79 sanctipauli Pseudanophthalmus 398 sanjuanensis Amara (Amara) 61 saphyrinus Loxandrus 318 sasajii Clivina (Antroforceps) 210 satanicus Trechus (Microtrechus) 509 satelles Bembidion 137 saturatus Elaphropus 260 sayanus Pterostichus (Gastrosticta) 435 sayi Calosoma (Castrida) 181 sayi Notiobia (Anisotarsus) (Anisotarsus piceus) 342 scalpta Lebia (Lebia) 304 scapula Lebia (Lebia) 304 scenicum Bembidion (Trepanedoris) 137 schaefferi Anchonoderus 67 schaefferi Aztecarpalus 87 schaefferi Calosoma (Microcallisthenes) 182 schaefferi Selenophorus (Selenophorus) 475 schaumii Oodes 352 schaumii Ardistomis 84 schaumii Sphaeroderus 480 scholasticus Pseudanophthalmus 398 schwarzi Amara (Bradytus) 61

578

André Larochelle & Marie-Claude Larivière

schwarzi beverlianna Nebria (Catonebria) 337 schwarzi Pseudomorpha 404 schwarzi saludae Trechus (Trechus) 509 schwarzi Scaphinotus (Maronetus) 460 schwarzi schwarzi Nebria (Catonebria) 337 schwarzi schwarzi Trechus (Trechus) 509 schwarzi scopulosus Trechus (Trechus) 509 scitula Amara (Zezea) 61 scitulus Paratachys 359 scitulus Poecilus (Poecilus) 379 scolopaceus Selenophorus (Selenophorus) 475 scopaeus Schizogenius (Schizogenius) 469 scopulinum Bembidion 137 scrutator Calosoma (Calodrepa) 182 scrutator Lophoglossus 309 scudderi Bembidion 137 sculptilis intricatus Dicaelus (Paradicaelus) 233 sculptilis Loxandrus 318 sculptilis sculptilis Dicaelus (Paradicaelus) 233 sculptilis upioides Dicaelus (Paradicaelus) 233 sculpturatum Bembidion (Odontium) 137 sculptus Dyschiriodes (Dyschiriodes) 253 sculptus Pterostichus (Abacidus) 436 scutellaris Pterostichus (Hypherpes) 436 scutilus Pseudanophthalmus 398 seclusus Pseudanophthalmus 398 sectator Elaphropus 260 sedulus Elaphropus 260 sejunctum sejunctum Bembidion (Peryphus) 138 sejunctum semiaureum Bembidion (Peryphus) 138 sejunctus Bradycellus (Stenocellus) 167 sejungendus Pterostichus (Hypherpes) 436 sellatus Dyschiriodes (Dyschiriodes) 253 sellatus Pericompsus 368 semicinctum Bembidion (Semicampa) 138 semilaeve Calosoma (Chrysostigma) 182 semiopacum Bembidion (Notaphus) 138 semiopacus Notiophilus 346 semipubescens Bradycellus (Lipalocellus) 167 semipunctatum Bembidion (Notaphus) 139 semistriatum Bembidion (Hydriomicrus) 139 semistriatus Notiophilus 346 semitinctus Stenolophus (Stenolophus) 490 septentrionis Patrobus (Patrobus) 365 sequax Paratachys 360

