Eriosyce. The Genus revised, amplified

Succulent

Plant

Research

Volume One

Eriosyce ( Cactaceae) The genus revised and amplified by

Fred Kattermann with contributions by Jonas M. Liithy, Robert S. Wallace David Hunt, Nigel Taylor and Daniela Zappi

edited by David Hunt & Nigel Taylor Royal Botanic Gardens Kew

1994

Eriosyce (Cactaceae): AUTHOR'S ACKNOWLEDGMENTS

© 1994 Fred K a t t e r m a n n and contributors

Alt rights reserved.

N o part of

this publication m a y

be

reproduced, in any form or by a n y m e a n s , without prior permission in writing from the Publisher.

ISBN 0 9517234 2 1 Published by David H u n t 5 4 Priory Road Richmond Surrey T W 9 3 D H England

Printed in E n g l a n d

The genua revined and

amplified

I want to thank all those who have helped me in the preparation of this book. I especially want to thank Heriberto and Adela Nichterlein and their family, of Vina del Mar, Chile. They provided my home away from home. It was always with enthusiasm that they put up with "el fanatico". Special mention must also be made of Hans Nichterlein, once an exchange student in our house, on whom I could always depend, just like a son. I am also very grateful to Sr Mario Pino, for many years the president of the Sociedad Cientffica de Valparaiso, a naturalist without equal: Sra Adriana Hoffmann, botanist, author, friend and travel companion, for sharing her knowledge and smoothing the way through Chilean bureaucracy; to Herr Walter Jung, cactophile, guide and friend, who made my first field expeditions possible; to Dr David Hunt and Nigel Taylor (Royal Botanic Gardens, Kew) for their editorial help and to Dr Danicla Zappi (Kew) for redrawing the distribution maps; to Prof. Otto Zoellner, Dept of Botany, Universidad Catolica de Valparaiso, who accompanied me on many of my trips; to Dr E.F. Anderson, formerly of Whitman College, Walla Walla, Washington, and now at the Desert Botanical Garden, Phoenix, Arizona, for his advice and assistance in preparing most of the seed, plant and pollen SEM pictures, and especially to Christine Thomson at Whitman for her technical help; to Dr Steven Koppe from Montclair State College, New Jersey, for allowing me use of the SEM facility at that college; and last but not least to Dr Robert Wallace of Iowa State University for preparing and writing the cladistic analysis. Fred Kattermann 719 Route 519, Wantage, NJ 04761, U.S.A.

Succulent Plant

Research

Volume One

CONTENTS Summary - Zusammenfassung - Resumen

6

Erioayce (Cactaceae): The genus revised and amplified Fred Kattermann Introduction Previous study Historical landmarks Scope of present study General conclusions Outlline of proposed classification Characters used in classification Reproductive system Allied genera Uses and economic importance Threats to the survival of Eriosyce species Cultivation Selected bibliography Systematic treatment

7 7 8 10 10 11 12 17 17 20 20 21 23 25

APPENDICES I.

Validation of new names

117

II.

Eriosyce laui J. Luthy sp. nov Jonas M. Liithy

120

III. Phylogenetic analysis of Erioayce Robert S. Wallace

125

IV. Documentation of accepted names and synonyms David Hunt, Nigel Taylor & Daniela Zappi

131

Alphabetical list of synonyms Index of accepted taxa Index of illustrations

170 174 175

В

Eriosyce (Cactaceae):

Summary.

T h e g e n u s Eriosyce

The genus revised and amplified

Phil. (Cactaceae-NotocacteacJ is revised and amplified to include 3 3 species

and various infraspecific taxa previously classified in Neoporteria

sens. lat. A s circumscribed here

Eriosyce

ranges along the western aide of South A m e r i c a , from central Chile to south-western Peru, and on the e a s t side of the A n d e s in western A r g e n t i n a , from M e n d o z a to Salta. T h e data upon which this study is based derive from field observations and herbarium collections m a d e by the principal author ( K a t t e r m a n n ) . Bibliographical and nomenclatural details (Appendix IV» and a pbylogeneLic a n a l y s i s (Appendix III) are supplied b y various collaborating authors, and a new species, E. laui J. Luthy, is described in Appendix II. A study of seed-morphology utilizing the S c a n n i n g Electron Microscope is presented and taxa at all ranks are keyed out, described and illustrated by m e a n s of colour photographs and/or line drawings. T h e g e n u s is divided into two sections and several subsections and all new n a m e s and combinations required are published and listed ' A p p e n d i x 11: sect. Eriosyce Pyrrhocactus

<E. bulbocalyx.

fE. islayensis); limariensis, napina,

E. confinis.

E. odieri,

E. tenehrica.

E. strausiana,

and sect. Neoporteria E. crispa,

E. taltalensis,

E. garaventae. E. occidta).

E. laui) and subsect. Neoporteria

Zusammenfaasung.

with subsect. Eriosyce

E. umadeave.

Die G a t t u n g Eriosyce

\E. aurata. E. rodent iophila).

E. andreaeana,

with subsect. Horridocactus E. aspdlagae.

E. marksiana.

subsect. Chileosycc IE. chilensis,

E. villicumensis)

(E. recondita,

E. engleri,

<E. esmeraldana,

E. subgibbosa,

Islaya

E. kunzei,

E. heinrichiana,

E. aerocarpa,

E. sociabilis,

subsect.

and subsect.

E. vurvispina,

E. senilis,

E. E.

E.

krausii.

E.

villosa).

Phil. ' C a c t a c e a e - N o t o c a c t e a e ( wird revidiert und erweitert und

umfasst nun 3 3 A r t e n und verschiedene infraspezifische T a x a . die fruher zu Neoporteria worden sind. In der hier vorgeschlagenen U m s c h r e i b u n g ist die G a t t u n g Eriosyce

sens. lat. geslelll

auf der westlichen Spite

von S u d a m e r i k a zwischen Zentral-Chile und dem siidwestlichen Peru verbreitet, sowie dstlich der A n d c n im westlichen Argentinien von M e n d o z a bis Salta. Die dieser Studie zugrundeliegenden Duten wurden im Feld und aulgrund d e r H e r b a r b c l e g e des Hauptautnrs < K a t t e r m a n n ) erarbeitet. Hibliographische und n o m e n k l a torische Details i A n h a n g I V ) sowie die phylogenetische A n a l y s e l A n h a n g Ш ) wurden von verschiedenen Mitarbeitern beigesteuert, und die neue A r t E. laui J. Luthy wird beschrieben i A n h a n g III. Zusatzlich wird eine S E M - S t u d i e der S a m e n m o r p h o l o g i e vorgestellt. Die Taxa aller R a n g s l u f e n werden

aulgeschlusselt.

beschrieben und mit Fnrbbildern nder Zeichnungen illustriert. Die G a t t u n g wird in zwei Sektionen und verschiedene Subsektionen eingeteilt. und zahlreiche neue N a m e n und Kumbinationen werden publiziert und aufgelistet ( A n h a n g I): Sektion Eriosyce Pyrrhocactus Islaya

curvispina. engleri,

<E. bulbocalyx.

(E. islayensis); E. kunzei.

E. limariensis,

E. heinrichiana.

esmeraldana,

Resumen.

E. umadeave,

E. confinis,

E. napina.

E. aerocarpa.

E. subgibbosa.

m i t Suhsektion Eriosyce

E. strausiana.

und die Sektion Neoporteria E. odien.

E. krausii.

E. sociabilis.

El genero Eriosyce

E. senilis,

E. crispa.

E.

E. rodentiophilu),

E. vitlicumensisl

E. garaventae.

E. taltalensis.

E. tenebrica.

<E. aurata.

E. andreaeana,

mit der Subsektion Horridocactus E. occulta*,

Subsektion

und Subsektion <E. recondita,

E. aspdlagae,

E. IE.

E. laui) und mit der Subsektion Neoporteria

>E.

chilensis,

villosa).

Phil. (("actaceae-Notocacteaei h a sido revisado у ampliado, aclualmente incluye

3 3 especies у diversas categories infraespecificas q u e anteriormente fueron clasificadas en Neoporteria lat. Eriosyce

E.

E. marksiana,

der Subsektion Chileosyce

sens,

s e encuentra distribuido a lo largo del oeste de A m e r i c a del S u r . de la parte central de Chile

hasta el Sudoeste de Peru, t a m b i e n en el lado este de la Cordillera de los A n d e s , oeste de A r g e n t i n a , de M e n d o z a a S e l l a . Los datos f u n d a m e n l a l e s en el presente estudio fueron obtenidos por el principal autor < K a t t e r m a n n i en el cam no. у d o c u m e n l a d o s con materia) de h e r b a n o . Detalles de bibliugrafiay nomenclatura lApendice I V ) у la filogenia del grupo lApendice III) fueron contribuidos por varios eolaboradores. S e describe una especie nueva, E. laui

J. Luthy lApendice II). S e efectuaron estudios de morfologia de las semillas

utilizandn Microscopio Electronic*) de barrido. S e presentan claves de identifkaciun para todos los niveles. descripciones у

folografias a color y/o dibujos. El genero se subdivide en

dos secciones

subsccciones. los nombres у combinaciones n u e v a s se encuentran en el Apendice I: secc. Eriosyce Eriosyce

IE. aurata.

andreaeana, IE. recondita. marksiana,

E. villicumensis) E. curvispina, E. engleri.

iE. esmeralduna, subgibbosa.

E. rodentiophilai,

subsecc. Pyrrhocactus

у subsecc. Islaya <E. islayensis); E. kunzei.

E. heinrichiana.

E. aerocarpa.

E. sociabilis.

>E. bulbocalyx.

E. limariensis, E. napina.

E. krausn,

E. senilis.

E.

у secc. Neoporteria

E. confinis, E. (xlieri.

E. tenehrica,

villosal.

E. strausiana,

E. crispa.

E. taltalensis,

con

con subsecc.

E. garaventae, E. occulta),

varias

con subsecc.

E. umadeave,

subsecc.

E. laui) у subsecc. Neoporteria

E.

Harridwactus

E. aspdlagae,

E.

ChUemyce

<E. chilensis.

E.

Succulent

Plant

Research

Volume One

Eriosyce (Cactaceae): The genus revised and amplified Fred Kattermann Introduction Eriosyce, as here redefined, is a genus of 33 species of subglobular, globular, elongated or rarely almost columnar form. It can be recognized by the presence of wool and bristles on the pericarpel and tube (except for Eriosyce laui J. Luthy sp. nov., in which the flower is naked: see Appendix II), a hollow, dehiscent fruit, which has a basal opening, the seeds loose within and seeds with a small hilum (hilum-micropylar region or HMRi. The hilum rim at the micropyle of the seed is often modified by a thickening, forming a partial to complete annular structure around the micropyle. Floral characters vary somewhat within the genus. The flower-tube is funnelform and usually has some wool or bristles in the axils of the bract-scales. Some flowers are adapted for hummingbird pollination, and in some of these the flower is almost tubular. Eriosyce belongs to tribe Notocacteae, and occurs from central Chile, with its southernmost limit near Concepcion at 37°S, to southern Peru, where it reaches the Rio Omas at 14 S, and eastward into north-west Argentina, from Mendoza to Salta Map 1). C

Previous study The most recent taxonomic treatment of this group of plants was that of Backeberg 1958-62). The later treatment by F. Ritter (1980) can only be viewed as a catalogue at population descriptions because it lacks a key to species. Backeberg's treatment separated these plants into the genera Eriosyce, Islaya, Horridwactus, Neochilenia, Neoporteria and Pyrrhocactus. Donald & Rowley (1966) consolidated the above genera in Neoporteria, except for Eriosyce. In the past Eriosyce was always excluded from any revision of this South American group, as in the work of Donald & Rowley, although they did suggest that Eriosyce was closely related. In a complete study one cannot exclude Eriosyce. Nevertheless, their work laid the foundation for the present treatment. The group as a whole has had a chequered taxonomic and nomenclatural history since the first of its species were discovered more than a century and a half ago. Some :>f the chief landmarks are as follows:

7

Kattermann:

Eriosyce

Historical landmarks 1829. Cactus curvispinus, the earliest known species now referred to Eriosyce, was named by Bertero. It was described by Colla in 1833. Meanwhile, in 1831, Haworth described Echinocactus subgibbosus, later to be the type of Neoporteria. Various other species, including the type of Eriosyce itself, Echinocactus sandillon Gay (= E. auratus Pfeiffer, described a few months earlier), were named before 1850. 1872. Dr R.A. Philippi (Descr. pi. nuev., 63. 1872) proposed the genus Eriosyce for E. sandillon, separating it from Echinocactus by reason of the floral tube, produced beyond the ovary and spiny at the apex, and the dry fruit Ctubo elongato supra ovarium producto, apice spinas cum petalis gerente; bacca exsucca'). 1922. Apart from Eriosyce sandillon, all known plants of this South American group had been included in Echinocactus Link & Otto until Britton & Rose's major sub­ division of the latter genus in vol. 3 of their monograph The Cactaceae. Britton & Rose recognized Eriosyce as a monotypic genus, using the name E. ceratistes (now considered misapplied) for the only species and referred all but one of the other relevant species to two other genera. Those with funnel-shaped flowers and bristles on the flower-tube were placed in the new genus Neoporteria, with Echinocactus subgibbosus Haworth as the type, and those with campanulate flowers in Malacocarpus Salm-Dyck (nom. illegit., ICBN Art. 64), which is now for the most part included in Parodia Speg. sens, lat.). Echinocactus villosus Monville was listed as an uncertain species, perhaps referable to Matucana. The exact identity of the original E. villosus remains uncertain, along with several of the seven species Britton & Rose recognized in Neoporteria. Relevant species placed by Britton and Rose in Malacocarpus were M. reichei (Schumann) B. & R., M. napinus (Philippi) B. & R., M. catamarcensis (Spegazzini) B. & R., M. islayensis (Forster) B. & R., M. strausianus (Schumann) B. & R., M. tuberisulcatus (Jacobi) B. & R. and M. curvispinus (Bertero ex Colla) B. & R. Their broadly based concept of Malacocarpus included plants from Bolivia. Peru, Brazil, Paraguay and Uruguay, as well as from Argentina and Chile. They distinguished it from Eriosyce by the absence of spines at the top of the fruit. 1929. Alwin Berger, in his book Kakteen, divided Malacocarpus, referring the hollowfruited species toNotocact'us (flowers yellow) and Pyrrhocactus (flowers flame-coloured or reddish yellow), with Echinocactus strausianus Schumann as its type. Berger also included E. catamarcensis Speg., E. tuberisulcatus Jacobi and Cactus curvispinus Bertero ex Colla. This classification distinguished the Argentinian forms now referred to Eriosyce from other Notocacteae in Argentina, like Parodia (Notocactus), whilst confirming the close relationship of plants from both sides of the Andes. 1934. Curt Backeberg proposed the genus Islaya, with Echinocactus islayensis Forster as type species, separating it from Malacocarpus on account of its hollow fruit. 1935. Backeberg (in Backeberg & F. Knuth, Kaktus-ABC, 299, without Latin diagn.) proposed the genus Chilenia for part of Neoporteria B. & R., but overlooked that one

В

Kattermann:

Eriosyce

of the species he included, Echinocactus subgibbosus, was the type of the latter name. After abortive attempts by Backeberg to undo this mistake himself, it was eventually rectified in 1942 by B. Doelz (in Fedde, Repert. Sp. Nov. 51: 60) who published Neochilenia Backeberg ex Doelz for the part of Neoporteria with funnelform, rotate flowers (though it is arguable that the name Nichelia Bullock (published in 1938), should take precedence). Meanwhile, Backeberg (Blatter f. Kakteenforschung 1938(6): [16, 21J) proposed another new genus, Horridocactus Backeberg, based on Cactus horrid us Colla, for Chilean species allied to the Argentinian Pyrrhocactus but with the ovary nearly naked. His proposal drew attention to the differences between plants growing on the two sides of the Andes, which he argued were a barrier between the two groups, but the basis for separating the Chilean plants into Horridocactus and Neochilenia (the amount of bristles etc on the floral tube) now seems inadequate. 1957. Y. Ito {Explanatory diagram of the Austroechinocactinae, 148, 292) proposed the genus Thelocephala for Neoporteria napina and another species with prominently tubercled stems. 1959. F. Ritter (in Cactus (Paris) 14(63): Suppl. [5]. 1959) took up Frit's proposed but undescribed genus Chileorebutia, based on Echinocactus reichei Schumann, for various species with tuberous roots and a stem reminiscent of Rebutia spp. These included Neoporteria napina, the type of Ito's Thelocephala, and so the name Chileorebutia Fric ex Ritter is illegitimate. In the same year Ritter (in Succulenta 1959( 10): 129-131) also rejected Backeberg's division of Pyrrhocactus, broadening the genus to include Horridocactus and Neochilenia. 1962. Backeberg's monumental work. Die Cactaceae, completed publication. He continued to place the plants from the east and west side of the Andes in different generic groups and restricted Pyrrhocactus to species from Argentina. His 'Pacific' genera included Eriosyce, Horridocactus, Islaya, Neochilenia (incl. Chileorebutia and Thelocephala), Neoporteria and Rodentiophila (an invalid name). In the 1962 version of his 'phylogenetic' classification, F. Buxbaum (in Krainz, Die Kakteen, Lfg. 21) accepted Pyrrhocactus, Islaya and Neoporteria (incl. Chilenia, Chileorebutia, Horridocactus, Neochilenia and the misplacedReicheocactus) and placed them in his tribe Notocacteae. At that time, Buxbaum regarded the relationships of Eriosyce and Rodentiophila as uncertain, grouping them next to Eulychnia in tribe Leptocereeae, but by the time the final version of his system appeared in 1974 (in Endler, J. & Buxbaum, F.. Die Pflanzenfamilien der Kakteen, ed. 3), he had realized :hat Eriosyce and Rodentiophila should be placed in the Notocacteae (I.e., page 161, note no. 48). 1966. Donald & Rowley (in Cact. Succ. J. Gr. Brit. 28: 54-58, 74-77, 83) reunited Hurridocactus, Islaya, Neochilenia, Pyrrhocactus and Thelocephala under Neoporteria. Their work was based on a statistical^analysis of floral and other characters. Plants

9

Kattermann;

Eriosyce

growing on the eastern side of the Andes were once again classified with plants growing on the western side. Islaya from Peru was also included, but Eriosyce was excluded, largely, it seems because of the nomenclatural implications: Notocactus... and Eriosyce run so close that they can be separated by combinations of characters, no one of which is absolute. Artificial considerations, like convenience of use and desire to retain old familiar names tip the balance here.' l

1980. In his monograph, Ritter (Kakteen in Sudamerika, vol. 3) retained the narrower generic concepts proposed in his earlier work, recognizing£>/osycc, Islaya, Neoporteria, Pyrrhocactus (including Horridocactus and Neochilenia) and Thelocephala as genera. He also described many new species. His locality data greatly helped the present author to collect the material used in this revision. 1986. For its 'consensus' list of accepted genera (Hunt & Taylor, eds., in Bradleya 4: 65-78), the IOS Working Party adopted the Donald & Rowley circumscription of Neoporteria, noting that Eriosyce is 'An older name than Neoporteria with which it might otherwise be combined'. The IOS list has been adopted for various subsequent treatments, including Adriana Hoffmann's work on the cacti of Chile (Hoffmann 1989). Scope of present study The writer's conclusions are based on more than a decade of studies of about 500 collections or population samples made during many field trips to Chile and Argentina. Observations in the field and greenhouse were documented with photographs of plants, flowers and fruits, herbarium vouchers and many notes and analytical drawings. A detailed study of seed-morphology was carried out using both the light microscope and Scanning Electron Microscope (SEM). Numerous vegetative, floral, fruit and seed characters were considered, and all the recognizable species scored for 58 characters considered to be of taxonomic value. The resulting data-matrix was subjected to a cladistic analysis to test my intuitive conclusions concerning the classification and phylogeny of the group. General conclusions As data accumulated, it became increasingly apparent to me that all the species under discussion should be included in a single genus. The data also suggested that at the species level a much broader approach was necessary than had been adopted of late. Many of the described taxa are local populations representative of much larger species complexes. Number of spines and their length, number of ribs, quantitative covering of the flower tube and ovary by bract-scales and wool etc. are environmentally influenced and are therefore of limited use for classification. Past misunderstandings, incomplete data, ignorance of the rules of nomenclature, and a tendency among authors to describe individual plant populations as separate species, have led to the publication of a large number of unacceptable or doubtful names and consequent taxonomic problems. Many of the names must now be consigned to synonymy or discarded for lack of adequate typification.