sequoiae Bembidion (Lionepha) 139 sequoyah Pseudanophthalmus 398 sera Amara (Amara) 61 seriatus Cymindis (Cymindis) 228 sericea Amphasia (Pseudamphasia) 65 sericeus Metrius contractus 320 sericeus regularis Chlaenius (Chlaenius) 201 sericeus sericeus Chlaenius (Chlaenius) 202 sericeus sierricola Chlaenius (Chlaenius) 202 sericeus viridifrons Chlaenius (Chlaenius) 202 sericus Pseudanophthalmus 398 serratus Carabus (Hemicarabus) 189 serripes Pterostichus (Hypherpes) 436 seticollis seticollis Schizogenius (Schizogenius) 469 setosum Piosoma 369 setosus Dyschiriodes (Dyschiriodes) 253 setosus Pterostichus (Hypherpes) 436 seximpressus Cyclotrachelus (Evarthrus) 220 sextoni Dyschiriodes (Dyschiriodes) 254 shilohensis boonensis Pseudanophthalmus 398 shilohensis mayfieldensis Pseudanophthalmus 398 shilohensis shilohensis Pseudanophthalmus 398 shulli Pterostichus (Pseudoferonina) 436 sidus Pseudanophthalmus 399 sierrablancae Nebria (Catonebria) 337 sierranus Notiophilus 346 sierranus Pterostichus (Hypherpes) 436 sierricola Bembidion (Plataphus) 139 sigillatus Cyclotrachelus (Evarthrus) 220 signata Phloeoxena (Oenaphelox) 369 silphoides Licinus 309 simile Agonum (Europhilus) 36 similis Anisodactylus (Anisodactylus) 79 similis Pterostichus (Cryobius) 437 simillimus Chlaenius (Chlaeniellus) 202 simplex Bembidion 139 simplex Calosoma (Chrysostigma) 183 simplex Pseudanophthalmus 399 simulans Pseudanophthalmus 399 simulator Notiophilus 346 sinaloae Stenomorphus 492 sinuaticollis Anillinus 68 sinuaticollis Selenophorus (Selenophorus) 475 sinuatus Anillodes 69 sinuatus Apenes (Apenes) 81

Index sinuosa Amara (Amara) 61 sinus Cyclotrachelus (Evarthrus) 220 siticum Bembidion (Trepanedoris) 140 smetanai Pterostichus (Pseudoferonina) 437 snowi roeschkei Scaphinotus (Scaphinotus) 460 snowi snowi Scaphinotus (Scaphinotus) 460 sodalicia Amara (Amara) 62 sodalis colossus Cyclotrachelus (Evarthrus) 220 sodalis lodingi Cyclotrachelus (Evarthrus) 221 sodalis sodalis Cyclotrachelus (Evarthrus) 221 solea Lebia (Lebia) 304 solidum Omophron 350 solitaris Harpalus (Harpalus fuliginosus) 290 solitarius Chlaenius (Chlaenius) 203 solivagus Pseudanophthalmus 399 somnulentus Harpalus (Harpalus pleuriticus; Harplus fallax) 291 soperi Pterostichus (Cryobius) 437 sordens Agonum (Europhilus) 36 sordidum Bembidion 140 spadicea Amara (Curtonotus) 62 spadiceus Harpalus (Euharpalops) (Harpalus carolinae) 291 spadix Paratachys 360 spathifer Pterostichus (Pseudoferonina) 437 spatulata sierrae Nebria (Catonebria) 337 spatulata spatulata Nebria (Catonebria) 337 spectabile Bembidion (Amerizus) 140 specularis Amara (Amara) 62 specus crinicollis Rhadine 447 specus gentilis Rhadine 447 specus specus Rhadine 447 speokoites Horologion 295 sphaericollis Dyschiriodes (Dyschiriodes) 254 sphaeroderum Bembidion (Cyclolopha) 140 sphodrinus Pterostichus (Hypherpes) 437 spinilunatus Loxandrus 318 splendidula Abaris 13 splendidulus Stenolophus (Stenolophus) 491 splendidum Calosoma (Calodrepa) 183 spoliatus Cyclotrachelus (Cyclotrachelus) 221 sponsor Amara (Curtonotus) 62 sponsor Pterostichus (Hypherpes) 437 sponsum Calosoma (Carabosoma) 183 spraguei Pterostichus (Hypherpes) 437