10

Kattermann:

Eriosyce

Outline of proposed classification Fieldwork carried out from 1977 to 1986 convinced me that Eriosyce in its narrow original sense is only part of a larger and closely related group which includes plants growing in north-western Argentina, central to northern Chile and southern Peru. Consequently I am proposing to enlarge Eriosyce by merging it with Neoporteria sensu Donald & Rowley. Unfortunately the rules of nomenclature prescribe that the larger group lNeoporteria) must be sunk into the smaller (Eriosyce), because Eriosyce is the older name. In defence of Eriosyce, however, it can be argued that it has been a wellknown name, used with a consistent circumscription, since Britton & Rose's mono­ graph, whereas the circumscription of Neoporteria has varied widely, from few to many species. Included in the expanded concept of Eriosyce are Chileorebutia Ritter, Islaya Backeberg, Horridocactus Backeberg, Neochilenia Backeberg, Neoporteria Britton & Rose, Pyrrhocactus A. Berger and Thelocephala Y. Ito. I am further proposing to divide the expanded genus into two sections, sect. Erio­ syce, and sect. Neoporteria. Sect. Eriosyce is split into subsection Eriosyce, subsect. Islaya and subsect. Pyrrhocactus. Sect. Neoporteria is divided into subsect. Chileosyce
11

Kattermann:

Eriosyce

subsect. Neoporteria, comprising E. chilensis, E. senilis, E. sociabilis, E. subgibbosa and E. villosa, has flowers which have incurved perianth-segments and are adapted for hummingbird pollination. Characters used in classification Previous classifications of Eriosyce and allied genera have largely relied on a relatively few characters, such as flower shape, style and stigma colour, tube armature, and whether the fruit is fleshy or dry (cf. Donald & Rowley 1966). In the course of my work I have identified useful additional characters, notably in the form of the nectary, the junction of style and stigma, the attitude of the stigma-lobes, and the mode of dehiscence of the fruit. The Scanning Electron Microscope has facilitated examination and comparison of the seeds, and several characters of taxonomic importance can now be recognized. On the basis of my fieldwork and observations of documented material in cultivation, I was able to distinguish 33 species. To test my ideas on their classification and inter-relationships, I scored them for numerous characters, eventually reduced to 58. After various refinements, the resulting data-matrix was subjected to a cladistic analysis by Dr Robert Wallace (see Appendix III). The characters, and the characterstates observable for each, are listed in Table 1 (pages 126-128), which is in itself a summary of the morphological diversity present in the genus. Brief notes are given below on the vegetative and floral characters employed in the analysis, followed by a systematic overview of fruit and seed characters in the genus. Vegetative characters (Matrix nos. 1-19) 1. Stem elongation. Most species of Eriosyce are globular or subglobular in habit but the stem may also be elongate or columnar, as in some forms of E. subgibbosa. 2. Stem diameter varies from 1.5—40 cm. 3. Stem firmness. In most species the stem-surface is not easily depressed, but soft stem tissue has been observed in E. aspillagae, E. crispa and E. villosa. 4. Taking the genus as a whole, rib number varies from 7-30 or more but the range in individual groups and species overlaps to such as extent that it is of little use as a 'key' character. In many populations the number of ribs increases with age. 5. Rib shape. In all species the ribs are somewhat notched or depressed between the areoles and in some the ribs are dissolved into elongate or isodiametric tubercles. 6. When the ribs are deeply notched, one can often observe a chin-like protrusion below the areole. 7. The areole itself may be superficial (slightly raised to almost flat) or sunken to a varying degree. 8. Areolar wool is always present, and may be very short or longer, in some species 3 mm or more and floccose. 10. Apex. Depending on the development of areolar wool, this varies from naked to densely woolly. The only spine characters used in the analysis were as follows. Radial spines: 10. Thickness, 11. Attitude and 12. Curvature. Central spines: 13. Presence I absence, 14. Thickness, 15. Attitude and 16. Curvature. Spine number is very variable within taxa and is of little value as a diagnostic character. 17. Roots. The root system in plants of this genus varies from fibrous to turbinate or tuberous. 18. In thick-rooted species, the presence or absence of a neck-like constriction where the root joins the plant-body is diagnostic. 19. The root-epidermis

12

Kattermann:

Eriosyce

is either firm, and not easily damaged, or very soft and easily damaged, as in in E. krausii and E. laui. Floral characters (Matrix nos. 20-37) 20. Flower production may be from old areoles, either lateral or around the periphery of the stem-apex, or from young areoles in a circle at the apex itself. 21. Flowers per areole. Usually one, rarely more than one. Eriosyce flowers are short to long funnelform. The perianth is usually erect to curved outward but curved inward in subsect. Neoporteria (character no. 34). Features of the pericarpel and tube have traditionally been found to be useful in the classification of this group. The pericarpel and tube have bract-scales which are usually more or less woolly in the axils and may also have bristles. 22. Wool length varies from very short (the tube appears ± naked) to long (with the wool extending to the next higher bract-scale, partially or fully obscuring tube or pericarpel epidermis). Very rarely it is absent (E. laui). 23-25. Pericarpel bristles may be present or absent, and vary in number, length, shape and attitude. 29-33. Tube bristles are always present (except in E. laui), and also vary in number, length, shape, thickness and attitude. Useful internal floral characters include 28. Style insertion, i.e. whether the ovary apex is drawn upward into the base of the style, or flat, or depressed (so that style appears pushed downward into the ovary apex). 35. Nectar-chamber shape. I distinguish three types: 'Simple', narrowly tubular < 1-5 mm long), with the walls of the chamber parallel to each other and its base rounded or angular; 'Modified Г, with the upper portion narrower than the base (this may be by thickening of the tube wall, narrowing of the tube just above the pericarpel, or widening at the base of the nectarchamber), and the base of the nectary is ± flattened; and 'Modified II\ where the base of the nectar-chamber is much widened, almost pear-shaped, or sometimes more angular (subsect. Neoporteria only). 36. Stigma-lobe attitude. This varies from spreading or recurved to erect or the lobes may appear clasped together. 37. Stigmabase. In several species there is a clear enlargement of the top of the style ('stepped'), while in others there is a smooth uninterrupted transition from style into stigma-lobes ''not stepped'). Fruit characters (Matrix nos. 38—43) There is considerable variety of fruit structure in the genus, related to mode of seed dispersal. The fruit may be dehiscent or indehiscent, and absciss or remain attached to the plant and several types of dehiscence and abscission are found (38-39). The pericarp (fruit wall) varies in thickness at maturity (40) and how quickly it dries up 141). The fruit may or may not elongate, but becomes enlarged and balloon-like in subsect. Islaya (42). The seeds may be retained in the fruit in various ways, or fall out freely on dehiscence (43). In subsect. Eriosyce the fruit when completely ripe detaches from the areole and is lifted upward exposing loose seed at the base of the fruit (N.P. Taylor, pers. comm., reports that fruits may not always dehisce and expose the seed directly). Fruit of E. rodentiophila from a population north of Taltal did not have a basal opening, but had all the other characters of a subsect. Eriosyce fruit.). The fruit remains fleshy for an

ta

Kattermann:

Eriosyce

extended period after dehiscence. Seed dispersal could be effected by rainfall, but also appears to be assisted by insects and rodents. The fruit of subsect. Pyrrhocactus dehisces by a partial circumscissile slit at the base of the fruit. The fruit wall dries out within a few days of ripening and this drying process partially lifts the fruit, creating a basal opening from which the loose seed falls on to the plant. Seed dispersal appears to be similar to that in subsect. Eriosyce. In subsect. Islaya the fruit dehisces by a basal opening. Abscission of the fruit is incomplete. Fruit detachment depends on external forces, possibly wind or also rodents and reptiles. The fruit expands balloon-like, facilitating removal of the fruit by wind. Seed dispersal appears to take place as the fruit is blown over the ground. Seed is often contained in a sac-like structure inside the top of the fruit. This may be a mechanism to retard seed dispersal until the fruit has travelled far from the mother plant. This sac-like structure with modifications can also be observed in subsect. Chileosyce and in some species of subsect. Horridocactus. In subsect. Chileosyce the fruit dehisces by complete abscission at the base leaving a basal opening. It dries within a short time after reaching maximum elongation. The fruit is densely covered with many porrect bristles facilitating wind removal and seed dispersal. In subsect. Horridocactus and Neoporteria seed dispersal appears to depend on removal of the fruit by animals. Its red colour supports this hypothesis. Dehiscence occurs when the fruit separates from the stem completely leaving a basal pore, except in E. aspillagae, which has a mode of dehiscence similar to that in subsect. Pyrrhocactus. In some instances seed dispersal is restrained by a sac-like structure inside the top of the fruit. Seed characters (Matrix nos. 44-58) The value of seed-characters in the taxonomy of the Cactaceae has long been known, and the diverse form and structure of Eriosyce seeds which is visible to the naked eye suggested their examination with the Scanning Electron Microscope (SEM) would be particularly interesting. A total of 95 population samples was examined, many of which are illustrated in the SEM Plates intercalated in Appendix IV (pages 133-163). Method. After brief rinsing in water, seed was gold-coated in a Pelco Model 3 Sputter Coater and then examined with a JEOL Scanning Microscope, model JSM-T300. After a preliminary study of three seed samples at various magnifications, standard magnifications were established: for a lateral view of the seed x 35; frontal view of the hilum-micropylar region x 75; testa-cell detail from central region of the lateral side of the seed x 750. Some pictures were taken at different magnifications to show specific details. In some seeds the testa-cells near the hilum may be different but this was not utilized in this analysis. Some taxa have seeds with a loose sheathlike cuticular layer. Removal of this was not attempted. To utilize the data obtained, various characters and character states listed in a draft paper by Barthlott & Hunt lined.) were scored. These include surface relief features (44—46); testa-cell size (47); visibility of anticlinal cell-boundaries (48); cellinterstices (49); shape and relief of periclinal walls (50—54); features of hilummicropylar region (HMR; 55-58).

Kattermann:

Erioayce

Subsection Eriosyce has the largest seeds in the genus, the majority being 1.9-2.7 mm. Testa-cell size varies from 0.0025-0.01 mm . Some testa-cells are elongated. At the cell interstices deep pits can be observed. The hilum region differs in the various subsections of Eriosyce. In subsect. Eriosyce the micropyle is not clearly delimited, or if it is, it does not have special cells surrounding or partially surrounding it. The cells of the hilum rim are the same on all sides. The cells end abruptly at the rim. Various hilum shapes can be observed. In subsect. Eriosyce the hilum is ovate to oval or widely keyhole-shaped. Testa-cell shape and surface features and the anticlinal borders vary from population to population. Deep pits at cell interstices are present in most populations. The anticlinal boundary is often bridged by transverse striations. The testa-cell surface can be highly sculptured with fine striations or the surface may be smooth. Sometimes a tertiary structure of folded striations covers or partially covers the testa-cell surface. The seeds of subsect. Pyrrhocactus are smaller, mostly 1.5—2.1 mm. Testa-cell size varies from 0.0036-0.0081 mm . All testa-cells are isodiametric. Pits at the cell interstices are lacking. The hilum/micropylar region is similar to that in subsect. Eriosyce. The micropyle is not clearly delimited or else it does not have special cells surrounding or partially surrounding it. The testa-cells of the hilum rim are the same on all sides, ending abruptly at the rim. The shape of the hilum varies from oval to ovate or wide keyhole-shaped. In general the cell surfaces are smooth, but partly hidden under tertiary striations. In some examples, the surface of the seed is provided with a thick cuticular sheathlike covering, making it difficult to evaluate testa-cell surface features. The anticlinal testa-cell boundaries are often bridged by parallel or branched, transverse striations. The seeds of subsect. Islaya studied are not of documented origin, being from various commercial seed sources. Flower characters suggest the possibility that more than one species exists and seeds observed may support this. However, due to insufficient field material and experience I will include all within a single species under the name E. islayensis. Most of the seeds are 1.3-1.4 mm, with testa-cell size in the range 0.0018-0.0049 mm . Some testa-cells are slightly elongated and in some seeds deep pits at the cell interstices exist. The hilum-micropylar region differs from the remainder of Eriosyce in having testa-cells at the hilum rim equal in size to those on the lateral surface of the seed. The shape of the hilum is wide keyhole-shaped, but narrower than in subsect. Pyrrhocactus and subsect. Eriosyce. The testa-cells near the micropyle are not modified as in sect. Neoporteria. The anticlinal cell boundaries are ± straight and without striations bridging the cell borders. In some, the periclinal surfaces are smooth, while others are finely striated. Tertiary appendages are absent in some and present in others. The seeds of sect. Neoporteria subsect. Horridocactus are 0.9-2.0 mm. Testa-cell size varies from 0.0025—0.01 mm . All cells are isodiametric and pits at cell interstices are lacking. The hilum region is modified at the micropyle, which is in a narrow groove surrounded or partially surrounded by inward growing cells different in shape and structure from the cells surrounding the rest of the hilum. Sometimes these cells completely surround the micropyle, separating the micropyle from the hilum. This surrounding tissue and the degree of isolation of the micropyle can be variable in seeds 2

2

2

2

15

Kattermann:

Eriosyce

of the same population. In some seeds the groove exists and is open, while in others it is completely surrounded. Several cell types can be observed in subsect. Horridocactus. Most have highly developed tertiary structures. The anticlinal cell boundaries are not always visible. When visible the boundary is bridged by parallel striations. In this analysis it is assumed that this holds true for all the species of this subsection. The seeds of sect. Neoporteria subsect. Chileosyce are the smallest in the genus. Most of the seeds are 0.9-1.1 mm. Testa-cell size varies from 0.0015-0.0049 mm . Pits at cell interstices are lacking. The micropyle is not clearly delimited or else it does not have special cells surrounding or partially surrounding it. The cells of the hilum rim are the same on all sides, ending abruptly at the rim. The shape of the hilum is oval to wide keyhole-shaped. The cell shape, surface features and anticlinal borders vary from population to population. The anticlinal boundary is sometimes bridged by parallel striations. The cell surface can be highly sculptured with fine striations or smooth. The seeds of subsect. Neoporteria vary little from those of subsect. Horridocactus. The length of the seed varies from 1.0-1.6 mm and testa-cell size from 0.0016-0.0081 mm*. The cells are isodiametric and pits at the interstices are lacking. Reviewing the seed data, similarities between the various sections and subsections of Eriosyce can be observed. Within the sections, differences may correlate with a diversity of habitats for a large species group, or be characteristic of the species group itself. The diverse cell surface characters of E. aurata include populations growing from almost sea level to above 3000 m elevation and a north-south distribution of almost 500 km. In subsect. Pyrrhocactus several of the species have similar testa-cell surfaces. Only E. umadeave has a different cell surface. Subsect. Islaya has in some of the seed samples deep pits at cell interstices, just as in subsect. Eriosyce. Testa-cell characters in subsect. Chileosyce suggest affinity with several of the other subsections in Eriosyce. The northern species group around E. krausii (including a plant of uncertain identity collected by Walter Jung near Totorab shows affinity to E. islayensis. E. laui from Tocopilla may be intermediate. E. tenehrica in turn suggests in its testa-cell characters a possible affinity to E. kunzei var. transitensis. In subsect. Neoporteria and Horrido­ cactus the seeds are similar. They show some relationship to the other subsections. The shape of the hilum suggests that if evolution of the Cactaceae followed a north-south direction in South America, the keyhole hilum shape is a state derived from the ovate hilum. The change from the ovate/oval shape to the keyhole shape occurs in subsect. Eriosyce, subsect. Pyrrhocactus and section Neoporteria. PI ate-tectonic studies suggest that the coast of Chile has been and is currently uplifting. It has been suggested that this uplifting may be as much as 1 cm per year. This would suggest that the tops of mountains 1000 m high may have been at sea level as little as 100,000 years ago. This scenario leads to the idea that most of the cacti on the western side of the Andes evolved within the last 10,000 to 500,000 years. It further suggests that subsect. Eriosyce and subsect. Pyrrhocactus may have existed at similar elevations, without the barrier of the present high Andes, which may explain their close similarities in seed characters. 2

16

Kattermann:

Erioayce

Reproductive system Reproduction within the genus Eriosyce is mostly by seed. Branching or offsetting has only been observed in E. aspillagae, a form of E. curvispina var. mutabilis from Pichidangui, a single population of E. esmeraldana, a population of E. krausii and a population of E. villosa. The various species appear for the most part self-incompatible and require a second clone for successful seed production. The open flowers, except for some plants in subsect. Neoporteria, have an appearance suggestive of insect pollination, and visits by Hymenoptera have been observed in several instances in the field. In subsect. Neoporteria the flowers are adapted for hummingbird pollination. Field observations suggest that there are large groups of populations which share the same genetic pool through regular gene-exchange. Transitional populations between species previously recognized were not infrequently found and these often made it difficult to delimit species. Hybrids between distinct taxa were not observed. Ritter (1980) suggests that hybrids exist, but I have been unable to confirm this. Allied genera Following Buxbaum (1962, 1974), there is general agreement that the South American cacti formerly included in Echinocactus are not closely related to their North American counterparts and should be placed in a separate tribe, Notocacteae F. Buxb. ('southern cacti'). Buxbaum further divided this tribe into four subtribes: Corryocactinae F. Buxb., a 'basal' group for genera linking the tribe with its cereoid ancestors, and three groups of predominantly 'globular' genera: Neoporteriinae F. Buxb. (mostly endemic west of the Andes), Notocactinae F. Buxb. (floral armature well-developed; genera endemic east of the Andes), and Gymnocalyciinae F. Buxb. (flowers naked except for scales; genera endemic east of the Andes). Members of this last subtribe are referred by the IOS Working Party (Hunt & Taylor (eds) 1986,1990) to tribe Trichocereeae F. Buxb., except for Melocactus Link & Otto, which is placed in tribe Cereeae Salm-Dyck and Neower­ dermannia Fric, which is placed in Neoporteriinae. The principal genera of the other 'eastern' group, subtribe Notocactinae, are Parodia Speg. and Notocactus (Schumann) Fric (now united by the IOS Working Party), which differ from Eriosyce (as redefined here) in their seeds, which have a large hilum and are frequently strophiolate. The genera regarded as most closely allied to Eriosyce are Austrocactus B. & R., Copiapoa B. & R., Corryocactus B. & R., Eulychnia Philippi and Neowerdermannia Fric. Of these, Eulychnia and Corryocactus (now thought to be closely related to one another), are very different in habit from Eriosyce, being shrubby or tree-like, often freely branching. In the present study, the type species of Corryocactus, C. brevistylus I Schumann ex Vaupel) B. & R., was chosen as the 'outgroup' for the cladistic analysis of Eriosyce (Appendix III). Its description is given below, followed by notes on Austrocactus, Neowerdermannia and Copiapoa. Note that measurements here and elsewhere in this paper refer to length, unless otherwise indicated.

17

Kattermann:

Eriosyce

Corryocactus brevistylus (Schumann ex Vaupel) Britton & Rose, The Cact. 2: 66 (1920); Cereus brevistylus Schumann ex Vaupel, Bot. Jahrb. 50: Beibl. I l l , 17 (1913). Stem columnar, freely branching from the base, erect, forming clumps, or often tree-like; ribs 6-7, prominent; areoles 3 cm apart; spines c. 15, unequal in length 1-31-24) cm; flowers yellow, 9 cm, broadly funnelform, constricted above the pericarpel, pericarpel and tube with tufts of wool and bristles from the axils of bract-scales; stigmas white, lobes 18-29, 10-15 mm; nectar-chamber simple, elongate, tubular; filaments inserted over И or more of tube; fruit indehiscent, ± globose, filled with funicular pulp, perianth remnants deciduous; seed medium in size, c. 1.7 mm; hilum oval, large, sunken; testa tuberculate, sometimes ribbed. DISTRIBUTION. S Peru and N Chile. Austrocactus B. & R. (5-6 spp.) The stem is ± cereoid and commonly soft-fleshy. Most form adventitious roots from the ± procumbent stems, which sometimes thicken to form tuberous roots. Adventitious roots have not been recorded in Eriosyce. In Austrocactus the stamens are disposed in two series. One set is arranged s hugging the tube wall, the other set upright around the style. This can also be seen in the genus Copiapoa but has not been observed in Eriosyce. Pollen data published by Leuenbergcr (1976) euggest the separation of Austrocactus from Eriosyce. The pollen of Austrocactus is 12-colpate, whereas it is mostly 3-colpate or rarely 6-colpate in Eriosyce. The fruit morphology is not entirely understood. One fruit of A. spiniflorus observed in habitat did not display the method of dehiscence or looseness of the seed within the fruit. Roger Ferryman (pers. comm.) reports that the fruit upon drying dehisces by a basal pore, somewhat as in the genus Eriosyce. I could not confirm this. A. Hoffmann (1989) notes for A. hibernus that the fruit is with mucilaginous flesh CFruto en forma de tonel, verde у blando, con la carne muscilaginosa у perfumada'). A similar reference is given for A. spiniflorus. The seed differs from Eriosyce in the shape of the hilum-micropylar region and the location of the hilum in relation to the longitudinal axis.

FIGURE I

1 2

Austrocactus spiniPorus lErditio spimflora).

3

Acanthocatycium brevispinum. C a f a y a t e ,

4

Copiapoa tenuissima. El

5

Neowerdermannia

Lagunillas. Chile. F K - 3 0 6 .

C

DISTRIBUTION. Both sides of the Andes, from c. 34 S in Chile to 40°S in Patagonia, Argentina. In the northern half of their range they parallel the distribution of Eriosyce. The most northern population I have studied is of A. spiniflorus. Additional material was received of A. hibernus collected by Walter Jung in Chile, and of Austrocactus populations from Mendoza and from Neuquen collected by R. Kiesling and by B. Leuenherger & S.C. Arroyo.

18

Corryocactus brevistylus. Zapnhuira, Chile.

Argentina, F K - 6 6 9 .

Cobre. C h i l e . F K - 7 7 9 .

chitenKis.

Belen, C h i l e ,

F K - 3 5 0 : fruit on right is from N.

vorwerkii.

Kattermann:

Eriosyce

19

Kattermann:

Eriosyce

Neowerdermannia Fric (2 spp.) This differs from Eriosyce in the naked pericarpel and tube without bract-scales, and by the method of fruit dehiscence. Dehiscence is by a partial circumscissile splitting near the top of the fruit with the partial fruit cap tilting outward at about 160° in N. vorwerkii, and by several longitudinal slits in the wall of the fruit in N. chilensis. The seed of Neowerdermannia differs in having a much larger hilum and a ± strophiolar protrusion of hilar tissue. D I S T R I B U T I O N . North and west of the range of Eriosyce. N. chilensis ranges from 18-19"'S, always above 3000 m and on the western side of the Andes in Chile. N. vorwerkii is reported from Bolivia and N Argentina. Its range does not overlap with that of Eriosyce. Copiapoa B. & R. 117-26 spp.) Copiapoa differs from Eriosyce in its ± naked pericarpel and tube EG solans has wool in the axils of the bract-scales), in its stamens disposed in two series (as in Austrocactus and Corryocactus). and in its mode of fruit dehiscence, which is similar to that of Neowerdermannia vorwerkii. The evolutionary history of Copiapoa is not yet entirely clear. It has been suggested that C. solans (genus Pilocopiapoa Ritter) might represent a link to Eriosyce. I cannot confirm this. See also the description of Eriosyce laui (Appendix II). D I S T R I B U T I O N . Chile, from 22-31 S. Populations oCCopiapim are often found near sea level, and they rarely grow more than a few kilometres from the coast. ;

Donald (in Hunt & Taylor 1990: 99) argued that the genus Acanthocalyciurn Racket), could be subsumed into Pyrrhocactus or Neoporteria, citing Pyrrhocactus andreaeanus (A. andreaeanum (Backeb.) Donald) as a 'good intermediate'. I cannot find evidence to support this. The flower of this species has the filaments evenly inserted over the lower one-third of the tube, their attitude ± parallel with the style. The filaments of Acanthocalycium sens. str. are appressed to the tube wall with their tips curved inward. Also its nectary is partly closed by a woolly plug, whereas Eriosyce andreaeana has an open nectary. The fruit of E. andreaeana dehisces by a basal pore, Acantho­ calycium by splitting of the side wall (see fig. 1.3). Uses and economic importance There is no evidence that members of the genus Eriosyce have been used by local peoples except for horticultural purposes. In Concepcion, near the southern limit of sect. Neoporteria, Eriosyce subgibbosa was and still is collected for sale at the local market near Christmas time, when they are in flower. This collecting has greatly reduced the populations of E. subgibbosa around Concepcion. There are a few local hobbyists who grow some of the species in their gardens. Representatives of the genus have been in horticultural production outside Chile for many years. Threats to the survival of Eriosyce species Most of the species of Eriosyce are threatened by the spread of human habitation. The populations of E. curvispina var. tuberisulcata near Valparaiso, the type locality, have only a few plants. These populations have as their neighbours large apartment complexes. The locality of E. napina at Huasco, the type locality, is now used as a dump. E. aspdlagae is probably extinct because of collecting and pulpwood production. In the drier areas in the northern part of the range of the genus grazing by goats has

20

Kattermann:

Eriosyce

damaged many of the habitats. Yet further north, the extremely dry climate over the past 25-50 years has destroyed many populations. Only dead plants could be observed. Commercial collecting has also been a problem. Fortunately the government of Chile is trying to prevent this kind of trade. It is hoped and indeed possible that some of the over-collected populations may recover. Cultivation In view of the wide distribution of Eriosyce, several climatic conditions must be considered. All Eriosyce require a well aerated, open growing medium. Extended periods of waterlogged conditions are a sure path to disaster. Plants of subsect. Eriosyce growing at high altitude may often be covered by snow. In dry condition they should be extremely frost hardy and they require a long winter rest period, with at most only very light misting. In the spring they have a short growing period during which the flowers develop. After flowering, watering should be done very sparingly. Heavy misting is probably the best method. They require lots of sun. The more northern populations of subsect. Eriosyce, specifically E. rodentiophila, grow in areas without normal rainfall. Heavy misting should be the only method of watering, ensuring that all moisture in the soil has evaporated before misting again. Plants of subsect. Eriosyce are difficult to flower and have to be well over 15 years old before reaching reproductive maturity. Plants of subsect. Pyrrhocactus are also for the most part growing in the snow belt. E. umadeave has regular snow cover and growing at 3000 m altitude is frost-hardy. Plants of E. umadeave must reach a large size before flowering, as in subsect. Eriosyce. Other species of subsect. Pyrrhocactus are easy to flower. They require a long resting period during the winter months. With good porous soil, watering is not a problem. Subsect. Islaya grows mostly on coastal hills, often in sand. They are the least coldhardy of Eriosyce, since they experience more tropical temperatures in their habitat. They grow in extremely dry conditions. Rainfall may occur only at 10-30 year intervals and they depend entirely on the coastal fogs for their water requirements. The coastal fogs, called camanchaca in Chile and garua in Peru, are the only source of moisture for the coastal vegetation of northern Chile and central-southern Peru. The fog is very dense and condenses on the spines and areolar trichomes. The plants may be totally wet, yet little water reaches the ground. The plants may be bathed in this fog for many hours each day, especially in the morning. Fog may be present for weeks at a time, interspersed with weeks or months without any fog. Associated vegetation often comprises small bromeliads and many types of lichens. In cultivation it is best to use a very porous growing medium, and use only misting as the method of watering. E. islayensis flowers freely under these conditions. However, it resents transplanting, which should only be attempted when absolutely necessary. A very long dry period with only the slightest misting is recommended after transplantation. Subsect. Horridocactus inhabits the widest range of climatic conditions. The species E. curvispina can be an alpine plant with several months of snow cover, or may grow u only a few metres above sea level. Most forms of E. curvispina are easy to grow. The <mly exception is var. tuberisulcata which should be grown in even more porous media and prefers less water. Heavy misting is always safer. E. garaventae grows in cracks

21

Kattermann:

Eriosyce

of rock outcrops and its roots, which penetrate deep into the rock fissures, may have only a few grammes of decomposed organic material available to them. Less watering is indicated by this type of habitat. E. heinrichiana, E. napina, E. taltalensis, E. crispa and similar species grow in dry areas where rainfall may not occur for several years at a time. They depend mostly on the moisture derived from coastal fogs. In the dry desert regions plants derive additional moisture from condensation. Temperature fluctuations between day and night are often great enough to reach the dew point at night. In the daytime the temperature may reach 35°C at a relative humidity of 3-5%. Night temperatures may drop to 5-10°C with a corresponding increase in relative humidity, often reaching the dew point. This high relative humidity at night also accommodates the best time for gas exchange of these Crassulacean Acid Metabolism (CAM) plants. Therefore, I always mist my plants in the evening. Subsect. Neoporteria has for the most part a coastal distribution, where, at its southern limit it grows in coastal areas with regular rainfall, often competing with dense vegetation. Some populations grow on rock outcrops close to the coast, such that sometimes mist from the high surf of the Pacific bathes the plants in moisture. Plants from these areas like lots of misting between normal applications of water. The more northern populations, especially those which have developed tuberous root systems, must be grown in a porous medium and watering replaced by heavy misting. Members of this subsection are probably not frost hardy except for possibly some of the populations of E. senilis which grow at up to 1900 m elevation. Subsect. Neoporteria in habitat flowers in the spring, never in autumn, yet in cultivation in the northern hemisphere flower-buds are developed in autumn (in New Jersey, U.S.A., around October). Some plants will actually flower at this time. Proper cultivation for the preservation of these plants through the winter months requires that dormancy is induced by not watering and by cool temperatures (З^С in my greenhouse). Under these conditions plants of subsect. Neoporteria produce a second crop of flowers, usually even more abundant than the autumn crop. Subsect. Chileosyce is probably the most difficult to grow. Good viable seed germinates extremely well. When given abundant moisture the first year from seed is easy. At the first transplanting the smallest damage to the root system spells certain death and this is even more the case with mature plants. Extreme care in trans­ planting is necessary. The plants grow in a particularly hostile environment. A very porous medium is advisable, with watering mostly by light misting. Extended waterlogged conditions mean loss of the plant. Propagation of all species of Eriosyce is best from seed, but E. aspillagae, forms of E. curvispina (from Pichidangui) and some populations of E. krausii and E. esmeraldana offset freely from the main stem. Germination differs between some of the subsections. Subsect. Eriosyce has a reputation for difficulty. Sometimes freshly collected seed germinates 50—60% at the first attempt, but at other times only IO-20'Я and sometimes no germination at all. Some seed samples germinated after extended freeze treatment by leaving the seed pan outside through the winter. Subsect. Neoporteria requires the longest time to germinate, especially plants from the more southern populations. Germination temperature appears, for all species of Eriosyce, best in the range of 20—25°C, with continuous moisture present.

22

Kattermann:

Eriosyce

Selected bibliography BACKEBERG, C. 11959-62). Die Cactaceac, vols 1-6. Gustav Fischer Vorlag, Jena. BARTHLOTT, W. (1984). Microstructural Features of Seed Surfaces. In Heywood, V.H., & Moore, D.M. (eds). Current Concepts in Plant Taxonomy. Systematics Association Special V o l . 25: 95-105. Academic Press, London. & V O I T , G. (1979). Mikromorphologie d e r Samenschalen u n d Taxonomic d e r Cactaceae: Ein raster-elektronmikroskopischer Uberblick. In PI. Syst. Evol. 132: 205-229. B E R G E R , A. (1929). Kakteen. E. Ulmer, Stuttgart. B O W M A N , I. (1924). Desert trails of Atacama. Amer. Geol. Soc. special publ. no. 5. New York. B R I T T O N N X . & R O S E , J . N . (1922). The Cactaceae, vol. 3. Carnegie Institute, Washington, D.C. B R O W N JK, G.W. (ed.) (1968). Desert Biology 1. Academic Press, N e w York, I ^ n d o n . BURNING, A. (1971). Eine Kakteen Reise durch Chile u n d Peru mit Friedrich Ritter. In Kakt. a n d . Sukk. 22(10): 191-195. D O N A L D , J. & R O W L E Y , G.D. (1966). Reunion of the genus Neoporteria. In Cact. Succ. J . Gr. Brit. 28: 54-58. E N D L E R , J . & B U X B A U M , F. (1974). Die Pflanzenfamilien d e r Kakteen, ed. 3. Albrecht Philler Vertag, Minden. G A Y , C. (1848). Cactaceae. In Flora Chilena 3: 8-31. H U N T , D. & T A Y I . O R , N . ( e d s ) (1986). The genera o f t h e Cactaceae: towards a n e w consensus. In Bradleya 4: 65-78. & (1990). The genera of Cactaceae: progress towards consensus. In Bradleya 8: 85-107. HUTCHISON, P.C. (1953). Studies of South American Cactaceae. In Cact. Succ. J . (US) 25: 34-37. •lOHOW, F. (1948). Flora de l a s plantas vasculares de Zapallar. In R e v . Chil. Hist. N a t . 49: 193. J U N G , W. (1978). Contribucidn a l o s Cactaceas Chilenas. In I n v . Agric. Chile vol. 4. K A T T E R M A N N , F. (1980). Neoporteria nigrihorrida, a review. In Cact. Succ. J . (US) 52: 211-214. (1982). Pyrrhocactus (1983). Pyrrhocactus

Garaventai. deherdtiana.

Ibid. 54: 109-111. Ibid. 55: 121-123.

R. (1984). Recopilacion, en edicion facsimilar, de todos l o s trabajos о referencias sobre cactaceas publicadas por el Dr. Carlos Spegazzini. Librosur Ediciones, Buenos Aires. L E M B C K E , H . & W E I S S E R , P. (1979 ]. T h e distribution o f t h e genera of t h e Chilean Cactaceae. In Aloe 17(1): 9-26. L E U E N B E R G E R , B . (1976). Die Pollenmorphologie d e r Cactaceae u n d ihre Bedeutung f u r d i e Systematik. Diss. B o t . 31, J . Cramer, Vaduz. L O O S E R , G. (1929). Catalogus cactacearum chilensium. In R e v . Chil. Hist. N a t . 33: 583-614. PHILIPPI, R.A. (I860). Florula Atacamensis. In: Viaje al desierto de Atacama hecho p o r orden d e l Gobiemo de Chile en el verano 1853-54: [append.) 23. Eduard Anton, Halle. (1891). Verzeichnis der v o n Friedrich Philippi a u f der Hochebene der Provinzen Antofagasta u n d Tarapaca gesammelten Pflanzen. [p. 27.1 Aufgestellt von Dr R.A. Philippi. F.A. Brockhaus, Leipzig. R U N D E L , P. (1976). Succulents in t h e coastal f o g zone of northern Chile. In Cact. Succ. J. (US) 48: 269-272. R I T T E R , F. (1980). Kakteen in Sudamerika, vol. 3. Selbstverlag, Spangenberg. W A G E N K N E C H T , R . (1955). Algunas Cactaceas Chilenas. In Rev. Univ. Catolica de Chile 40: 97-107. W E I S S E R , P. & S C H R E I E R , K. (1976). Die Kakteenwelt Mittel-Chiles u n d ihre Begleit Vegetation. In Kakt. a n d . Sukk. 27: 224-228. WHITE).EY, D.W. (1978). Echinocactus occultus Philippi - The Hidden Hedgehog Cactus'. Some thoughts as to its true identity. In Ashingtonia 3: 62-71. KlESLING,

23

Kattermann:

Eriosyce

MAP 1. The genus Eriosyce: General distribution and species-density.

Kattermann:

Eriosyce

SYSTEMATIC TREATMENT Eriosyce Philippi, Descr. pi. nuev., 63 (1872). Type: E. sandillon (Gay) Philippi (= E. aurata (Pfeiffer) Backeberg). (For synonymy see page 131). Stem subglobular, globular or sometimes elongated, variable in size, 1-100 x 2-50 cm; roots fibrous or including a turbinate tap root, sometimes connected to the above-ground part by a thin neck-like extension of the stem; ribs 7-30 or more, 20—30 mm high, depressed or notched between the areoles or tuberculate; areoles woolly, sometimes with long floccose wool covering stem apex when young. Spines numerous to few, stiff acicular to thin tortuous or bristle-like, 2-50 mm. Flowers arising from nascent or older areoles, rarely > 1 per areole, funnelform to almost tubular, 22-60 mm; tube and pericarpel with numerous scales subtending dense tufts of wool, sometimes with only very short wool or wool absent (pericarpel naked in E. laui), bristlespines present to absent, sometimes only near top of tube or extending to pericarpel, thin and flexible or thick and spine-like, 1-12, appressed or porrect; nectar-chamber ± tubular, often widened at the base; peri a nth-segments pale yellow to deep carmine; stamens numerous, inserted over lower two-thirds of tube; style 10—40 x 1-3 mm; stigma-lobes numerous, diverse in length, thickness and attitude, pale cream to pale carmine red. Fruit a hollow berry, seeds loose within or sometimes some held in a sac-like structure inside the apex of fruit (this structure retains some of the seed and appears to serve as a retardant for seed dispersal), floral remnants persistent, fruit dehiscent by a variously derived basal opening, indehiscent in E. bulbocalyx. Seed 0.7-3.0 mm; hilum ± ovate to keyhole-shaped; micropyle not always obvious, or ± enclosed by special hilum tissue. DISTRIBUTION (MAP 1): Eastern and western slopes of the Andes, from sea level to 3000 m, Argentina, Chile and Peru, 14—37 S. The distribution is almost continuous along the western coastal areas from southern Peru to southern Chile, while at higher elevations populations are more disjunct. The subsections Eriosyce and Pyrrhocactus are separated from each other by the high Andes which reach or exceed 4000 m. N

ARTIFICIAL K E Y T O SECTIONS A N D SUBSECTIONS

1. Floral tube with thick spine-like bristles near apex . . . sect. Eriosyce subsect. Eriosyce 1. Floral tube with thin flexible bristles, often distributed over the pericarpel and tube . . . 2 2. Fruit dehiscent by partial circumscissile slit at base, greenish (red, indehiscent, in E . bulbocalyx) sect. Eriosyce subsect. Pyrrhocactus 2. Fruit dehiscent by complete circumscissile slit at base, or reddish 3 3.

Fruit elongating, >3 cm diam. and or enlarging balloon-like, thin-walled sect. Eriosyce subsect. Islaya 3. Fruit elongating, 1-3 cm diam., not enlarging and balloon-like 4 4.

Perianth-segments upright or directed inward . sect. Neoporteria subsect. Neoporteria

4. Perianth-segments strongly out-curved 5 5. Flower-tube and pericarpel bristles variable, but never porrect sect. Neoporteria subsect. Horridocactus 5. Flower-tube and pericarpel bristles porrect or completely absent sect. Neoporteria subsect. Chileosyce

25

Kattermann:

Eriosyce

KEY TO SPECIES

1. 1. 2.

2.

3.

3. 4. 4. 5. 5. 6.

6. 7. 7. 8. 8. 9.

9. 10. 10. 11. 11. 12. 12. 13. 13. 14. 14.