579

spretus Stenolophus (Stenolophus) 491 spuria Amara (Amara) 62 stantonensis Pterostichus (Cryobius) 438 stapedius Pterostichus (Hypherpes) 438 stapedius yosemitensis Pterostichus (Hypherpes) 438 steensensis Nebria (Catonebria) 337 steevesi Anillinus 68 steevesi Pseudanophthalmus 399 stenolophoides parallelus Pelmatellus (Pelmatellus) 365 stenolophoides stenolophoides Pelmatellus (Pelmatellus) 365 stenostomus Sphaeroderus stenostomus 481 stephani Harpalus (Glanodes) 292 stephensii Bembidion (Peryphanes) 140 stigmula Clivina (Paraclivina) 210 stillaguamish Bembidion (Plataphus) 141 straneoi Loxandrus 318 strenuus Dicheirus 235 strenuus Pasimachus (Pasimachus) 362 strenuus Pterostichus (Phonias) 438 striata Calleida (Calleida) 174 striatium Calosoma (Microcallisthenes) 183 striatopunctata Clivina (Paraclivina) 210 striatopunctata Diplocheila (Isorembus) 239 striatopunctatum Agonum (Circinalidia) 36 striatopunctatus Scaphinotus (Brennus) 460 striatum Tanystoma 497 striatus Pseudanophthalmus 399 striola Bembidion 141 stupkai Trechus (Microtrechus) 509 stygicus Patrobus (Patrobus) 365 stygicus Pterostichus (Euferonia) 438 suavis Bradycellus (Stenocellus) 167 subacutus Pterostichus (Gastrosticta) 439 subaeneum Calosoma (Microcallisthenes) 183 subaerarium Bembidion (Hirmoplataphus) 141 subangulata Aspidoglossa 85 subangustatum Bembidion (Peryphanes) 141 subarcticus Dyschiriodes (Dyschiriodes) 254 subasperatum Calosoma (Microcallisthenes) 183 subcarinatus Cymindis (Pinacodera) 228 subcordatus Bradycellus (Stenocellus) 167 subdeletus Apristus 83

580

André Larochelle & Marie-Claude Larivière

subdola Lebia (Loxopeza) 305 subgrandis Lebia (Loxopeza) 305 sublaevis Brachinus (Neobrachinus) 157 sublaevis Dyschiriodes (Dyschiriodes) 254 sublaevis Pasimachus (Pasimachus) 362 sublaevis Pterostichus (Metallophilus) 439 sublustris Rhadine 447 submarginatus Piesmus 369 submarinus Badister (Baudia) 90 subrugosa Lebia (Lebia) 305 subsericeum Agonum (Agonum) 37 substrenuus Lophoglossus 310 substriatus Cyclotrachelus (Evarthrus) 221 subsulcatus Apristus 83 subsulcatus Pasimachus (Pasimachus) 363 subterranea mitchelli Rhadine 447 subterranea subterranea Rhadine 448 subterraneus Scarites (Scarites) 464 subtilis Scaphinotus (Brennus) 461 subtilis Trechus (Microtrechus) 510 subtinctus Amblygnathus 64 subtropicus Selenophorus (Celiamorphus) 476 subviolaceus Discoderus (Discoderus) 242 suffusus Dicaelus (Liodicaelus) 233 suffusus Pterostichus (Hypherpes) 439 sulcatum Tanystoma 497 sulcifrons Schizogenius (Schizogenius) 469 sulcipenne Agonum (Stereagonum) 37 sulcipenne hyperboroides Bembidion (Plataphus) 141 sulcipenne prasinoides Bembidion (Plataphus) 141 sulcipennis Clivina (Paraclivina) 211 superciliosus Pterostichus (Eosteropus) 439 superioris Agonum (Europhilus) 37 supplex Bradycellus (Stenocellus) 168 surgens Pterostichus (Cryobius) 439 suturale Agonum (subsericeum Agonum) 37 suturalis Loxandrus 318 suturalis Nebria (Reductonebria) 337 sycophanta Calosoma (Calosoma) 183 sylvaticus Notiophilus 346 sylvaticus Pseudanophthalmus 399 sylvosus Carabus (Tanaocarabus) 189 symbolicus Discoderus (Discoderus) 242 symetricus Bradycellus (Stenocellus) 168

tadorcus Harpalus (Opadius) 292 taedatus agassii Carabus (Tomocarabus) 190 taedatus taedatus Carabus (Tomocarabus) 190 taeniatus Loxandrus 318 tahoensis Bradycellus (Liocellus) 168 tahoensis Elaphropus 260 tahoensis Pterostichus (Hypherpes) 439 talequah Trechus (Microtrechus) 510 tantillus Bradycellus (Stenocellus) 168 tareumiut Pterostichus (Cryobius) 440 tarsalis Pterostichus (Hypherpes) 440 tartarea Amara (Curtonotus) 62 tartaricus Lophoglossus 310 tellkampfii Neaphaenops tellkampfii 324 templetoni Pseudanophthalmus 399 tenebrionella Amara (Amara) 62 tenebroides Pseudomorpha 405 tenebrosa mckenziei Rhadine 448 tenebrosa tenebrosa Rhadine 448 tenebrosus Discoderus (Discoderus) 242 tenebrosus Pseudanophthalmus 400 tener Acupalpus (Anthracus) 17 tenesensis Pseudanophthalmus 400 tennessensis tauricus Trechus (Microtrechus) 510 tennessensis tennessensis Trechus (Microtrechus) 510 tenue Agonum (Agonum) 37 tenuicollis Brachinus (Neobrachinus) 158 tenuicollis Platynus (Platynus) 374 tenuicollis Pseudaptinus 403 tenuipes Rhadine 448 tenuis blatchleyi Pseudanophthalmus 400 tenuis jeanneli Pseudanophthalmus 400 tenuis morrisoni Pseudanophthalmus 400 tenuis Pterostichus (Pseudomaseus) 440 tenuis Scaphinotus (Maronetus) 461 tenuis stricticollis Pseudanophthalmus 400 tenuis tenuis Pseudanophthalmus 400 tenuiscapus Trechus (Trechus) 510 tenuispinus Dyschiriodes (Eudyschirius) 255 tepidum Calosoma (Chrysostigma) 184 terminata Notiobia (Anisotarsus) 342 terminatus Dyschiriodes (Paradyschirius) 255 termitiformis Stomis (Neostomis) 492 terricola terricola Laemostenus (Pristonychus) 296

Index tessellatum Omophron 350 testacea Rhadine 448 testaceus Acupalpus (Tachistodes) 17 testaceus Pterostichus (Hypherpes) 440 teter Dicaelus (Paradicaelus) 234 tetracolum Bembidion (Peryphus) 142 tetraspilotus Mochtherus 323 texana Amara (Amara) 62 texana Stenocrepis (Stenocrepis) 482 texanum Agonum (Stictanchus) 38 texanum Bembidion (Peryphanes) 142 texanus Anisodactylus (Gynandrotarsus) 80 texanus Brachinus (Neobrachinus) 158 texanus Chlaenius (Chlaeniellus) 203 texanus Discoderus (Discoderus) 243 texanus Harpalus (Pseudoophonus) 292 texanus Helluomorphoides 294 texanus Poecilus (Poecilus) 379 texanus Pogonus 380 texanus Scarites (Scarites) 465 texensis Cyclotrachelus (Cyclotrachelus) 222 thomasi Pseudanophthalmus 400 thoracica Amara (Curtonotus) 63 thoracicus Apristus 83 thoreyi Agonum (Europhilus) 38 tibialis Schizogenius (Schizogenius) 470 tibialis Stenocrepis (Stenous) 483 tigrinum Bembidion (Leuchydrium) 142 tiliaceoradix Pterostichus (Cryobius) 440 timidum Bembidion 142 timidus Dyschiriodes (Dyschiriodes) 255 timidus Plochionus (Menidius) 375 tiresias Pseudanophthalmus 401 tomentosus tomentosus Chlaenius (Eurydactylus) 203 tomentosus zunianus Chlaenius (Eurydactylus) 204 tonitru Trechus (Microtrechus) 510 torrida Amara (Curtonotus) 63 torvus deceptus Cyclotrachelus (Evarthrus) 222 torvus torvus Cyclotrachelus (Evarthrus) 222 transfluvialis Pseudanophthalmus 401 transparens Bembidion (Diplocampa) 143 transversale Bembidion 143 transversus Badister (Trimorphus) 90