20

Bristles on flower-tube stiff, spinelike, erect, often constricting the perianth-segments 2 Bristles on flower-tube not stiff and spinelike 3 Stigma-lobes erect or clasped together; stem 12-50 cm diam., apex naked or covered by short wool; ribs 24—40 or more; fruit greenish, covered or partially covered with dense wool; bract-scales large to small; seed 1.9-3.0 mm, hilum region ovate to keyhole-shaped, testacells not arranged in rows or ribs 1. aurata Stigma-lobes spreading; stem 12-32 cm diam., apex covered by wool; ribs 18-32 or more; fruit reddish, with slight wool; bract-scales small; seed 1.8-1.9 mm, hilum oval, testa-cells arranged in distinct rows of single cells 2. rodentiophila Fruit indehiscent; stem 20 x 8-12 cm; ribs 13; flowers 2.5 x 2 cm, perianth-segments narrow lanceolate, 2 mm wide, pale yellow to pale pink; seeds with a large hilum, hilum rim curved outward 3. bulbocalyx Fruit dehiscent; stems, ribs, flowers etc. various; seeds never with outward curving hilum rim 4 Fruit greenish to red. not elongating, dehiscent by a partial circumscissile slit at base, the unequal drying of the fruit wall forming a basal opening 5 Fruit red, elongating, dehiscent by a full circumscissile slit around base 8 Flowers red, narrow funnelform: stem 5-7 cm diam.; ribs 12-14 6. andreaeana Flowers yellow; stem 4-20 cm diam.; ribs 7-21 6 Flowers narrow funnelform to campanulate; stem subglobular to globular, 4-9 cm diam., pearl-grey; ribs 7-11, broad obtuse; perianth-segments 2-3 mm wide, narrow lanceolate 7. villicumensis Flowers wide funnelform; stem, ribs etc. various 7 Stem to 40 x 10-20 cm; ribs 18-21; spines greyish white; perianth-segments 3—4 mm wide, narrow lanceolate 5. umadeave Stem to 20 x 8-15 cm; ribs 12-14; spines yellowish to dark grey; perianth-segments 5-6 mm wide, broadly lanceolate 4. strauaiana Fruit elongating 3-8x diam., sometimes halloon-like 9 Fruit elongating 1—4x diam., never balloon-like 10 Flower-tube and pericarpel covered in very dense wool; bristles always present on tube and sometimes also on pericarpel; stem 5-20 cm diam., apex covered in dense wool; roots fibrous 8. islayensis Flower-tube and pericarpel naked; stem 1.5-2 cm diam., apex not as above; roots large, tuberous 28. laui Perianth-segments strongly curving outward, colour various 11 Peri a nth-segments erect to curving inward, fuchsia red 31 Nectary simple, tubular, not widened at base 12 Nectary modified, widened at base 24 Wool in pericarpel and tube bract-scale axils very short, 1-3 mm, bristles borne only on upper part of tube, 5-10 mm, white, appressed, rarely with >1 per scale axil 13 Wool in pericarpel and tube bract-scale axils 5-15 mm, accompanied by bristles throughout, these 10—20 mm, white, not tightly appressed, 2 or more per scale axil 18 Flower-tube 15-25 mm, wide funnelform, medium to thick-walled; perianth-segments 15-30 mm, variable in width; spines medium thickness, curved 14 Flower-tube 10-15 mm, campanulate, thick-walled; perianth-segments 1.25-1.75 mm wide; spines thickly acicular, strongly curved 17. marksiana Fruit globular or sometimes elongating to 1.5x diam 16 Fruit elongated, 1.5-2.5x diam 17

Kattermann:

Eriosyce

15. Base of stigma-lobes sharply expanded outward at top of style 16 15. Base of stigma-lobes not as above 18 16. Stem to 30 x 14-18 cm; ribs 16-20; seed 1.2-1.7 mm, hilum with micropyle sunken 18. engleri 16. Stem to 20 x 8-20 cm; ribs 14—24; seed 0.9-1.62 mm, micropyle on same plane as or slightly raised above hilum rim 10. curvispina 17. Stem 101-20) x 13-14 cm; spines yellowish; stigma-iobes erect to spreading, 4 mm, creamyellow; seed 1.3 mm, testa tuberculate, sometimes ribbed 11. kunzei 17. Stem 8-13 x 5-15 cm; spines greyish; stigma-lobes clasped together, 4-6 mm, yellowish to orange; seeds c. 1.07 mm, testa ribbed 12. limariensis 18. Ribs not dissolved into tubercles, but often deeply notched 19 18. Ribs dissolved into tubercles 21 19. Spines black to dark greyish, hairlike and tortuous to stiff and acicular; stem subglobular to globular, 5-10 cm diam., blackish grey; ribs 13-16; flower reddish, tube narrow funnelform; stigma-lobes spreading, pale red 14. crispa 19. Spines straw-yellow to white, thin to medium acicular, never tortuous 20 20. Roots fibrous or with only a small taproot; stem 6-12 cm diam., yellowish green, unbranchcd; ribs 11-16, narrow; spines straw-yellow, stiffly acicular; flower greenish yellow, tube narrow funnelform; stigma-lobes clasped together or erect, pale cream 15. garaventae 20. Roots large-tuberous; stem 8-15 cm diam., bright green, freely branching from lower areoles; ribs 5-81-14), broadly obtuse; spines whitish, thin flexible; flower whitish to pale yellow, tube wide funnelform; stigma-lobes pale yellow, spreading 16. aspdlagae 21. Stem 4-8 cm diam., subglobular to globular, often dark brown; flowers trumpet-shaped; pericarpel much longer than wide; stigma-lobes clasped together, 5-6 mm, orange to yellow 19. heinrichiana 21. Stem 2—4 cm diam., grey to reddish brown; flowers funnelform; pericarpel not much longer than wide; stigma-lobes various 22 22. Bristles on pericarpel and tube ± appressed; stem grey, tubercles with short obtuse chin below areole, 4-5 mm diam.; spines 5—8, tightly appressed, black 20. napina 22. Bristles on pericarpel and tube porrect; stem, tubercles and spines various 23 23. Floral bristles red brown; stem often elongating, tubercles 2-3 mm diam.; spines honeycoloured, centrals sometimes present 25. aerocarpa 23. Floral bristles dark brown; stem never elongating, tubercles 4-5 mm diam.; spines black, centrals lacking 27. tenehrica 24. Nectary widened at its ± rounded base 25 24. Nectary widened at its ± flat angular base 26 25. Stem to 100 x 6-12 cm; spines thinly acicular, ± radiating;flowerswide-funnelform, fuchsia red, often with centre white; stigma-lobes spreading 29. chilensis 25. Stem little elongated, 5-15 cm diam.; spines thinly to thickly acicular, curved upward; flowers narrow-funnelform. perianth-segments white with reddish central stripe; stigmalobes clasped together 13. confinis 26. Bristles on pericarpel and tube appressed 27 26. Bristles on pericarpel and tube porrect 30 27. Ribs dissolved into tubercles 2—4 mm diam., with short, obtuse chin below areole; stem 3-5 cm diam.; taproot large; spines rarely to 8 mm; spines horn-coloured, radiating to appressed 21. odieri 27. Ribs dissolved into tubercles with long, acute chin below areole; other characters not in the above combination 28

27

Kattermann: 28. 28. 29. 29. 30. 30. 31. 31. 32.

32.

33.

33.

Eriosyce

Spines often absent, when present to 7, 3-11 mm; stem 3-8 cm diam., subglobular; flowers white 23. occulta Spines always present, to 24, 1-40 mm; flowers various 29 Stem 3-14 cm diam., tubercles with large acute chin below; spines 10-30 mm, centrals 3-6 22. taltalensis Stem 2-5 cm diam., tubercles with short, more obtuse chin; spines 5-15 mm, centrals 0-1 (-2) 9. recondita Bristles on flower-tube dark brown, porrect; flowers white to pink; stigma-lobes clasped together 24. esmeraldana Bristles pale yellowish brown; flowers yellow; stigma-lobes spreading 26. krausii Spines never hair-like or contorted 32 Spines, or some of them hair-like or contorted 33 Stem yellowish or dark green to glaucous green, to 100 x 5—20 cm; spines stiff, acicular; roots mostly fibrous; flowers 22-50 mm, pericarpel and tube with scant wool and few bristles; nectary large, rounded drop-shaped at the base 30. subgibbosa Stem red brown, to 20 x 3-10 cm; spines stiff, acicular, 8-13 mm; flowers 20-30 mm, pericarpel and tube with dense wool and bristles; nectary small, flattened at base 31. sociabilis Spines mostly hairlike, very thin, intermixed with stiff straight acicular spines; stem to 20 x 4-10 cm; flowers 22-30 mm; pericarpel and tube with dense wool and bristles 33. villosa Spines contorted, often interwoven over stem apex; stem to 15 x 4-14 cm; flowers 40-50 mm; pericarpel and tube with variable covering of wool, from almost naked to densely covered, tube often very long 32. senilis

Note: All new names required by the following treatment are validated in Appendix I (pp. 117-119). Synonyms, with places of publication and details of typification, are given in Appendix IV (pp. 131-169). Material of the Kattermann (FK» field collections cited is deposited at the Desert Botanical Garden, Phoenix, Arizona (herbarium acronym: DBG).

Sect. I. Eriosyce Stem medium to large; areoles with white wool of variable length. Spines variable in number and size, mostly upcurved (except in subsect. Islaya, where the central spines are often deflexed). Flowers from young or old areoles, shortly funnel­ form, yellow to reddish; pericarpel with bristles from the axils of bract-scales, these sometimes only from uppermost axils and stiffly spinelike, never porrect; nectar-chamber simple. Fruit various, dehiscent by a basal or modified basal pore (enlarging 3-10x in subsect. Islaya >. Seed variable in size, hilum sunken, micropyle not or scarcely Burrounded hy special cell structures, hilum rim never raised at micropyle, intercellular depressions sometimes present on testa. D I S T R I B U T I O N . Chile, Peru and Argentina. Diagnostic features: Seeds with large, often deeply sunken hilum, lacking special cell structures around micropyle.

28

FIGURE 2

1-4 E. aurata var. aurata 1, F K - 1 7 1 ; 2 , F K - 1 7 ; 3. FK-5.45; 4. F K - 3 I

E. aurata mr. spinibarbia FK-808 fi

E. rodentiophila

FK-36

Kattermann:

Eriosyce

Kattermann:

Eriosyce

Sect. I. Eriosyce subsect. 1. Eriosyce (spp. nos. 1, 2) Stem large, globular, rarely elongated, 15-50 cm diam.; ribs numerous, 18—42 or more, the number increasing with age, slightly to deeply depressed between areoles, thus forming tubercles, 1-3 cm high; areoles with wool, but never long floccose; spines numerous, stiff acicular, always ± upcurved; roots fibrous. Flowers from areoles formed in previous years; buds, open flowers and ripe fruit sometimes contemporaneous; pericarpel and tube covered in long, very dense white wool, with stiff spine like bristles from upper bract-scale axils, tips of bristles often projecting beyond perianth; stigma-lobes clasped together, upright or spreading; nectary simple. Fruit ovoid, thick-walled, greenish, without apparent elongation upon ripening, dehiscent by a basal pore. Seed large, 1.5—3 mm, black or reddish black; hilum sunken, ovate to keyhole-shaped; micropyle not clearly separated within hilum region. DISTRIBUTION (MAP 2). Chile, western side of the Andes, 25-34\S. At the northern end of this distribution E, rodentiophila inhabits the coastal area between 25-26 S. E. aurata ranges from 26°S, near the town of Copiapo at c. 1000 m above sea level, southward. In the central part of this distribution plants can be found at elevations as high as 2800 m. South of the town Coquimbo at ± 30°S, populations of E. aurata also reach down to sea level. c

Diagnostic features: Plants very large; flower and fruit very woolly; fruit thick-walled; seeds very large. 1. Eriosyce aurata (Pfeiffer) Backeberg Stem subglobular to globular, rarely elongated, green, 10-50 cm diam. or more; apex naked to slightly woolly; ribs 24-42 or more, 1.5-3 cm high, barely to deeply depressed between the areoles; roots fibrous. Flowers from older areoles, arranged in spirals, with buds opening sequentially over an extended period; buds, flowers and fruit contemporaneous; flowers 3-3.5 x 2.2 cm, funnelform; pericarpel and tube wall thick-fleshy, with many bract-scales, completely covered by dense wool, bristles near top of tube stiff, spinelike, erect, 15 mm, often overtopping the perianth-segments; tube 12 mm; perianth-segments 15 x 3 mm, erect, yellow to reddish; style 13-16 x 1.5-2 mm, cream to pink; stigma-lobes 10-14, 2-4 mm, tips clasped together or upright, not constricted at base, cream; ovary locule apex updrawn; nectary simple, 2-3 mm. Fruit 4 x 2 cm. not elongating upon ripening, greenish but densely or loosely covered by wool, thick-walled, fleshy, bract-scales on fruit large (var, aurata) or small (var. spinibarbis), the seeds loose within; dehiscence by a basal pore, seed dispersal mechanism not known; perianth remnant attachment area not visible, 45-60 /( of fruit diam. r

E. aurata was previously known by the misapplied name, E. veratistes, and more recently by the junior, correctly applied synonym, E, sandillon (see Appendix IV). The following varieties can be distinguished: Fruit covered by dense wool, bract-scales on fruit large Fruit loosely covered by wool, bract-scales on fruit small

a. var. aurata b. var. spinibarbis

a. var. aurata Stem 10-50 cm diam. or more; areoles large, 14-18 x 6^8 mm, 10-15 mm apart, indumentum long but not Ooccose, covering stem apex; spines thickly acicular, curved upward, yellowish to later greyish, radials 12-15, 30-40 x 0.82 mm, centrals 4-6, strongly curved, 35—45 x 1.5 mm, base bulbous. Seed ISEM Plates I, II| 1.9-2.9 x 1.3-2 mm. asymmetry ratio 0.45-0.6; testa-cells isodiametric, testa not ribbed, black; hilum deeply sunken, keyhole-shaped (pyriform) to almost ovate; micropyle in narrow section of hilum, hilum rim thickened and modified, not raised. (Seed size and testa appear to vary from population to population).

30

Kattermann: DISTRIBUTION (MAP 2). Chile, from 28-33°S. At the southern end of the distribution the plants grow mostly above 1800 m. Near Coquimbo populations almost reach the coast at about 300 m altitude.

Erioayce

•

£. aurata van aurata

•

E. aurata vat.

•

E.

spinibarbts

rodentioohila

SPECIMENS EXAMINED. CHILE. Huasco: El

Transito, La Pampa, FK-822 iE. lapampaensis Ritter); S of Conay, 2300 m, FK-823; S of Mina Algarrobo, FK-171 iE. algarrobensis Ritter); Domeyko, Pajonales, FK-400 {E. algarrobensis Ritter). Elqui: La Serena, Tres Cruces, FK-454, 536 (£. ausseliana Ritter, invalid); Junta de Chingolos, FK-535. Limari: Ovalle, Rio Los Molles, FK-17; Soco, FK-449 iE. ihotzkyanae Ritter); Estancia Frai Jorge, FK-20a; Portezuela Pangue, FK-31; Combarbala, FK-190 iE. combarbalensis Backeberg); Pta Teniente, FK-486. Choapa: Salamanca, Cuncumen, FK-421. San Felipe de Aconcagua: Putaendo, FK-310. Rudolfo Wagenknecht (La Serena) reports a yellow-flowering, yellow-spined form from Cuesta Las Breas, Prov. Limari. b. var. spinibarbis Ritter Stem 16-25 cm diam., apex naked; areoles with short wool, 8 x 4 mm wide, 8 mm apart; spines thinner than in var. aurata. 0.55—0.75 mm diam., dark horn-coloured, radials 14—16, 25 mm, centrals 6-8, 25-30 mm, strongly curved upward. Fruit with wool only partly covering epidermis, bract-scales small. Seed ISEM Plate U 2.04 x 1.81 mm, asymmetric ratio 0.5; testa red-blackish-brown, slightly ribbed; hilum wide keyhole-shaped to ovate, deeply sunken, 0.36 x 0.48 mm; micropyle in narrow section of hilum, hilum rim barely enlarged at micropyle. DISTRIBUTION. Chile, from N of Copiapd, S to the Rio Jorquera. SPECIMENS

EXAMINED. CHILE. Copiapo:

Los Loros, FK-831; Rio Jorquera, FK-524; Puquios, FK-808. This variety is more or less intermediate with E. rodentiophila.

M A P 2. Distribution of Eriosyce

subsect.

Eriosyce.

Kattermann:

Eriosyce

2. Eriosyce rodentiophila Ritter Stem subglobular to globular, 12-32 cm diam.; apex covered by long white wool; ribs 18-35, obtuse, 12-18 mm high, more or less deeply notched to form tubercles; areoles 10 x 5-6 mm, 6-10 mm apart, with long wool; spines thick acicular, 1 mm thick, horn-coloured later grey, curved upward, radials 12-15, 20-30 mm long, centrals 4-8, 30-50 mm; roots fibrous from a short turbinate taproot. Flowers from older areoles, 25—40 x 22-30 mm, reddish; pericarpel and tube covered by dense white wool, with many bract-scales whose tips are long and almost spinelike, bract-scale axils near top of tube bearing stiff erect spinelike bristles to 15 mm, projecting above perianth; peri a nth-segments 8 x 2.5 mm; style 8-15 x 2 mm at base, cream; ovary locule apex updrawn; stigma-lobes 10—12, 5-7 mm, spreading, cream; nectary simple, 1.5-3 mm. Fruit (based on F. Rttter's description! 25-30 x 20-25 mm, reddish yellow, half covered by wool, wall 3-4 mm thick; not self-dehiscent, seed loose within. Seed [SEM Plate II] 1.9 x 1.36 mm, asymmetry ratio 0.3; testa tuberculate to finely ribbed, blackish to greyish brown; hilum ovate, 0.5 x 0.3 mm, deeply sunken; micropyle at narrow end, hilum rim without modification. DISTRIBUTION ( M A P 2t. Chile, along the coast beginning S of Chanaral to N of Taltal (Quebrada Matancillas). SPECIMENS EXAMINED. CHILE. Antofagasta: Taltal. FK-36; Quebrada Matancillas, 880 m, 12 June 1990, Taylor 740 (K, photos.); N of Paposo, Anderson in FK-851 (seeds); Cifuncho, FK-118, 119, 124. Chanaral: Caleta Flamenco, FK-136; Barquito, FK-394. 832.

PLATE 1 1

Fieldwork in Chile: searching for E.

napina

south of Freirina. L to R: Beatrice and Gerry Barad. Betty and E d G a y . 2

Chilean botanist A d r i a n a Hoffmann at the author's home, with a collection of Chilean cacti laid out according to latitude of origin.

:) E. aurata FK-17 at Rio Molles ( 1 9 7 7 ) .

32

Kattermann:

Eriosyce

Sect. I. Eriosyce subsect. 2. Pyrrhocactus (spp. nos. 3-7) Stem semi-globular to globular to elongated, 4-20 cm diam.; ribs many, 7-18, depressed or deeply notched between areoles; areoles large, with whitish wool; spines variable, 10-35, 1.5—4.5 cm; roots not always fibrous, sometimes with a shortly tapering, cone-shaped taproot. Flowers from old or young areoles; pericarpel and tube with short wool, bristles sometimes present on pericarpel, bristles thin hairlike, contorted or straight; nectary simple. Fruit short-ovoid, thinwalled, slightly or not elongating upon ripening, immediately drying out when ripe, dehiscent by a modified basal pore (indehiscent in E. bulbocalyx); attachment area of perianth remnant large. Seeds large 1.6-2.3 mm, testa dark brown or black; hilum large, sunken, oval to keyshaped; micropyle not clearly separated within the hilum region. D I S T R I B U T I O N ( M A P 3). Endemic to Argentina; eastern side of Andes from 24-38°S. The northernmost species is E. umadeave, which grows at Puerta Tastil, W of Salta (Salta), at about 2600 m elevation and possibly in Jujuy. There are no further populations recorded until 29 S, from where populations of the various species are ± continuous southwards until 38 S. b

U

Diagnostic features: Fruit dry when ripe; seed with large, often deeply sunken hilum. 3. Eriosyce bulbocalyx (Werdermann) Kattermann Stem globular to elongated, glaucous green, to 20 x 8-12 cm; ribs 13, notched, 1 cm high, obtuse; spines up to 20, to 30 mm, all strongly curved upward, radials 8-12, centrals 5-8; roots fibrous. Flowers produced from stem apex, sometimes from older areoles, 2.5 x 2 cm; peri a nth-segments 12 x 2 mm, narrow lanceolate, tip acute; pericarpel with tufts of wool and occasionally with bristles from the bract-scale axils; tube with tufts of wool and 3-4, 3 mm bristles; style 14 mm including 8, 1.5-3 mm stigma-lobes, stigma-lobes clasped together; nectary simple, narrow tubular, 2-3 mm. Fruit short oval elongating c. 1.5x at maturity, 10-15 x 7.5-10 mm, at first bright red and fleshy but within a week of maturity wall drying, becoming thin and almost translucent, indehiscent, seed loose within but sticking to the inside of the fruit wall; mode of seed dispersal not known. Seed (SEM Plate IV] 1.6 x 1.3 mm, black, not ribbed; hilum very wide oval, with hilum rim curved outward, with some strophiolar tissue projecting above rim. D I S T R I B U T I O N ( M A P 3I. Argentina, from Marayes to Patquia. S P E C I M E N S E X A M I N E D . ARGENTINA. San Juan: Marayes, FK-709. La Rioja: Los Colorados, near Patquia, FK-706, 680 m. Hunt s.n. (photo., see Plate 4.1.). 4. Eriosyce strausiana (Schumann) Kattermann Stem globular to elongated, to 20 x 8-15 cm, apex naked; ribs 12-14, notched halfway, 1 cm high, obtuse; spines black-grey with purplish tints to dark horn-coloured, all upcurved; roots fibrous from short turbinate taproot. Flowers from young areoles, 3.5-4 x 5 cm; pericarpel and tube with tufts of wool and bristles from bract-scale axils; tube broad funnelform, 15 mm; perianth-segments 25 x 5-7 mm, pale yellow or sometimes pale reddish; style 22-25 x 2 mm, light yellow; ovary locule apex flat to slightly updrawn; stigma-lobes 8-12, 1.5-3 mm, erect, cream-yellow; nectary simple, 2-3 mm. Fruit short oval, 7-10 x 5-10 mm, without elongation upon ripening, greenish, thin-walled, hard and dry when ripe, seed loose within, dehiscent by a basal pore, seed dispersal by freefall from fruit; perianth remnant attachment area wide, 95'# of fruit diam. Seed 1SEM Plate III 1.93 x 1.48 mm, asymmetry ratio 0.8; testa smooth, tuberculate to slightly ribbed, sometimes keeled, brownish, cells isodiametric, low domed; hilum keyhole-shaped, 0.64 x 0.4 mm, deeply sunken; micropyle in groove, hilum rim modified, not raised.

34

FIGURE S 1

I-'., strausiana

var. strausiana

FK-576

2

E. strauniana

var. tttrauniana

FK-572

.

— ........

3

E. strauHiana var. pachacoensis

....

.

.

™,

E

For captions D

O

pages

to colour plates

on

36 and 37. see page ЗИ

FK-582

SB

Kattermann: P L A T E 2 'page

Eriosyce

The following varieties of E. strausiana can be distinguished:

36)

Stem to 20 x 8-15 cm; ribs 12-14 Stem to 10 x 8 cm; ribs 13 1

2

3

4

a. var. strausiana b. var. pachacoensis

a. var. strausiana Stem to 20 x 8-15 cm, dark blackish green; areoles 10 x 5 mm, 8 mm apart; radial spines 12-14, 25-30 mm, centrals 6-8, 30—40 mm, with bulbous bases. Flowers 4 cm. DISTRIBUTION. Argentina, c. 30-40 S, the northern limit near San Jose de Jachal. a

5

S P E C I M E N S E X A M I N E D . ARGENTINA. Mendoza: Potrerilla,

6

FK-576 (П., fr„ seed, photos); Uspallata, FK-572; Meglioli in FK-583 (П., seed).