581

trechiforme Bembidion (Trechonepha) 143 trepidus Selenophorus (Selenophorus) 476 tricarinatus Scaphinotus (Steniridia) 461 tricolor Lebia (Loxopeza) 305 tricolor tricolor Chlaenius (Chlaeniellus) 204 tricolor vigilans Chlaenius (Chlaeniellus) 204 tridentatus Dyschiriodes (Eudyschirius) 255 trifaria pasquineli Nebria (Catonebria) 338 trifaria trifaria Nebria (Catonebria) 338 trifoveolatus Platynus (Platynus) 374 trigeminum Agonum (Agonum) 39 trinarius Pterostichus (Bothriopterus) (Pterostichus ohionis) 440 trinitensis Pterostichus (Hypherpes) 441 tripunctatus Elaphropus 260 tristis Pterostichus (Hypherpes) 441 triviale Bembidion 143 trivittatus Euproctinus (Neoeuproctus) 269 troglodytes Pseudanophthalmus 401 trossulus Elaphrus (Elaphrus) (Elaphrus (Elaphrus) parviceps) 267 truncaticollis Carabus (Aulonocarabus) 190 truncatus Dyschiriodes (Dyschiriodes) 255 tuberculatus Cychrus 214 tuberculatus Elaphrus (Elaphrus) 267 tuberculofemoratus Pterostichus (Hypherpes) 441 tuckaleechee Trechus (Microtrechus) 510 tuckeri Apristus 83 tuckeri Lebia (Lebia) 306 tullahoma Pseudanophthalmus 401 tumescens Pterostichus (Gastrosticta) 441 turbata Amara (Amara) 63 turmaduodecima Nebria (Boreonebria) 338 turneri Anillinus 69 tusquitee Trechus (Microtrechus) 511 ulkei Bembidion 144 umbra Rhadine 448 umbratilis Pseudanophthalmus 401 umbratum Bembidion (Notaphus) 144 umbripennis Paratachys 360 umiatense Bembidion 144 uncifer Trechus (Microtrechus) 511 undulata Diplocheila (Isorembus) 239 unicarum Gastrellarius 272 unicolor Cyclotrachelus (Cyclotrachelus) 222

582

André Larochelle & Marie-Claude Larivière

unicolor Cymindis (Cymindis) 228 unicolor dolosus Stenolophus (Agonoleptus) 491 unicolor floridanus Scaphinotus (Scaphinotus) 461 unicolor heros Scaphinotus (Scaphinotus) 461 unicolor shoemakeri Scaphinotus (Scaphinotus) 461 unicolor unicolor Scaphinotus (Scaphinotus) 462 unicolor unicolor Stenolophus (Agonoleptus) 491 unifasciatus Somotrichus 478 uniformis Loxandrus 318 unilobus Loxandrus 319 unionis Pseudanophthalmus 401 unipunctatus Dyschiriodes (Eudyschirius) 255 uniseriatus Cymindis (Cymindis) 228 unistriatus Scaphinotus (Maronetus) 462 utahense Bembidion (Lymneops) 144 utahensis Axinopalpus 87 vafer Chlaenius (Chlaeniellus) 204 valentinei Anillinus 69 valentinei Pseudanophthalmus 401 valentinei Trechus (Microtrechus) 511 valentinei Troglanillus 513 vanburenensis Pseudanophthalmus 401 vandykei Badister (Badister) 91 vandykei Bembidion (Plataphus) 144 vandykei Platidiolus 370 vandykei Pseudomorpha 405 vandykei Pterostichus (Hypherpes) 442 vandykei Scaphinotus (Scaphinotus) 462 vandykei Trechus (Microtrechus) 511 vandykei vandykei Nebria (Catonebria) 338 vandykei wyeast Nebria (Catonebria) 338 vaporariorum Cymindis (Cymindis) 228 variabilipes Chlaenius (Chlaeniellus) 205 varicornis Eucaerus (Eucaerus) 267 varidens Dyschiriodes (Eudyschirius) 256 variegatum Bembidion (Notaphus) 144 vegasensis Amara (Bradytus) 63 velocipes Loxandrus 319 velox Loxandrus 319 velutinus Brachinus (Neobrachinus) 158 velutinus Scaphinotus (Stenocantharus) 462 ventralis Harpalus (Euharpalops) 292 ventralis Pterostichus (Gastrosticta) 442 ventricosus Scaphinotus (Brennus) 462 ventricosus ventricosus Pterostichus (Cryobius) 442