1

E. aurata № 4 1 ,

2

/ . aurata F K - 8 2 3 . /-.. aurata var. spini­ barbis F K - B 0 8 .

3 4

£ . aurata var. spini­ barbis F K - 5 2 4 .

5

E. rodentiophila

6

i?. rodentiophila Taylor

FK-36.

7 4 0 fpAofo:

Taylor)

P L A T E 3 (род.' S7J

1

2

3

4

1-3

5

E. rodentiophila Taylor

7 4 0 /photos:

Taylor) 4. 5

38

E. bulbocalyx

FK-706.

b. var. pachacoensis (Rausch) Kattermann Stem 10 x 8 cm, light grey-green; ribs 13, notched to form tubercles to 10 mm; areoles oval, 5 mm, with white wool; radial spines 8-12, in 4—6 pairs and one directed downward, 10-15 mm, ft appressed, whitish grey to pink, centrals 1-2, curved upward, grey-pink with black tip. Flowers 3.5 cm, yellow; pericarpel and tube green, bract-scale axils with wool and long reddish brown bristles; stigma-lobes 8, yellow. Fruit 8 x 5 mm, red brown. Seeds 1.5 x 1 mm. DISTRIBUTION. Argentina, San Juan, near Pachaco. SPECIMEN E X A M I N E D . ARGENTINA. San Juan: near Pachaco, FK-582 (fl., fr., seed, photos). 5. Eriosyce umadeave (Werdermann) Kattermann Stem globular to slightly elongated, to 40 x 10-20 cm, dark green, apex naked; ribs c. 18-21 or more, deeply notched, about 1 cm high, somewhat spiral, areole with a small chin below, large, elongated, 9-12 x 4-7 mm, 6-10 mm apart, with white wool; spines 20—35 or more, to 45 mm, white to violet, in older plants not easily divisible into radials and centrals, all ± curved upward, base bulbous; roots fibrous. Flowers from older areoles, 3-3.5 cm, funnelform; pericarpel and tube with bractscales bearing wool in their axils and near apex with 1 or more bristles to 15 mm, these straight to slightly contorted; perianth-segments 15 x 3 mm, pale yellow; filaments numerous; style 15 x 1.5 mm, cream-yellow; ovary locule apex fiat; stigma-lobes 12, 5 mm, spreading, cream yellow; nectary simple, 3 mm. Fruit (descr. from literature) elongated, 3—4 x 1.5-2 cm, light brownish, surface like the pericarpel. Seed I SEM Plate II] large, 1.79 x 1.36 mm, asymmetry ratio 0.4; testa deeply ribbed, brownish, cells not easily distinguished; hilum ovate, 0.48 x 0.36 mm, deeply sunken; micropyle in slightly modified portion of hilum rim, rim not raised. DISTRIBUTION. Argentina, prov. Salta, Andes to the W of Salta.

Kattermann: SPECIMENS

EXAMINED.

Erioayce

ARGENTINA.

Salta, Puerta Tastil, 2650 m, FK-652; loc. cit., Hunt s.n. (photo; see Plate 4.2.); loc. cit., R. Kiesling (seed), cult. FK-843. Werdermann's description was based on material from F. Ritter and Stumer (lectotype, В I. However, the picture furnished by Ritter does not represent E. umadeave, but rather a form of E. curvispina. It is possible that Bodeker, from whom Werdermann received the material, mixed up the information. 6. Erioayce andreaeana Kattermann Stem globular, later cylindric, 15-18 x 5-7 cm. dark grey green, apex naked; ribs 11-14, barely notched, 6 mm high; areoles 6 x 3 mm, 7 mm apart; spines horn-coloured, some upcurved, radials 12, 12-18 x 0.6 mm, ± radiating, slightly curved, centrals 4, cross wise, 19-40 x 0.8-1 mm, with bulbous bases; roots fibrous from a short turbinate taproot. Flower from young areoles, 3 x 3.7 cm, funnelform; pericarpel and tube with clusters of 10-15 brown, 10 mm bristles and short white wool from bract-scale axils; perianthsegments 20 x 3-5 mm, reddish; style 15 x 1.6 mm, cream-yellow; ovary locule apex updrawn; stigma-lobes 8-10, 2.5-4 mm, cream-yellow, erect; nectary simple, base slightly flattened, angular, 2 x 1 mm. Fruit with slight elongation upon ripening, 1.3 x 1 cm, covered by many bristles, hard and dry shortly after ripening; dehiscent by basal opening with lid like cap remaining on the fruit, seed free within but tightly packed; perianth remnant attachment area wide, 55% of fruit diam. Seed (SEM Plate II] 1.45 x 1.26 mm, asymmetry ratio 1.0; testa evenly tuberculate, not ribbed, brown, cells low domed, isodiametric; hilum oval, 0.6 x 0.45 mm, deeply sunken; micropyle not distinct, hilum rim not modified, flat. D I S T R I B U T I O N ( M A P 3). Argentina, in the higher mountains of the southern Sierra Famatina (La Rioja). SPECIMEN

EXAMINED.

ARGENTINA.

Rioja: Sierra Famatina, FK-593.

La M A P 3 . Distribution of subsect.

Pyrrhocactus.

Kattermann: E. andreaeana

iPyrrhocactus

andreaeanus

Eriosyce

(Backeberg) Ritter)

was included in Acanthocalycium by Donald (in Ashingtonia 1: 125. 1975). who has maintained as recently as 1990 (in Bradleya 8: 99) that the genus 'could be subsumed into Pyrrho­ cactus I Neoporteria'. As noted earlier (p. 20), I cannot find evidence to support this. The flower of E . andreaeana has the filaments evenly inserted over the lower one-third of the tube, their attitude ± parallel with the style. The filaments of Acanthocalycium sens. str. are appressed to the tube wall with their tips curved inward. Also, its nectary is partly closed by a woolly plug, whereas E . andreaeana has an open nectary. The fruit of E . andreaeana dehisces by a basal pore, Acantho­ calycium by splitting of the side wall. In its distribution, Acanthocalycium is partly sympatric with populations of E . subsect.

Pyrrhocactus.

When he named this species (as Neochilenia andreaeana) Backeberg failed to preserve the type, and so that name and all those based on it, including Acanthocalycium

are technically invalid. The name Eriosyce mann is validated on page 117.

andreaeana

andreaeanum,

Katter­

7. Eriosyce villicumensis (Rausch) Kattermann Stem subglobular, 4—9 cm diam., light grey-green with whitish powdery surface, apex naked; ribs 7-11, broad obtuse, 7 mm high, barely notched between areoles; areoles oval, 5—8 x 2-3 mm, 6-15 mm apart, with white wool; spines black with bulbous base, radials 3—4 pairs, ± appressed, 8-15 mm, centrals 1-2, 10—20 mm, curved upward; rootstock a short thick turbinate taproot. Flowers from young areoles, 28-35 x 30 mm, funnelform; tube and pericarpel with reddish bractscales, with white wool and brown 5 mm bristles in their axils; perianth-segments 11 x 3 mm, narrow lanceolate, orangeyellow; style 15 x 1.5 mm, cream-coloured to slightly pink towards top; ovary locule apex updrawn; stigma-lobes 10, 3 mm, clasped together, cream-yellow; nectary simple, 3 mm, base somewhat flattened. Fruit globular, 12 mm diam., dark brown, not elongating upon ripening, wall hard and dry shortly after dehiscence by basal pore with lid attached to fruit, seeds loose within; perianth remnant attachment area wide, 83% of fruit diam. Seed [SEM Plate II] 1.28 x 1.12 mm, asymmetric ratio 0.7; testa black-brown, not ribbed, cells isodiametric, medium domed; hilum keyhole-shaped, 0.5 x 0.36 mm, deeply sunken; micropyle in groove, hilum rim modified, slightly raised. DISTRIBUTION (MAP

3). Argentina, in the Sierra Villicum,

San Juan. SPECIMEN EXAMINED.

FK-584.

40

ARGENTINA. San Juan: loc. cit,

FIGURE 4

1 2

E. umadeave andreaeana I К •

3

E. villicumensis

4

E. bulbocalyx

FK-584

FK-706

Kattermann:

Eriosyce

Kattermann:

Eriosyce

Sect. I. Eriosyce subsect. 3. Islaya (sp. no. 8) Stem globular to elongated, 5—20 cm diam.; ribs many, to 25; areoles with long floccose wool, covering stem apex; spines many, 16-30, 1.2-1.6 cm, radial spines radiating, central spines porrect or deflexed; roots fibrous. Flowers from young areoles, in most forms sulphur yellow; pericarpel and tube covered by dense wool, bristles sometimes present on pericarpel, always present on tube, stiff, almost needle-like; stigma-lobes clasped together or spreading, yellow; nectary simple or modified by widening at base. Fruit clavate, thin-walled, red, much enlarged upon ripening, balloon-like, seed sometimes in a sac-like structure apex of fruit, dehiscent by a small basal pore; attachment area of perianth remnants variable, mostly small. Seeds 0.75-1.5 mm, black; hilum large, sunken, ovate to keyhole-shaped; micropyle partly surrounded by slightly modified hilum rim. D I S T R I B U T I O N ( M A P 4). Coastal area of southern Peru and northern Chile, from 14-18°S. Diagnostic features: Fruit much elongated, balloon-like; flowers mostly yellow; pericarpel and tube with dense wool and many bristles. Note: I have not yet studied sufficient field-collected material and can as yet recognize one variable species only, E. islayensis. The seed material studied (obtained from commercial sources; see SEM Plate III, rows 1-4) indicates a close relationship to subsect. Eriosyce and a very close relationship between the different populations, but the flower material studied indicates otherwise. The flowers show every form of development of internal and external characters as observed in parts of subsect. Eriosyce and subsect. Horridocactus. 8. Eriosyce islayensis (Forster) Kattermann Stem globular, 5-20 cm diam., or sometimes elongated to more than twice the diam.; ribs numerous, to 25; areoles with long floccose wool covering stem apex; radial spines 12-22, short, radiating, 6-10 mm; centrals 4-7, 12-16 mm, thicker than radials; roots fibrous. Flowers from young areoles [Rauh & Backeberg report that in Islaya grandis flowers are also borne on older areolesl, wide funnelform and in most forms sulphur-yellow; pericarpel and tube covered by dense wool, bristles sometimes present on pericarpel, always present on tube; style variable in length, cream-coloured; ovary locule apex downdrawn to updrawn; stigma-lobes mostly creamyellow, clasped together or almost spreading; nectary simple or widened. Fruit clavate, thinwalled, balloonlike, red, 5-10x elongated upon ripening; dehiscent by a basal pore, seed loose or sometimes in a sac-like structure at apex of fruit; perianth remnant attachment area small, usually less than 50% of fruit diam. Seed [SEM Plate III] 1-1.5 x 0.9-1.1 mm, asymmetric ratio 0.4-0.55; testa mostly keeled, cells isodiametric to slightly elongated and then arranged ± in rows, colliculate; hilum ovate to keyhole-shaped, relatively large; micropyle in a region slightly or not modified by hilum rim. D I S T R I B U T I O N ( M A P 4). N Chile to S Peru, N along the coast to Rio Omas S of Lima. SPECIMENS EXAMINED. CHILE. Arica: Poconchile, FK-341. Specimens shown in Figure 5 (page 43) and Plate 6 (page 52) were raised from seed from various commercial sources.) Subsect. Islaya represents the part of the genus least understood by me. I have only collected the form described as /. krainziana in northern Chile. Until new field observations and collections are made I am lumping all species described up to now under E. islayensis. Seed and flower material studied suggests that several species and varieties may exist. There is also the question of the affinity of E . recondita. It is possible that the most northern population of this species may belong to subsect. Islaya. I could not resolve this question.

12

Kattermann;

FIGURE б

Eriosyce

Б. islayensis

1

tlslaya

copiapoidvs

2

(Islaya

divaricaliflora)

i

3

(Islaya

chalaensis)

P L A T E 4 (page

1 2

44)

E. bulbocalyx E. umadeave Tastil tphoto:

at

Puerta

Hunt)

48

Kattermann:

M A P 4. Distribution of subsect. P L A T E 5 (page

1

Islaya.

45)

2

1

/<:. strausiana

FK-576

2

E.

andreaeana

FK-593

3

3

5

4

6

/.. strausiana pachacoensis

var. 4

£. andreaeana

5

£. villicumensis

6 £. islayensis (Islaya

46

Eriosyce

FK-582

FK-593

FK-341 kraimiana)

Kattermann:

Eriosyce

Sect. II. Neoporteria Stem small to medium sized; areoles white-woolly, but never long-floccose. Spines never deflexed. sometimes thin and contorted. Flowers from young areoles only, funnelform to somewhat elongate-funnelform, yellow to reddish or fuchsia red; bract-scales on tube subtending bristles, these never spinelike but sometimes porrect; nectar-chamber simple or modified at base. Fruit various. Seed variable in size, hilum not sunken, micropyle surrounded by special cell structures (except in subsect. Chileosyce), testa lacking intercellular depressions. DISTRIBUTION. Chile.

Diagnostic features: Seeds with small, never deeply sunken hilum, often with special cell structures around micropyle. Sect. II. Neoporteria subsect. 1. Horridocactus (spp. nos. 9-23) Stem semi-globular to globular or sometimes elongated, 2-20 cm diam.; ribs many, 8-24, 0.3-2 cm high, notched or depressed between areoles, sometimes tubcrculate; areoles variable in size, often with a protruding chin below; spines few to many, 6-25, 0.2-5 cm, appressed to porrect or curved upward; roots fibrous or small to large napiform, or tuberous. Flowers from young areoles, wide to narrow funnelform, yellow to sometimes deep reddish; pericarpel and tube never with stiff, porrect bristles (see subsect. Chileosyce), wool not completely covering pericarpel and tube, wool from the bract-scale axils sometimes scarcely visible, the pericarpel and tube appearing ± naked; stigma-lobes variable in attitude, tightly clasped together or spreading; nectary simple to modified and widened at base, then ± angular, not rounded bowl-shaped as in subsect. Neoporteria. Fruit ovoid or sometimes clavate, thick- to thin-walled, red or greenish red, sometimes elongating before ripening, fruit wall not drying shortly after ripening; dehiscent by a basal pore; attachment area of perianth remnants small to large. Seed 0.7-1.7 mm, black or brown, testa colliculate, sometimes ribbed; hilum relatively small, not sunken, keyhole-shaped; micropyle in a clearly defined groove at the upper end of the hilum, testa-cells sometimes enclosing the micropyle completely and separating it from the hilum tissue. D I S T R I B U T I O N . 18-36^8, endemic to Chile and range similar to that of subsect. Eriosyce, but extending further north and south. Populations of both subsections can be found in the same hahitat. Subsect. Horridocactus hae a broader east-west distribution, and in altitude ranges from near sea level to 2800 m. Diagnostic features: flowers broadly funnelform, fruit a red berry, sometimes elongated, and ± naked. 9. Eriosyce recondita I Ritter > Kattermann The following subspecies can be distinguished: Stem 2-5 cm diam.; flowers white to pinkish; pericarpel and tube not covered by dense wool, bristles on tube and pericarpel brown 9.1. subsp. recondita Stem 3-10 cm diam.; flowers yellow, with variably dense wool and white bristles (sometimes absent) from axils of bract-scalea 9.2. subsp. iquiquensis 9.1. E. recondita subsp. recondita (Ritter) Kattermann Stem subglobular, 2-5 cm diam., sometimes branching, dark grey green, apex sunken, spiny; areoles 4-6 x 2-3 mm, 5-10 mm apart, sunken; ribs 10-12, 5 mm high, dissolved into tubercles, with a chin below each areole; spines thin, needlelike, black, radials 6-10, 5-10 mm, centrals 1-2, 10-15 mm; rootstock a long turbiniform taproot with slight neck near apex. Flowers borne from young areoles, funnelform, 3-4 x 2.5-3.5 cm; pericarpel and tube bract-scale axils with grey

17

Kattermann: floccose wool and brown, porrect bristles; style 20-33ram,pale reddish; ovary locule apex downdrawn; stigma-lobes 8-12,3-5 mm, erect or clasped together, pinkish; nectary widened at base, 1-2 mm. Fruit elongating, seed contained in a sac, dehiscence by a basal pore, fruit attachment period unknown; perianth attachment small? Seed 1.4 x 1.0 mm, testa very finely tuberculate, dull black; hilum keyhole-shaped, 0.40 x 0.20 mm, micropyle in modified area of the barely raised hilum rim. DISTRIBUTION (MAP 5). Chile, region of Antofagasta. SPECIMENS

EXAMINED.

CHILE.

Eriosyce

•

Iquique:

Caleta Junin, FK-338. Tocopilla: S of Tocopilla, FK-776; Guanillo del Sur, FK-366. I presently place this species in subsect. Horridocactus even though it could belong to sect. Eriosyce subsect. Islaya. More morpho­ logical studies based on authentic material are necessary to reach a sound conclusion.

recona la iquique

ISIS

Гs

VF i s a g u a

i

ъ

Iquique

I \

>

\ \

1 |

\

\

\

° Calama

\ 'v San P de A t a c MM

69° of Eriosyce

6 recondita

i

FIGURE 6 1

E. recondita

2

T. recondita

subsp. recondita subsp.

FK-369.

iquiquensis

FK-366 3

/•. recondita

4

E. recondita

5

E. chilensis

subsp. iquiquensis subsp.

sens. str.

iquiquensis

from a more northern population, FK-776. FK-3 (see page

stigma 4-5 mm.

48

I

E. rec vnaita v a r . E. rec ONDITA v a r .

of

DISTRIBUTION. Chile, Prov. Iquique and Tocopilla. CHILE.

iquiquensts

• A

л

21 9.2. subsp. iquiquensis (Ritter) Kattermann Stem 3-10 cm diam., dark green; ribs 13, notched into tubercles with a chin below each areole; areoles sunken; rootstock a short, conical turbiniform taproot. Flowers borne from young areoles, narrow funnelform to rooopilla sometimes broader funnelform, 1.5-2 cm; pericarpel and tube bract-scale axils with white floccose wool, bristles lacking; style 10-12 mm; ovary locule apex unknown; stigma-lobes 2-3 mm, erect or clasped together; nectary 1-2 mm. Fruit elongating, 23" seed loose within, dehiscent by a basal pore, attachment firm; perianth remnant attach­ ment area small, 45% of fruit diam. Seed yym An ofagasta (SEM Plates XIII, XIV1 1.25 x 1.25 mm, I asymmetric ratio 0.5; hilum ± reverse 70° keyhole-shaped, with micropyle area larger, 0.30 x 0.18 mm, micropyle not modified. MAP 5. Distribution

EXAMINED.

recondite

Anto­

fagasta: Morro Moreno, FK-369; N Antofagasta, FK-777.

SPECIMENS

var. var.

\

Ъ А г к X/ IV

E. recondite

A £. recondite

102); П. 3.2 cm,

Kattermann:

Eriosyce

49

Kattermann:

Eriosyce

10. Eriosyce curvispina (Bertero ex Colla) Kattermann Stem subglobular to globular, rarely branching, rarely elongated, 8-20 cm diam., apex naked; ribs many, 14-24, variable in height from 1-2 cm, deeply notched or depressed between areoles, with an often large chin-like protrusion below each areole; areoles variable in size, with short wool; spines many, horn-coloured, tipped darker, later grey, thin to thickly awl-shaped, mostly curved upward, radials 10-15, 10-30 mm, centrals 4-8, 20-30 mm, mostly curved upward to inward, often with bulbous bases; roots ± fibrous, but sometimes with a short turbinate taproot. Flowers produced from young areoles, forming a circle around stem apex, wide funnelform, 3-5 x 3-5 cm; pericarpel and tube with variable density of small bract-scales rarely > 2 mm, their axils with very short tufts of wool barely visible, upper axils with thin white bristles; perianthsegments variable in length, 7-12 mm wide, yellow-edged with a reddish midstripe of variable width; filaments inserted over two-thirds of tube; style 16-25 x > 2 mm, thick, cream-yellow to reddish, often more reddish towards apex; ovary locule apex downdrawn (annulated); stigmalobes 10-15, 2-5 mm, erect or sometimes clasped together, tips ending abruptly, base often constricted, mostly cream-yellow to red; nectary simple (tubular), not widened at the rounded base, 2—4 mm, top opening by 1-2 mm. Fruit short-oval, elongating about l.l-1.4x upon ripening, fleshy, thick-walled, reddish when ripe, hollow, seed loose, dehiscent hy a basal pore, fruit remaining attached long after ripening; perianth remnant attachment area broad. Seed ISEM Plate IV| variable in size 0.9-1.62 x 0.88-1.43 mm, asymmetric ratio of 0.6-1.0; testa evenly tuberculate to ribbed, black to brown, cells medium to high-domed; hilum keyhole type, micropyle in a groove, hilum rim modified, raised, micropyle sometimes separated from hilum by fused testa-cell tissues. DISTRIBUTION ( M A P 6). Chile, 27°15'S, between Copiapo and Caldera (Prov. Copiapo), extending S almost uninterrupted to Rio Maule (Prov. Talca) at 35 30'S. At various points of its distribution branches of evolutionary development in a downhill direction can be observed. Some of these branches evolved slightly different characters and the taxa concerned are here recognized as varieties. These varieties are also often an intermediate development toward different species. In total about 150 population samples of this species were collected, propagated and studied. Herbarium specimens of most of these are in preparation and will be deposited at DBG. C

K U Y T O V A R I K T I K S OK E. 1. 1.

Plants never branching . . . Plants sometimes branching

2. Nectary simple (tubular), 3 mm 2. Nectary simple, 4—5 mm 3. Spines medium in thickness, strongly curved 3. Spines stouter, to 3 mm thick, barely curved

CURVISPINA

2 f. var. mutabilis 3 g. var. robusta 4 e. var. tuberisulcata

5

4, Spines 20 mm or longer 4. Spines 10-15 mm

d. var. choapensis

5 Flowers 5.5 cm, perianth-segments 6 mm wide 5, Flowers 4.5 cm

a. var. curvispina 6

6

50

Perianth-segments yellow Perianth-segments with deep red midstripe

b. var. aconcaguensis c. var. armata

Kattermann:

Eriosyce

a. var. curvispina Stem semiglobular to globular, 8-12 cm in diam., apex naked, dark green; ribs 14-16, 1-1.5 cm high, notched, with chin-like protrusion below areole; areoles 6 x 4 mm, 6 mm apart; spines horn-coloured, later grey, thin to thickly awl-shaped, strongly curved upward, radials 10-12, 10-20 mm, centrals 4, 20 mm, mostly curved upward to inward, bases bulbous; roots fibrous. Flowers produced from young areoles, borne in a circle around stem apex, wide funnelform, c. 5.5 cm long and wide; perianthsegments 35 x 6 mm, yellow-edged, with a central reddish stripe of variable width; style 25 x 2.7 mm, pale reddish, often more reddish towards apex; ovary locule apex down drawn (annular); stigma-lobes 10, 4 mm, erect or clasped together, cream yellow to pale red; nectary 3 mm, base rounded. Fruit short oval, 1.5 x 1.4, fleshy, thick-walled, reddish when ripe, hollow, seed loose; perianth remnant attachment area broad, 56% of fruit diam. Seeds 1.32 x 1.19 mm, asymmetric ratio 0.6; testa evenly tuberculate, not ribbed, black, cells medium to high domed; hilum narrow keyhole type, 0.53 x 0.16 mm; micropyle in a groove, hilum rim modified and raised. DISTRIBUTION. Chile, throughout the range of the species. Some populations grow at up to 2000 m, while others can be found at sea level. SPECIMENS EXAMINED. CHILE. Choapa: Tilama, FK-318, 319; Cuncumen, FK-420. Petorca: Alicahue, FK-308, 309; Chincolcolo, FK-312; Petorca, FK-313; Pedegua, FK-314; Zapallar, FK-561. San Felipe de Aconcagua: Las Coimas, FK-265; Los Pates, FK-268, 269. Quillota: Cerro Campana, FK-I01; Cuesta Melones, FK-428. Los Andes: Los Andes, FK-2; Rio Blanco, FK-194. Chacabuco: Cuesta La Dormida, FK-256, 258, 259; Caleu, FK-257, 260; Rungue, FK-261. Melipilla: Cholqui, FK-305; Horcon de Piedra, FK-447. Santiago: Farellones, FK-240, 243; Cuesta Las Chilcas, FK-87. Cordillera: El Volcan, FK-274; La Ohra, FK-276; Lagunilla, FK-307. Talca: A. Hoffmann

s.n.,R. Ferryman

PlJVTE 6 (page

1

2

3

4

5

6

E. islayensis (Maya

divaricatiftora)

2

E. islayensis

3

E. islayensis

(I.

11,

4

chataensis)

copiapoides)

/•.'. islayensis (I. islayensis

f.

brevicylindrica)

5. В E. curvispina FK-279

P L A T E 7 (page

100. Cauquenes: Rio Cachapuel,

FK-279. b. var. aconcaguensis (Ritter) Kattermann Flower lemon yellow, perianth-segments to 10 mm wide; pericarpel and tube with fewer bract-scales. DISTRIBUTION. Chile, Cuesta Las Chacras, N of Llaillay. S P E C I M E N E X A M I N E D . San Felipe de Aconcagua: Cuesta las Chacras, FK-263.

52)

1

1

53)

1

2

3

4

5

6

E. curvispina var. curvispina F K - 1 0 I var.

curvispina

FK-274

var.

curvispina

FK-313

var. aconcaguensis FK-263 var.

horrida

var.

odoriflora

FK-208 FK-427

51

Kattermann: с. var. armata (Ritter) Kattermann Flower reddish, perianth-segments 24 x 4.5 mm. Fruit elongating to 1.5x diam. Seed much smaller, 0.95 x 0.98 mm, asymmetric ratio 0.7, testa finely tuberculate. DISTRIBUTION. Chile, from El Paico (Santiago) S and W, including mountains near Horcon de Piedra. S P E C I M E N S E X A M I N E D . CHILE. Talagante:

El Paico, FK-28I; Melipilla: Horcon de Piedra, FK-447.

Eriosyce Pichidangui, FK-6; Quilimari, FK-321; W of Tilama, FK-320; Cerro Santa Ines, FK-323, 427; Los Viloa, FK-411; Cuesta Cavilolen, FK-412. Petorca: Pta Molles, FK-4, 85. g. var. robueta (Ritter) Kattermann Pericarpel much elongated; nectary 4-5 mm. DISTRIBUTION. Chile, known only from the type locality in the region of Ocoa (Quillota). SPECIMEN

EXAMINED.

CHILE.

Quillota:

Ocoa, FK-262.

d. var. choapensis (Ritter) Kattermann Spines short, rarely > 20 mm, thin acicular. Seed 0.94 x 1.0 mm, asymmetric ratio 0.9; testa-cells arranged in ribs. DISTRIBUTION. Chile, from S of Salamanca to Cuncumen in the Rio Choapa valley. SPECIMENS EXAMINED.

Quelen, FK-424; Chuchini, FK-426.

CHILE. Choapa:

Salamanca,

FK-425;

e. var. tuberisulcata (Jacobi) Kattermann Spines very thickly awl-shaped, only slightly curved upward, radials 8-12, 5-12 x 8 mm, centrals 4-8, 10-25 x 1.5-2.5 mm. Flower reddish; perianth-segments 20 x 7 mm. Seeds small, l.l x 1.07 mm, asymmetric ratio 0.8; testa tuberculate, slightly or not ribbed, brownish; hilum keyhole-shaped, 0.36 x 0.25 mm, micropyle in modified area of the raised hilum rim. DISTRIBUTION. Chile, region of Valparaiso. S P E C I M E N S E X A M I N E D . CHILE. Valparaiso:

S of Valparaiso, FK-251 (neotype), 300; Quilpue, FK-208. f. var. mutabilis (Ritter) Kattermann Stem often branching by forming little plantlets from lower areoles; spines stiff, sharp-tipped, horn-coloured. Perianthsegments with variable reddish midstripes. Seed 0.94-1.23 x 1-1.09 mm, asymmetric ratio 0.5—1.0; testa ribbed, brown; hilum 0.44 x 0.24 mm, micropyle in groove, hilum rim modified, sometimes closed around micropyle. DISTRIBUTION.

Chile, N, S and E of Pto

MAP

FIGURE 7 1

54

var.

E. curvispina

Erioayce curviapina

curviapina

F K - 2 7 9 ; flower 5 . 5 cm. s t i g m a

4 mm. 2

var.

aconcaguensia

F K - 2 6 3 ; flower 4 . 2 c m ,

stigma 3.5 m m . 3

Pichidangui. SPECIMENS EXAMINED.

6. Distribution of

var.

armata

F K - 2 8 1 ; flower 4 . 2 c m . s t i g m a 4

mm.

CHILE. Choapa:

4

var.

choapensis

FK-426.

Kattermann:

Eriosyce

Kattermann:

FIGURE 8 1

E. curvispina

Eriosyce

P L A T E 8 I page var. tuberisulcata

FK-251

1

Flower 3 . 5 cm; s t i g m a 3 . 4 m m . 2

E. curvispina

3

E. curvispina

var. mutabilis

FK-427.

Flower 3 . 2 cm; s t i g m a 3 . 5 m m ; fruit 9 m m . var. robusta

Flower 5 . 2 cm; fruit 1.7 cm.

E. curvispina armata

FK-2G2.

2

57) var.

FK-281

E. curvispina chitapensie

(above)

var. FK-426

(below)

Kattermann:

FIGURE 9

(above)

F I G U R E 1 0 (page

1

E. curvispina

FK-313

2

E. curvispina

FK-101,

from Ccrro C a m p a n a ,

1

Eriosyce

59)

E. kunzei var. kunzei F K - 4 5 9 . Fl. 3 . 5 cm.

2

60)

1

2

3

4

5

6

E. kunzei var.

w h e r e it is sympatric with

trail si tenets

E.

3 . 5 cm.

garaventae.

P L A T E 9 Ipage

F K - 1 6 7 . Fl.

3

E. kunzei

FK-170

4

E. kunzei

F K - 1 9 . Fl. 3 cm.

E. curvispina var. robusta F K - 2 6 2 E. kunzei

FK-459

E. kunzei FK-19 E. kunzei var. transitensis F K - 1 6 7 /-.'. kunzei var. transitensis F K - 8 2 4 E. limariensis

58

FK-20

Kattermann:

Eriosyce

58

Kattermann: P L A T E 10 I page

ft"

1

E. limariensis

2

E. limariensis FK-13

3

£.

confinis

FK-443

4

E. confinis

FK-528

5

£.

aspillagae

FK-19a

FK-197

Eriosyce

11. Eriosyce kunzei (Forster) Kattermann Stem globular, somewhat flattened to elongated, green, 10(-20) x 13-14 cm, apex somewhat sunken and almost without spines (but spines sometimes covering apex); ribs 16-20, at base ± spiral, vertical above, obtuse, enlarged at areoles, notched into tubercles, the tubercles interconnected, below areoles forming a protruding chin; areoles 10—15 x 2.5 mm, 5-10 mm apart, wool dirty white, later naked; spines all curved upward to inward, at first yellowish with honey-brown tips, then greyish honey-brown, later ash grey, completely obscuring stem, radials 10-12, the lower ones shortest, fairly straight and pointed downward, 13-25 mm, the upper ones 40-45 mm, centrals 2-4, longer and thicker, 45-50 mm. Flowers 3.5 cm long and wide; perianth-segments 20 x 7 mm, edges whitish yellow with variable reddish midstripe; style 22 x 2.2 mm, cream-yellow; ovary locule apex down drawn; stigma-lobes 10-11, 4 mm, erect to spreading, tip ending abruptly, not constricted at base, cream-yellow; nectary simple. Fruit shortly oval. 2 x 1.5 cm, fleshy, thick-walled, reddish when ripe, hollow, seeds loose, dehiscent by basal pore, remaining attached long after ripening; perianth remnant attachment area broad, 55% of fruit diam. Seeds [SEM Plate VI 1.31 x 1.00 mm, asymmetric ratio 0.8; testa evenly tuberculate to ribbed, black, cells medium to high domed; hilum narrow keyhole type, 0.4 x 0.33 mm, micropyle in groove, hilum rim modified and raised, micropyle sometimes separated from hilum by coalescent testa tissues. DISTRIBUTION (MAP 7). Chile, mountains N and S of the Elqui valley, mostly above 1500 m. Ritter's interpretation of Fbrster's description as applying to plants from the Sierra Hornillos cannot be correct. Ritter's use of the name is based on the plant figured by Rumpler in Forster's Handbuch de Cacteenkunde, ed. 2, 571 (1886). The description does not match the plant figured. The original description (1845) mentions that the plants are covered by snow for long periods. It never snows in the Sierra Hornillos. The following varieties can be distinguished: Spines thinly acicular; stigma-lobes erect to slightly spreading: seeds only sometimes ribbed a. var. kunzei Spines thicker; stigma-lobes clasped together; seed deeply ribbed b. var. transitensis

62

Kattermann:

Eriosyce

a. v a r . kunzei DISTRIBUTION. Chile, mountains N and S of the Elqui valley, mostly above 1500 m. SPECIMENS EXAMINED.

CHILE. Huasco:

Domcyko, FK-170, 173,399. Elqui: Portezuela Fangue, FK-30, 457 (Horridocactus vallenarensis); Balala, FK-460; Junta de Chingoles, FK-533; Guanta, FK-459 (neotype). Limari: upper Rio Molles, FK-19; Portezuela Las Tres Cruces, FK-456. b. var. transitensis (Ritter) Kattermann Fruit elongating more, in some populations to > 2x diam., perianth remnant attachment area smaller. Seed 1.13 x 1.14 mm, asym­ metric ratio 0.6; testa very deeply ribbed. DISTRIBUTION. Chile, from c. 25 km E of Vallenar to Conay, at c. 2000m. S P E C I M E N S E X A M I N E D . CHILE. Huasco: El

Terron, FK-167, 820; Santa Juana, FK-817; Conay, FK-824. The related Eriosyce crispa apparently evolved at lower elevations. The transition from one species to the next occurs at about 800 m. At this point climatic conditions change from alpine to coastal. This elevation is also the upper range of Eulychnia. The difference between Eriosyce crispa and E. kunzei var. transitensis is the frequent occurrence in the latter of large tuberous roots, narrower funnelform flowers, spreading stigma-lobes, long wool on pericarpel and tube, and bristles over wider area of the tube and over most of the pericarpel.

MAP

7. Distribution of spp. nos. 1 1 - 1 4 , E.

E. limariensis,

E. crispa

and E.

kunzei,

conftnis.

12. Eriosyce limariensis (Ritter) Kattermann Stem grey-green, globular, 5-15 cm diam., apex without spines, flattened; ribs 13-20(-30), 8-15 mm high, notched halfway, widened at, and with chins below the areoles; areoles with white wool, 6-12 x 3-5 mm, 10-20 mm apart; spines brown with darker tips, somewhat upcurved, seldom straight, awl-shaped, radials 6-10, ± pointing sideways, 15-40 mm, the upper longest, centrals 1-4, when 4 arranged crosswise, 20—45 mm; roots fibrous from a short conical taproot, or sometimes large taproot with slight neck (see note below). Flower 3.5-5.5 cm long and wide, somewhat fragrant; pericarpel elongated, 6-12 mm, olive green to reddish, bract-scales 1-2 mm, with short wool, tube 8-15 mm, funnelform, at apex 14-20 mm diam., with 10-20 mm white bristles from upper bract-scale axils; perianth-segments 25-30 x 5-8 mm, sulphur yellow with wide purplish midstripes; filaments light yellow, reddish at apex, 5-12 mm, the uppermost

88

Kattermann:

Eriosyce

inserted longest, anthers light yellow, insertion over two-thirds of tube; style cream-yellow to red; ovary locule apex down drawn, much elongated, 8.5 x 3.5 mm; stigma-lobes 8-10, red-brown to light yellow, 4-6 mm, clasped together; nectary simple, 2-5 mm. Fruit short-oval, 2.7 x 1.5 cm, fleshy, thick-walled, reddish when ripe, hollow, seed loose, dehiscent by a basal pore, remaining attached long after ripening; perianth remnant attachment area broad, 73% of fruit diam. Seed (SEM Plate V] 1.00 x 1.07 mm, asymmetric ratio of 0.6; testa ribbed, black to brown, cells medium to high-domed; hilum narrow keyhole type; micropyle in groove, hilum rim enlarged, raised. DISTRIBUTION ( M A P 7). Chile, in the Rio Molles and Rio Limari valleys, from 2300 m toe. 200 m at the coast, S to Combarbala, N to Samo Alto and Coquimbo. S P E C I M E N S KXAMINED. CHILE. Limari: Molles, 1500 m, FK-18, loc. cit., 2300 m, FK-19a; Soco, FK-12; Embalse La Paloma, FK-13; Combarbala, FK-189; Guanpulla, FK-451; Pichasca, FK-453; Rio Limari, FK-480; road from Pan American Highway to Estancia Frai Jorge, FK-20. E. limariensis shows transition from E. curvispina to E. heinrichiana. Some populations have plants with turbinate taproots and sometimes the formation of a slight neck above the root occurs. 13. Eriosyce confinis (Ritter) Kattermann Stem green, semiglobular, later elongated, 6-8 cm diam., apex naked to spiny; ribs 13-15, small obtuse, with a small protruding chin below, and deeply notched ahove areoles; areoles with white to greyish wool, 5-12 x 3-7 mm, 5-10 mm apart, not sunken; spines grey brown to greyish black, only slightly curved, radials 10-12, 10-25 mm, centrals 4-7, 20-40 mm; roots fibrous from a short taproot without a neck. Flowers 3 cm diam., funnelform; pericarpel grey green with small reddish bract-scales, with small flocks of white wool in their axils; tube 12-13 mm, 3 mm wide at base, 11 mm wide at apex, funnelform, narrowed above pericarpel, outside grey olive green, with bract-scales and wool as for pericarpel, tube apex with few white, thin bristles; perianthsegments 18-20 x 5 mm, narrower at base, tips pointed, silky, shiny with reddish midstripes, the outer segments redder, merging into red olive bract-scales; filaments white, the lowest 7 mm, the uppermost 10 mm; anthers lemon yellow; style exceeding anthers, c. 20 x 1.5 mm, carmine red; ovary locule apex down drawn; stigma-lobes 8, rose-yellowish to reddish, 4 mm; nectary simple to modified by slight to much enlargement at base, 3-4 mm. Fruit hollow, red. fleshy, 1.5 x 1 cm, dehiscent by a basal pore, remaining firmly attached for a long time; perianth remnant attachment area small, 44% of fruit diam. Seed |SEM Plate VI| 1.47 x 1.33 mm, asymmetric ratio 0.5; testa black to brown, dull, evenly tuberculate, not ribbed; hilum keyhole-shaped, 0.55 x 0.40 mm, micropyle in groove, hilum rim much enlarged, almost closing micropyle area, slightly raised to almost fiat. DISTRIBUTION (MAP 7). Chile, throughout the Copiapo Valley, beginning E of Copiapo, extending S to the Sierra Hornillos and W almost to the coast E of Caldera. S P E C I M E N S E X A M I N E D . CHILE. Copiapo: E of Copiapo, FK-141, 810; c. Sierra Hornillos, FK-143, 396, 528; Monte Amargo, A. Hoffmann 178 in FK-443, FK-807; E of Caldera, FK-487. The various populations of E. confinis clearly show a transition from E. curvispina to E. taltalensis. The seeds are still part of E. curvispina, but the flowers change toward the flowers of E. taltalensis by a slight widening at the base of the nectary and narrower funnelform tube. I place the plants from Morro Copiapo with E. taltalensis because of the reduction in size of the seed.

01

Kattermann:

Eriosyce

F I G U R K 11 1

E. limariensis

2

E. confinis

FK-443

F K - 2 0 , П. 5 c m , s t i g m a в m m .

.1

E. confinis

F K - 5 2 8 . П. 3 . 5 cm.

№

Kattermann:

Eriosyce

14. Eriosyce crispa (Ritter) Kattermann Stem flattened, subglobular or slightly elongated, 5-10 cm diam., blackish grey green; ribs numerous, 13-16, 0.5-1 cm high with a chin below each areole, notched or depressed, forming tubercles; spines black to brown, sometimes hairlike, curved up and sometimes contorted, radials C. 6-10, 15-50 mm, centrals 1-5, 20-80 mm; rootstock a thick, almost tuberous taproot, often narrowed and necklike at apex. Flower borne from young areoles, 3.5-5 cm long and wide, funnelform; axils of pericarpel and tube bract-scales with floccose wool and white bristles; perianth-segments 20-25 x 5-7 mm, whitish with reddish midstripes; style 25—30 x 1.9-2.2 mm, reddish; ovary locule apex down drawn; stigma-lobes 8-12, 5-7 mm, spreading, pale red; nectary simple, 2-4 mm. Fruit elongated oval, elongating 2-3x upon ripening, bract-scale axils with floccose wool tufts, sometimes with bristles, thin-walled but fleshy, reddish when ripe, dehiscent by a basal pore, remaining firmly attached for a long time; perianth remnant attachment area small, 35-457r of fruit diam. Seed [SEM Plate VI] 0.98-1.1 x 0.97-1.1 mm, asymmetric ratio 0.6-0.8; testa deeply ribbed to smooth-tuberculate, blackish to brownish; hilum keyhole-shaped, 0.44 x 0.24 mm; micropyle two-thirds surrounded by enlarged hilumrim,rim not raised. DISTRIBUTION (MAP 7). Chile, from E of Vallenar, W toward the coast, and from Huasco to N of Totoral Bajo. Further east, E. crispa

merges into E. kunzei

var.

transitensis.