ventricosus Paratachys 360 ventus Pseudanophthalmus 402 vermiculosus Pterostichus (Lenapterus) 442 vernicatus Elaphropus (Tachys unionis) 261 vernilis Paratachys 360 vernix Lophoglossus 310 veronianus Bradycellus (Stenocellus) 168 versicolor Bembidion 145 versutum Bembidion (Notaphus) 145 verticalis Anisodactylus (Spongopus) 80 verus Trechus (Microtrechus) 511 vespertina Clivina (Semiclivina) 211 vespertinum Bembidion (Plataphus) 145 vexatus Pterostichus (Pseudoferonina) 442 viator Bembidion (Plataphodes) 145 viator Neaphaenops tellkampfii 325 vicarius Pseudanophthalmus 402 vicina Pseudomorpha 405 vicinus Pterostichus (Hypherpes) 442 viduus Chlaenius (Chlaenius) 205 viduus Scaphinotus (Scaphinotus) 462 vietinghoffii vietinghoffii Carabus (Megodontus) 191 vigilans Selenophorus (Selenophorus) 476 vile Bembidion (Emphanes) 146 vinctus Carabus (Carabus) 191 vinctus Cyclotrachelus (Cyclotrachelus) 222 vindicata Pseudomorpha 405 violaceus carolinae Scaphinotus (Steniridia) 463 violaceus violaceus Scaphinotus (Steniridia) 463 virescens Nebria (Nipponebria) 339 virescens Nipponebria (Vancouveria) 340 virginiae Anillinus 69 virginicus Pseudanophthalmus 402 virgo Tachys 495 viridans Pasimachus (Pasimachus) 363 viridescens Anisodactylus (Anadaptus) 80 viridicolle Bembidion 146 viridicollis Calleida (Philophuga) 174 viridipennis Brachinus (Neobrachinus) 158 viridipennis Calleida (Calleida) 174 viridipennis Lebia (Lebia) 306 viridis amoena Calleida (Philophuga) 175 viridis Elaphrus (Elaphrus) 267 viridis horni Calleida (Philophuga) 175

Index viridis klamathea Calleida (Philophuga) 175 viridis Lebia (Lebia) 306 viridis Semiardistomis 476 viridis viridis Calleida (Philophuga) 175 vitiosus Loxandrus 319 vittata Lebia (Lebia) 307 vittatus Axinopalpus 87 vittiger Tachys 496 vivax Elaphropus 261 vividum Bembidion 146 volatilis Amara (Amara) 63 vorax Paratachys 360 vulcanoides Brachinus (Neobrachinus) 159 vulneratus Loxandrus 319 vulpecula Bembidion (Notaphus) 146 vulpeculus Trichotichnus (Trichochichnus) 513 walkeri Anillodes 69 wallacei Pseudanophthalmus 402 wallowae Nebria (Reductonebria) 339 walteri Nelsonites 339

583

webbi Scaphinotus (Scaphinotus) 463 westcotti Trechoblemus 501 whitcombi Cyclotrachelus (Evarthrus) 222 wickhami Bembidion 146 wilcoxi Calosoma (Calodrepa) 184 wilkesii Calosoma (Microcallisthenes) 185 wingatei Bembidion (Amerizus) 146 woodi Pterostichus (Cryobius) 442 xanthopus Elaphropus 261 ybousqueti Pterostichus (Hypherpes) (Pterostichus (Hypherpes) parens) 442 yosemitensis Pterostichus (Hypherpes) 443 youngi donaldsoni Pseudanophthalmus 402 youngi youngi Pseudanophthalmus 402 yukonense Asaphidion 85 yukonum Bembidion (Peryphanes) 147 zephyrium Bembidion (Bracteon) 147 zioni oasis Nebria (Reductonebria) 339 zioni zioni Nebria (Reductonebria) 339 zunianus Pterostichus (Hypherpes) 443