KEY TO INFRASPKCIFIC TAXA OF E . CRISPA

1. Stem subglobular, to globular

2

1. Stem globular or often elongated, spines thickly acicular 14.2a. subsp. & var. atroviridis 2. Seeds coarsely ribbed; spines thin, hairlike, tortuous, black

14.1. subsp. crispa

2. Seeds evenly tuberculate; spines various

3

3. Spines stiffly acicular

4

3. Spines tortuous 4. Ribs 10-12; spines to 15 mm 4. Ribs 15; spines to 25 mm 14.1. subsp. crispa Stem semiglobular, rarely globular, purplish blackish-brown; areoles 5 x 3 mm, 7 mm apart; spines all hair like, contorted, c. 16-24, not divisable into radials and centrals, 30-50 x 0.075 mm. Seeds 1.1 x 1.1 mm, asymmetric ratio 0.8; testa strongly ribbed. DISTRIBUTION. Chile, W of Freirina. SPECIMEN EXAMINED. CHILE. Huasco: Freirina, FK-78. Subsp. crispa is a very localized population within those of subsp. atroviridis, which surrounds it on all sides with intermediate populations between the two. Nomenclatural priority requires that E. crispa be used for the species name.

1 l.2d. var. totoralensis 14.2b. var. huascensis 14.2c. var. carrizalensis

FIGURE 1 2 E.

crispa

1 subsp. crispa FK-78, fl. 5 cm, stigma 5 mm. subsp. atroviridis var. huascensis

FK-fio FK-76,

П. 5 cm. var. carrizalensis

П

1.2

cm

FK-58.

Kattermann:

Eriosyce

Kattermann:

им

Eriosyce

1

v 1f if * 2> 4Ш 2

FIGURE 13 B.
var n

2

'

14.2. subsp. atroviridis (Ritter) Kattermann

» ' ' • » - ' - FK-526, c m

'

var. totoralensis

|

4 2 a

. var. atroviridis (Ritter) Kattermann

Stem often elongated; ribs 12-14, 1.2 cm high; areoles 7 x 6 r

FK-. i2

mm, 6 mm apart; spines stiffly or thickly needlelike, radials 12-14, 30 x 0.23 mm, centrals 3^4, 40-60 x 0.75 mm. DISTRIBUTION. Chile, from S to slightly N of Vallenar, then E to W of Freirina. SPECIMENS EXAMINED. CHILE. Huasco: S of Canto de Agua, FK-65; N of Freirina, FK-67, 69; S of Vallenar, FK-105; S of Freirina, FK-160; N of Vallenar, FK-397. 14.2b. var. huascensis (Ritter) Kattermann Stem subglobular; ribs 10-12 broader; spines shorter and fewer, mostly straight, radials 6-8, 5-12 mm, centra] 1, 15 mm. Seed not ribbed. DISTRIBUTION. Chile, E of Huasco. SPECIMENS EXAMINED. CHILE. Huasco: FK-76; N of Huasco/ FK-74.

Kattermann:

Eriosyce

14.2c. var. carrizalensis (Ritter) Kattermann Stem purplish black, ribs 15, 1 cm high; areoles not sunken, 6 x 4 mm, 12 mm apart; radial spines 10—13, 10—18 mm, centrals 4, 15-25 mm. DISTRIBUTION. Chile, c. Carrizal Bajo. SPECIMENS EXAMINED. CHILE. Huasco: E of Carrizal, FK-61; N of Carrizal Bajo, FK-813; Carrizal Bajo, FK-58.

PLATE 11 'page 70)

1 var. totoralensis (Ritter) Kattermann Stem small; ribs fewer; spines thin, sometimes contorted. Seed with a smoothly tuberculate testa. DISTRIBUTION. Chile, Copiapo, c. Totoral Bajo. SPECIMENS EXAMINED. CHILE. Copiapo: S of Totoral Bajo, FK-52, 150; N of Totoral Bajo, FK-526. 1

Merging into forms of var. carrizalensis end of its range.

1

2

3

4

5

6

E.

heinrichiana

towards the southern var. setosiflora 2-6

15. Eriosyce garaventae (Ritter) Kattermann Stem grass green to bluish green, globular to elongated, 6-12 cm in diam., apex spiny; ribs 11-16, deeply notched above and with a small chin below each areole; areoles 7.5—10 x 5 mm, 7.5-10 mm apart, with greyish wool; spines straw yellow, fairly thick, radials 12, centrals 1-6, 25—45 mm, curved upward; roots fibrous, from a short taproot. Flowers borne from young areoles, not fragrant, 3.5-5 x 4 cm, funnelform, yellowishgreenish; pericarpel grass green, bract-scales narrow, 2—4 mm, brownish, axils with small (loccose wool tufts and white, straight 5-15 mm bristles; tube covered as for pericarpel; perianth-segments 17.5-25 x 5-8 mm, 2-3 mm wide at base, pointed at apex, pale yellow, reddish toward base, sometimes with reddish midstripes; filaments greenish to white, 7.5-12.5 mm, resting against style, inserted to within 3 mm from top of tube; style 25 x 1-1.5 mm, pale greenish; ovary locule apex updrawn; stigma-lobes 10, yellowish, 4 mm, erect to spreading; nectary simple, 3—4 x 4 mm, white at base. Fruit elongating, 2.5 x 1.3 cm, hollow, seed free within, yellowish with pale red tint, dehiscent by a basal pore, remaining firmly attached for a long time; perianth remnant attachment area large, 65'# of fruit diam. Seed [SEM Plate VIII 1-14 x 1.12 mm, asymmetric ratio 0.5; testa black to dark brown, dull, finely tuberculate, not ribbed; hilum keyhole-shaped, 0.33 x 0.24 mm, not sunken; micropyle in wide groove, hilum rim only very little modified, not raised. DISTRIBUTION (MAP 8). Chile, known only from Cerro Campana (Prov. Valparaiso). SPECIMENS EXAMINED. CHILE. Valparaiso: loc. cit., FK-102, 103, 249.

E.

FK-11

crispa

2

subsp. crispa

3

var. huascensis

FK-78

4

var. carrizalensis

5

var. totoralensis

FK-52

6

var. totoralensis

FK-526

FK-76 FK-58

PLATE 12 'page 71)

1

E. aspillagae

FK-197

2

E. marksiana

KK-755

3

/•;. marksiana

FK-203

4

A',

marksiana

var. lissocarpa

FK-206

5

E. garaventae

FK-103

6

E. heinrichiana

FK-465

09

Kattermann:

F I G U R E 14 1

I

garaventae

2

/

aspillagae

3

E. marksiana

1-1

F K - 1 0 2 , fl. 3 . 5 cm. F K - 1 9 7 , П. 3 cm FK-755

Eriosyce

Kattermann:

Eriosyce

16. Eriosyce aspillagae iSohrens) Kattermann Stem dark green, at first flattened later more globular, 8-15 cm diam., the sunken apex partly covered by curved spines, tissues softly fleshy, freely caespitose forming clumps; ribs few, 5-8, later to 14, 1-3 cm wide, depressed or notched between the areoles, forming tubercles 12 x 10-25 mm, with impressed lines around; areoles round to oval, with white wool, later naked, on lower part of plant more linear and sunken, 3-4 mm, 8 mm apart; spines white tipped darker, later grey, radials 4-12, 10-20 mm, thin, needlelike, centrals 1-4, 20-30 mm, thicker, straight to curved, longer in older plants; rootstock a large napiform taproot. Flowers from young areoles, 4 cm, funnelform; pericarpel brownish green, 7 mm, bract-scales 2 mm, with plentiful white, loose wool and bristles from axils; tube short, with closely set bract-scales, these larger and more oval toward apex, wool and hristles as for pericarpel, bristles 15 mm; perianth-segments 20-30 x 8 mm, яра tula te, obtuse at apex, light silky yellow, with red midstripes, outer segments more reddish, inner segments silky yellowish green; filaments white; style thin, white, 30 x 1.3 mm, cream; ovary locule apex updrawn; stigma-lobes 4, 6 mm, spreading, cream-white; nectary simple. 2-3 mm. Fruit 1 cm, oval to elongated, reddish, hollow, fruit wall at first fleshy but soon dry, dehiscent by a partial circumscissile slit at base (somewhat as in suhsect. Pyrrhocactus), seed loose within loften partly germinated within the fruit); perianth remnant attachment area large, 69*# of fruit diam. Seed [SEM Plate VII | 1.14 x 1.12 mm, asymmetric ratio 0.3; testa dull black, tuberculate, not ribbed; hilum small, 0.36 x 0.16 mm; micropyle in groove, hilum rim slightly enlarged, not raised.

PLATE 13 (page 1

E.

74)

heinrichiana

subsp. simulans

FK-82

(above) 2

/-.'. crispa

var.

atroviridis

FK-65 (below)

PIRATE 14 (page

75)

1

2

DISTRIBUTION (MAP 8). Chile, Colchagua, Hacienda Tanume. SPECIMEN EXAMINED. CHILE. Colchagua: San Fernando, Hacienda Tanume, FK-197.

3

4

FK-755 from Cabo Carranza (Prov. Talca) may belong here on the basis of its fruit dehiscence and seed similarities. At present I place it with E. marksiana var. marksiana.

5

6

1-6 1

/•.'. subsp.

heinrichiana heinrichiana

FK-465 2

subsp.

heinrichiana

FK-177 3, 4

subsp. intermedia

FK-473

5

subsp. intermedia

FK-2S

6

subsp. intermedia

FK-458

73

Kattermann:

Eriosyce

17. Eriosyce marksiana (Ritter) Kattermann Stem flattened to subglobular, later elongated, grey green, 12-24 cm diam., apex sunken and naked; ribs 16-28, 1.5 cm high, sunken above and below areole, with a small to very large chin; areoles oval, 5-10 mm, with white wool, 10-15 mm apart; spines thick, grey brown with darker tips, radials 8-12, directed sideways to partly curved upward, 15-25 mm, centrals 1-6, thicker, strongly curved upward, not flattened; roots fibrous. Flowers many, from near stem apex, 3-4 cm long and wide; pericarpel light green, with very small yellowish bract-scales with white flocks of wool from their axils; tube funnelform to bell-shaped, 9-13 x 1-1.5 cm, covered as for the pericarpel, inside pale yellow, toward apex with straight bristles; perianth-segments 15-27.5 x 3-5 mm, narrower at base, apex lanceolate, margins not serrate, lemon yellow, sometimes reddish yellow; filaments pale yellow, reddish toward apex, 4-12.5 mm, inserted to within 1.5-3 mm of tube apex; style 20—25 x 1.9 mm, light yellow, thinner toward apex; ovary locule apex flat to updrawn; stigma-lobes 7-10.1.5-2 mm, erect to clasped together, light yellow; nectary simple, 2-4 x 4 mm. base honey-yellow, pale yellow towards apex, filled with nectar. Fruit 1.5 x 1 cm, brownish red or more greenish, covered as for pericarpel, fleshy, dehiscent by a basal pore, remaining firmly attached for a long time; perianth remnant attachment area wide, 86% of fruit diam. Seed |SEM Plate VIII 1.47 x 1.44 mm, asymmetric ratio 0.5; testa brown-blackish, smooth to finely tuberculate, not ribbed; hilum keyhole-shaped, small, 0.36 x 0.14 mm; micropyle in groove, hilum rim flat to regressed at micropyle (not raised), only very slightly enlarged. DISTRIBUTION (MAP 8). Chile, from Rancagua S to Rio Maule, E into the foothills of the Andes to 2000 m, W to the coast S of Constitucion. K E Y TO VARIETIES OK E . MARKSIANA

1. Spines thickly acicular; perianth-segments narrowly lanceolate, tips acute, 1.5-2.75 mm wide a. var. marksiana 1. Spines thinly acicular; peri a nth-segments almost spatulate, to 6 mm wide 2 2. Spines horn-coloured to grey 2. Spines yellow

b. var. lissocarpa c. var. gracilis

a. var. marksiana DISTRIBUTION. Chile, as for the species, to c. 1500 m. SPECIMENS EXAMINED. CHILE. Curicd: Villa Prat, FK-203. Talca: Botalcura, FK-277; Cabo Carranza, FK-755 (see note under E. aspillagae). b. var. lissocarpa (Ritter I Kattermann Differs from above: spines thinner; perianth-segments wider. DISTRIBUTION. Chile, Prov. Colchagua and Prov. Curico. SPECIMENS EXAMINED. CHILE. Curico: Rio Teno, 750 m, A. Hoffmann in FK-434, ibid., 1500 m, A. Hoffmann & R. Ferryman in FK-435; W of San Fernando, FK-762. Colchagua: Sierra Bella Vista, FK-206. c. var. gracilis (Ritter) Kattermann Stem smaller; spines yellow and longer. DISTRIBUTION. Chile, Cerro Horcon de Piedra and S to Cerro Nacimiento. SPECIMEN EXAMINED. CHILE. Melipilla: Horcdn de Piedra, 2000 m, A. Hoffmann s.n.

70

Kattermann:

Eriosyce

FIGURE 15 1

/.'. marksiana

v;ir. marksiana

F K - 2 0 3 , fl. 3 . 5 cm.

2

E. marksiana

var. lissocarpa

FK-206

3

E.

engleri

Left, seed of A Hoffmann

s.n. in F K - 5 6 7

C e n t r e a n d right, hilum and seed of F K - 5 6 8

77

Kattermann: 18. Eriosyce engleri (Ritter) Kattermann Stem large and slightly elongated, to > 30 x 14-18 cm, densely covered by spines; ribs numerous, 16-20, notched above the areoles, with a small chin below; areoles with white wool, 7.5-12.5 x 5-7 mm, 10-15 mm apart, later only 5 mm apart; spines curved upward, thick, the lower half of the spines white or yellowish, upper half brown to black, radials 12-20, shorter, thinner, and lighter coloured than the 5-8, 40—70 mm centrals; roots (ibrous. Flowers borne near stem apex, not fragrant, 5-6 x 4—4.5 cm, more funnelform than in E. curvispina: pericarpel green, with small, narrow, brownish red bract-scales, with flocks of wool from their axils; tube covered as for pericarpel, with a few, thin, white 20 mm bristles near apex, there 25 mm wide, funnelform; peri a nth-segments 25-30 x 3 mm at base, to 7-14 mm wide near midpoint, apex spalhulate, some segments ± linear, pale lemon yellow, with reddish to deep red midstripes Vt-Vi of width, abaxial surface of outer segments olive green; filaments pale greenish yellow, to reddish near apex, 15-20 mm. inserted to within 1-2 mm of tube apex; style 30—35 x 2-3 mm. pale greenish white, hollow above; ovary locule apex unknown; stigma-lobes exserted beyond anthers; nectary pale, 2-3 x 5-6 mm, partly closed by the filaments. Fruit reddish, covered as for the pericarpel, perianth remnant persisting (Ritter mentions a wide attachment for the perianth remnant). Seed |SEM Plate Villi 1.65 x 1.65 mm, asymmetric ratio 0.6; testa dull black, finely tuberculate, ribbed, similar to seed of E. marksiana var. lissocarpa and E. garaventae; hilum small, micropyle in a ± regressed portion of the hilum.

Eriosyce The picture of the sectioned flower (from R. Ferryman) places this plant close to E. curvi­ spina var. curvispina. E. marksiana var. gracilis, from Horcon de Piedra, has seeds almost exactly like E. engleri. Until flowers can be more closely observed E. engleri is treated here as a separate species.

DISTRIBUTION (MAP 8). Known only from the type locality, Cerro Robles Alto/Laguna Chicauma, on high mountains between Santiago and Valparaiso. SPECIMENS EXAMINED. CHILE. Chacabuco:

Laguna Chicauma, Л. Hoffmann s.n., cult. R. Ferryman (photo.); A. Hoffmann s.n. (anno 1985) in FK-567. M A P 8. Distribution of E. garaventae, E. marksiana a n d E. engleri.

78

E.

aspillagae.

Kattermann:

Eriosyce

19. Eriosyce heinrichiana (Backeberg) Kattermann Stem subglohose to globular or elongated, dark brownish to blackish green, 4—10 cm diam.; ribs 14—22, depressed between the areoles to form tubercles, 5-10 mm high, obtuse with a long drawn out chin below each areole; areoles sometimes slightly sunken, to 20 mm apart; spines straight to curved, stiffly acicular, radials 6-10, 3-12 mm, centrals 1-4, 3-15 mm, spines sometimes absent; rootstock of large tuberous taproots, slightly narrowed to a neck where joining plant stem. Flowers from young areoles, 3-5 cm long and wide; pericarpel and tube bract-scale axils with often long, floccose tufts of wool and bristles, tube base slender, narrow funnelform, upper part of tube broad funnelform, trumpet-shaped, with an often very much elongated pericarpel; style 25-30 mm, cream-yellow to reddish; ovary locule apex downdrawn to updrawn; stigma-lobes 8-12, 5—8 mm, clasped together or erect, reddish to orange; nectary simple, 3—5 mm. Fruit oval, thin-walled, fleshy, seeds free within, elongating 1.5-3x upon ripening, 1-1.5 cm diam., red, dehiscent by a basal pore, remaining firmly attached to plant; perianth remnant attachment area small, 46% of fruit diam. Seed [SEM Plate Villi 0.75 x 0.83 mm. asymmetric ratio 0.8; testa brown, tuberculate, sometimes ribbed, keeled; hilum keyhole type, 0.36 x 0.23 mm, almost flat; micropyle in enlarged section of the raised hilum. DISTRIBUTION (MAP 9). Chile, 28-31°S, from S of Huasco to Quebrada Las Palmas. At the N and S end plants grow only along the coast. In the Choros Valley populations almost reach Tres Cruces; in the Elqui Valley populations reach Herradura. E. heinrichiana is closely related to E. limariensis and south of Vicuna subsp. intermedia merges with some populations of that species. KEY TO INFRASPECIFIC TAXA OF E. HEINRICHIANA

1. Stigma-lobes 5-7 mm, clasped together 2 1. Stigma-lobes 4 mm, erect to spreading; (stem elongated, dark green; flowers funnelform, often with dense wool on pericarpel and tube) 19.3. subsp. simulans 2. Ovary locule updrawn

19.2b. var. setosiflora

2. Ovary locule downdrawn 3 3. Stem brown, rarely elongated 19.1. subsp. heinrichiana 3. Stem often green, or purplish grey in the populations in the middle Elqui valley 19.2a. subsp. & var. intermedia 19.1. E. heinrichiana subsp. heinrichiana Stem subglobose to only sometimes globular, dark brownish to blackish green, 4-8 cm diam.; rihs 14-22, tubercles 5 mm high. Bristles not always present on the pericarpel, not porrect. DISTRIBUTION. Chile, 28°30-29°30'S, from S of Huasco to El Tofo. At the northern end of the distribution plants grow only along the coast. SPECIMENS EXAMINED. CHILE. Elqui: N of Trapiche. FK-401; S of Trapiche. FK-81; E of Mina El Tofo, FK-176, 177; E of Choros Bajos, FK-470; W of Tres Cruces, FK-537; Huasco: Sarco, Quebrada Honda, FK-465.

79

Kattermann:

Eriosyce

Kattermann:

\

it

fl\\

Eriosyce

1Q

v f t '

/

1 4'

til ^

j

1 /

\ 2

F I G U R E 17 Габоне E.

heinrichiana

1

var. setosiflora

FK-11,

fl. 5 cm. 2

subsp. nimulans

FK-82,

fl. 3 . 5 cm, s t i g m a 4 m m . F I G U R E 1 6 (left) 1

E.

heinrichiana

s u b s p . heinrichiana

FK-465,

П. 4 cm, s t i g m a 6 m m . 2

s u b s p intermedia

FK-473

3

subsp. intermedia

FK-458,

4

subsp. intermedia

fl. 4 cm, s t i g m a 8 m m . FK-28,

fl. 4 . 5 c m , s t i g m a 7 m m .

81

Kattermann:

Eriosyce

19.2. E. heinrichiana subsp. intermedia (Ritter) Kattermann 19.2a. var. intermedia (Ritter) Kattermann Stem sometimes more elongated, dark green to sometimes purplish-blackish; spines often more curved upward. DISTRIBUTION. Chile, throughout the Elqui Valley until Rivadavia; along the coast populations extend N to Cruz Grande and S to Quebrada Las Palmas. SPECIMENS EXAMINED. CHILE. Elqui: W of Vicuna, Las Rojas, FK-28; Cuesta Buenos Aires, FK-104; S of Cruz Grande, FK-180; Rivadavia, FK-458; El Tambo, FK-461; Herradura, FK-476; near Pafiuelas, FK-534; Panuelas, FK-473. 19.2b. var. setosi flora (Ritter) Kattermann Ovary locule apex updrawn. DISTRIBUTION. Chile, coastal region of the Elqui and Limari valleys. SPECIMENS EXAMINED. CHILE. Elqui: Punta Lengua de Vaca, FK-23; Quehrada Las Palmas, S of Coquimbo, FK-11. Limari: S of Soco, FK-448. 19.3. E. heinrichiana subsp. simulans (Ritter) Kattermann Stem elongated, 7-15 x 5-7 cm, dark green; ribs 13-16, 7 mm high; areoles 8 x 5 mm, 5 mm apart, with long white wool, wool covering stem apex. Flowers not narrow funnelform at base of tube; pericarpel and tube covered by dense white wool and with many bristles from bract-scale axils; ovary locule apex updrawn; stigma-lobes shorter, 4 mm, reddish to cream, erect to spreading. DISTRIBUTION. Chile, throughout the lower Rio Choros valley. SPECIMENS EXAMINED. CHILE. Elqui: W of Trapiche, FK-82; E of Los Choros, FK-472. 20. Eriosyce napina (Philippi) Kattermann Stem miniature, 2-4 cm diam., sometimes elongated; ribs dissolved into tubercles only a few mm high, 2-5 mm diam., with obtuse chin like protrusion below areoles; areoles sunken, with white wool, stem apex naked or sometimes covered by wool; spines few to many, 2-5 mm, black to horn-coloured. + appressed, radiating or sometimes ± pectinate, mostly all radials, but sometimes 1 central present; rootstock of large, tuberous taproots with a neck like restriction between root and stem. Flower from young areoles, funnelform, pale yellow to pale reddish; pericarpel and tube bract-scale axils with long, brownish to whitish floccose wool and black to brownish or whitish bristles; style 15-20 mm; ovary locule apex updrawn; stigma-lobes 7-10, 3—4 mm, erect to slightly spreading; nectary simple, tubular. 2-3 mm. Fruit elongated, seed restrained in sac­ like structure, dehiscent by a basal pore, firmly attached; perianth remnant attachment area 25-60% of fruit diam. Seed [SEM Plate X] 0.95-1.21 x 0.85-1.12 mm, asymmetric ratio 0.6; testa brown to black, tuberculate, ribbed; hilum keyhole-shaped, flat, 0.47-0.59 x 0.24-0.33 mm; micropyle completely or almost completely surrounded by hilum rim enlargement, rim not raised. DISTRIBUTION (MAP 9). Chile, in the Rio Huasco valley from Huasco to Vallenar. KEY TO SUBSPECIES OF E. NAPINA

1. Tubercles large, 5-6 mm diam.; stem 3-5 cm diam., never elongated; spines black, 20.1. subsp. napina appressed 1. Tubercles small, 1-2 mm diam.; stem smaller, 1-3 cm diam., often elongated; spines horncoloured (subsp. lembekei) 2 2. Stem globular, rarely elongating; tubercles 2-3 mm diam. 20.2b. var. duripulpa 2. Stem globular to elongated; tubercles 1-2 mm diam. . . . 20.2a. subsp. & var. lembekei

82

Kattermann:

Eriosyce

20.1. E. napina subsp. napina Stem Ыаск-grecn; ribs divided into tubercles 5-6 mm diam. Flower yellow to reddish; peri­ carpel and tube with dark brown to pale brown wool and brown bristles. Seed small, 0.95 x 0.85 mm, asymmetric ratio 0.6; hilum 0.47 x 0.24 mm. DISTRIBUTION. Chile, in the Rio Huasco valley from Huasco to Freirina, and extending C. 15 km S of Freirina. SPECIMENS

EXAMINED.

CHILE.

Huasco:

Huasco, с. 1 km from beach, FK-75; near Freirina, FK-70; S of Freirina, FK-158, 159, 162. 20.2. E. napina subsp. lembekei FCattermann 20.2a. var. lembekei Kattermann Stem 1-2 cm diam., sometimes elongated; ribs in greater number, tubercles much smaller, 1-2 mm diam.; spines horn-coloured, radi­ ating to pectinate. Flower yellow. Perianth remnant attachment area 23'Й of fruit diam. Seed 1.00 x 1.00 mm, asymmetric ratio 0.6; hilum 0.53 x 0.33 mm. DISTRIBUTION. Chile, known only from the type locality W of Freirina. SPECIMEN EXAMINED. CHILE. Huasco: loc.

cit., FK-77 (holotype). According to Pablo Weisser, who was shown the plant by Lembcke, the type locality of Neochilenia lembekei Backeberg (invalid name) was west of Freirina (cf. FK-77).

M A P 9. Distribution of E. heinrichiana. and E. odieri

E.

napina

P L A T E 1 5 (page 84)

20.2b. var. duripulpa (Ritter) Kattermann Stem rarely elongating, 2-3 cm diam. Seed 1.21 x 1.12 mm high, asymmetric ratio 0.6; hilum larger, 0.59 x 0.30 mm. DISTRIBUTION. Chile, from Vallenar to Huasco.

1

2

3

4

5

6

SPECIMENS EXAMINED. CHILE. Huasco: N

of Huasco, FK-72; N of Maitencillo, FK-80; Vallenar, FK-398.

1-2

/•.'. recondita

subsp.

iquiquensis

1, F K - 3 3 8 a ; 2, F K - 3 6 6

Ritter illustrates plants which are much elongated, but my observations indicate that elongation happens only rarely in habitat, although it may occur in cultivation.

3

E. recondita

4

E. napina

subsp. napina

subsp. recondita

5

E. napina

subsp. lembekei

6

E. napina

subsp. napina

FK-369

FK-75 FK-77 FK-70

83

Kattermann:

Eriosyce

If

• ^ 1

к 0

V \1 1 ч

1

^

1С

2

w% P L A T E 16 (ред.- 8 5 ; 1 - 4 £ . napina

var.

F I G U R E 18 duripulpa

1

6

£ . « № r i Gay 4 2 0 0 £ . odieri

Jung s.n.

2

subsp. lembekei

3

var. duripulpa

napina

1

subsp. odieri

F K - 7 7 , П. 3 . 5 cm. F K - 3 9 8 , fl. 3 cm.

F I G U R E 1 9 (right)

88

E.

F K - 7 5 , П. 4 c m ,

s t i g m a 3—4 m m .

1. F K - 7 2 ; 2 , F K - 8 0 ; 3 - 4 , F K - 3 9 8 5

(above}

subsp. napina

E.

odieri

F K - 8 0 4 , fl, 3 . 5 c m .

2

var. monte-amargeneia

3

subsp. glabrencenH F K - 5 3

4

subsp. fulva

FK-429

FK-806

Kattermann:

Eriosyce

87

Kattermann:

1

2

3

4

5

6

1-4

1-2

var. monte-amargensis FK-806 subsp. FK-53

glabrescens

subsp. fulca E. occulta I

FK-529

21. Eriosyce odieri (Salm-Dyck) Kattermann Stem subglobular, rarely globular, 3-6 cm diam., brownish to almost greyish; ribs completely dissolved into tubercles arranged ± spirally, tubercles 4—6 mm diam.; areoles sunken, with a chin below; spines mostly radial, ± appressed, 6-12, thin to thick, 2-5 mm, when centrals present these to 10 mm, much thicker; roots short to long turbinate. Flowers borne from young areoles, 3—4 cm, funnelform, pale yellow to pale reddish; pericarpel and tube bract-scale axils with white floccose wool, axils of tube with bristles; style 20 x 1.8 mm, reddish; stigmalobes 6-8, 4—6 mm, spreading, pale red, lighter than style; ovary locule apex updrawn; nectary modified, widened at base, 3 mm. Fruit elongating 3—4x diam. upon ripening, seed restrained in sac-like structure in fruit apex, dehiscent by basal pore, attachment firm; perianth remnant attachment area 35% of fruit diam. Seed |SEM Plate DC] 0.90-1.12 x 0.71-1.00 mm, asymmetric ratio 0.4-0.5; testa brown, tuber­ culate ribbed; hilum keyhole-shaped, 0.30 x 0.21 mm; micro­ pyle in groove partly surrounded by enlarged, flat hilum rim. DISTRIBUTION (MAP 9). Chile, from Morro Copiapo E to c. railway station of Monte Amargo, S to Carrizal Bajo.

FK-391

enmeraldana

Eriosyce

FK-436

KEY TO INFRASPECIFIC TAXA OF E. ODIERI

1. Stem subglobular, 3-5 cm diam.; ribs 18-21; spines radiating, not appressed (subsp. odieri) 2 1. Stem smaller, 2-3 cm diam.; spines appressed 3 2. Spines only radial 2. Central spines present 3. Tubercles 3-5 mm diam 3. Tubercles smaller and more numerous, 2-3 mm diam

21.1a. subsp. & var. odieri 21.1b. var. monte-amargensis 21.2. subsp. glabrescens 21.3. subsp. fulva

21.1. E. odieri subsp. odieri 21.1a. var. odieri Stem 3—5 cm diam.; ribs 18—21. tubercles 3 mm diam., 2-3 mm high; areoles 1.5 x 0.5 mm. 1.5 mm apart; radial spines 8-10, 1.5-2 x 0.05 mm, appressed. DISTRIBUTION. Chile, known only from Morro Copiapo S of Caldera. SPECIMENS EXAMINED. CHILE. Copiapo: top of Morro Copiapo, FK-804; loc. cit., base, FK-802 Ineotype); Morro Copiapd, Gay 4200, Jung s.n. 21.1b. var. monte-amargensis Kattermann Differs from the above: spines often thicker and longer; central 1, porrect; flower more reddish. DISTRIBUTION. Chile, Monte Amargo railway station E to c. Caldera and c. 10 km S. SPECIMENS EXAMINED. CHILE. Copiapu, SW of Monte Amargo, FK-806; S of Monte Amargo. FK-522 (holotype).

89

Kattermann:

Erioayce

21.2. E. odieri subsp. glabrescens (Ritter) Kattermann Stem sometimes almost spineless, soft; ribs 7-8, dissolved into tubercles 8-6 mm diam.; roots longer, narrowed to a neck above; pericarpel and tube with white dense wool; nectary modified. DISTRIBUTION. Chile, border region of Prov. Copiapo and Huasco. SPECIMENS EXAMINED; CHILE. Copiapo: N of Totoral, FK-525; S of Totoral, FK-53, 144. Huasco: S of Carrizal Bajo. FK-815. 21.3. E. odieri subsp. fulva (Ritter) Kattermann Spines thicker and longer, appressed to ± recurved; roots longer, narrowed to a neck above. DISTRIBUTION. Chile, known only from E of Totoral. SPECIMEN EXAMINED. CHILE. Copiapo: loc. cit., FK-529. 22. Eriosyce taltalensis (Hutchison) Kattermann Stem subglobular to globular to elongated, 3-14 cm diam.; ribs 8-13. notched into tubercles with a chin below each areole, 1 cm high; areoles sunken, variahle in size; spines irregular, straight to strongly curved, dark brown to blackish, radials 6-12, 3-20 mm, centrals 6-12, 20-40 mm; rootstock a shortly conical turbiniform taproot. Flowers borne from young areoles, narrow funnelform or sometimes more broadly funnelform. 3-4 x 2-3 cm; pericarpel and tube bract-scale axils with white floccose wool and white to brown, tortuous to straight bristles; perianthsegments lanceolate, fuchsia red to pale yellow; style 15-20 mm, cream-yellow to deep purplish red; ovary locule apex unknown; stigma-lobes 3-5 mm, erect or clasped together, cream-yellow to dark purplish red; nectary widened at base, 1-2 mm. Fruit elongating, seed loose within, dehiscent by a basal pore, fruit attachment firm; perianth remnant attachment area small, 35-45% of fruit diam. Seed ISEM Plates XI, XIII 1.0 x 0.9-1.0 mm, asymmetric ratio 0.5-0.7; testa tuberculate, sometimes strongly ribbed; hilum keyhole-shaped, 0.3 x 0.21 mm; micropyle in narrow part of keyhole, hilum rim enlarged, not raised. DISTRIBUTION (MAP 10). Chile, from S of Antofagasta to Caldera (Prov. Copiapo). Populations are mostly near the coast or on coastal hills. KEY

TO INFRASPECIEIC TAXA OF E. TALTALENSIS

1. Perianth-segments concolorous

3

1. Perianth-segments white below, uppermost one-third fuchsia red (subsp. echinus)

2

2. Stem apex with short areolar wool

22.3a. subsp. & var. echinus

2. Stem apex with long floccose areolar wool 22.3b. var. floccosa 3. Stem 6-15 cm diam., often elongated, glaucous green to green or brownish; flowers variable, yellow to reddish 22.2. subsp. paucicostata 3. Stem 3-8 cm diam., mostly subglobular to globular; flowers fuchsia red to white 4 4. Spines straight; flowers red, pink or white, style 1.2-1.5 mm diam. (subsp. taltalensis) 4. Spines tortuous, flower white, style 2-2.5 mm diam 5. Flowers red or cream 5. Flowers white or pink

90

.5

22.4. subsp. pilispina 22.1a. subsp. & var. taltalensis 22.1b. var. pygmaea

Kattermann:

Eriosyce

22.1. E. taltalensis subsp. taltalensis 22.1a var. taltalensis Stem subglobular to globular; central spines mostly straight, ± erect. Flower fuchsia red or cream; style purplish red. DISTRIBUTION. Chile, Prov. Antofagasta, S almost to Chanaral, N to Taltal (Paposo). SPECIMENS EXAMINED. CHILE. Antofagasta: Paposo, A. Hoffmann 194 in FK-440, E. Gay 4183; E of Paposo, FK-108; Quebrada Matancillas, Taylor s.n. (K, photos. I. Taltal, Breas, FK-116; Quebrada de Guanillos, FK-511; Chanaral: N of Pan de Azucar, FK-792; S of ibid., FK-798. The flowers of var. taltalensis are mostly carmine red, or sometimes cream. This variation can sometimes be observed in the same population. There exists a close relationship with E. confinis, which adjoins at the southern end, W of Caldera. 22.1b. var. pygmaea (Ritter) Kattermann Perianth-segments mostly white. DISTRIBUTION. Chile, mountains N of Chanaral to S of Caldera. SPECIMENS EXAMINED. CHILE. Chanaral: N of Chanaral, near sea level, FK-45, 130, 800, Gay 4192; ibid., mountains, FK-132. Copiapo: N of Caldera, FK-771, 765, Gay 4180; Morro Copiapo, FK-803. 22.2. E. taltalensis subsp. paucicostata (Ritter) Kattermann Stem larger and almost always elongated, 6-15 cm diam., dark green to glaucous green; ribs less depressed between areoles; spines mostly curved upward. Perianth-segments yellow to whitish, sometimes with a faint pinkish midstripes; tube and pericarpel bristles brown to white. Seed 0.95 x 0.90 mm, asymmetric ratio 0.8; testa tuberculate, not ribbed; hilum 0.35 x 0.21 mm; micropyle in narrow groove, hilumrimenlarged at micropyle, not raised. D I S T R I B U T I O N . Chile, Prov. Antofagasta, from Taltal N to c. 30 km N of Paposo. SPECIMENS EXAMINED. CHILE. Antofagasta: loc. cit., FK-34, 109, 110, 385, 386, 387, 388, 389, 489, 496, 499, 501, 785. 22.3. E. taltalensis subsp. echinus (Ritter) Kattermann 22.3a. var. echinus (Ritter) Kattermann Spines often thick. Flowers red with whitish centre. DISTRIBUTION. Chile, from S of Antofagasta along the coast to Blanco Encalada (Prov. Antofagasta). SPECIMENS EXAMINED. CHILE. Antofagasta: near El Cobre, FK-373, 374, 778, 780.

PLATE 18 (page 92)

/•.'.

1

2

3

4

5

6

taltalensis

1

var taltalensis

FK-116

2

var. taltalensis

Gay 4192

3

var. pygmaea

4

var. paucicostata

Gay 4180 FK-110

5

var. paucicostata

FK-389

6

var. paucicostata

FK--199

PLATE 19 (page 93)

1

2

3

4

5

6

1

E. taltalensis

2

A',

Taylor

var. floccosa 3

/'•-

4

E. sociabilis

FK-781

taltalensis

subsp. pilispina

5

/•:'.

FK-772

FK-147

taltalensis

var. pygmaea 6

s.n.

taltalensis

E. laui

FK-765

Lau 1541

91

Kattermann:

Eriosyce

22.3b. var. floccosa (Ritter) Kattermann Areoles with long wool, sometimes covering apex. DISTRIBUTION. Chile, known only from the region of Blanco Encalada, Prov. Antofagasta. SPECIMENS EXAMINED. CHILE. Antofa­ gasta: Blanco Encalada, FK-380, 381, 781. 22.4. subsp. pilispina I Ritter) Kattermann Spines thin, contorted. Flower-tube broadly funnelform; stigma-lobes 2-3 mm, spreading; nectary tubular, widened at base, 2 mm. Fruit much elongated; perianth remnant attach­ ment area 31% of fruit diam. Seed 0.90 x 0.86 mm, asymmetric ratio 0.6; testa ribbed; hilum 0.40 x 0.23 mm; micropyle in wide groove, hilum rim enlarged at micropyle, not raised. DISTRIBUTION. Chile, S of Barquito. SPECIMENS EXAMINED. CHILE. Chanaral: S of Barquito, FK-515; near Caleta Flamenco, FK-772.

FIGURE 20 1

E. taltalensis

var. pygmaea

FK-765.

Infraspecific categories_ of E taltalensis:

'.

F I G U R E 21 E.

•

subsp. taltalensis var. taltalensis

•

subsp. taltalensis var.

•

subsp echinus var. echinus

1

taltalensis

subsp. taltalensis

F K - 1 1 6 , П. 3 c m ,

stigma 3 m m . pygmaea

A subsp echinus var.

floccosa

О subsp. pilispina

f

\

2

3 О

subsp.

|

V,

occulta.

91

10. Distribution

F K - 1 1 0 , fl. 4 c m .

subsp. echinus

var. floccosa

FK-380,

/

"

fl. 3 . 2 cm.

paucicostata^ 4

MAP

»мЫ\>. paucicostata stigma 4.5 m m .

of E.

taltalensis

subsp. pilispina

F K - 7 7 2 , fl. 2 . 5 cm.

stigma 2.5 m m .

a n d E. 5

var. pygmaea

Cay 4 1 8 0

Kattermann:

Eriosyce

Kattermann:

FIGURE 22 1

E. occulta

2

E. esmeraldana

F'K-391, II 4 cm, s t i g m a 4 m m .

3

E. aerocarpa

FK-437 F K - 6 3 , Л. 3 . 5 cm.

Eriosyce

Kattermann:

Eriosyce

23. Eriosyce occulta Kattermann Stem semiglobular (to globular in cultivation), sometimes branching with new heads formed from upper part of rootstock, 3-Я cm diam.; ribs to 14, notched into rhombic tubercles, with chin below, 5 mm high; areoles 3-6 x 1.5 mm, 5 mm apart, sunken; spines sometimes lacking, black to dark brown, slightly curved, somewhat appressed. radials 4-6, 3-11 x 0.12 mm, centrals 0-1, 5-10 x 0.25 mm; rootstock large turbinate, not narrowed towards apex. Flower from young areoles, funnelform, 2.3—4 cm; pericarpel and tube bract-scale axils with white floccose wool and white contorted bristles; tube 13 mm; perianth-segments 18-20 x 5-6 mm, whitish with pale reddish midstripes; style 20 x 2 mm, purplish red at base, lighter above; ovary locule apex updrawn; stigma-lobes 8-10, 3—4 mm, erect or clasped together, orange-red; nectary 2 mm, widened at base. Fruit red, elongating 1.5-2x upon ripening, wall medium thick, fleshy, seed contained in a sac, dehiscent by a basal pore, attachment firm; perianth remnant attachment area 50% of fruit diam. Seed [SEM Plate XIIIJ 1.2 x 1.12 mm, asymmetric ratio 0.6; testa hlackish, deeply ribbed, slightly keeled; hilum 0.36 x 0.24 mm, slightly sunken; micropyle in groove of enlarged hilum rim. DISTRIBUTION (MAP 10). Chile, known from region of Taltal. SPECIMEN EXAMINED. CHILE. Antofagasta: Taltal, Breas, FK-39I (holotype). Note: This is Echinocactus occultus, Neoporteria occulta etc. of authors, following the use of the name employed by Schumann (Sbhrens coll.), and not Echinocactus occultus Philippi. which may be an older name for Eriosyce heinrichiana. Sect. II. Neoporteria subsect. 2. Chileosyce (spp. nos. 24-28) Stem very small, 2-3 cm diam.; ribs dissolved into tubercles; areoles small, with short wool; spines few, 5-8, short, 1-3 mm; roots long turbiniform, sometimes with delicate, easily damaged root epidermis. Flowers from young areoles. yellow to reddish; pericarpel and tube covered by dense wool, with porrect, stiff bristle clusters; stigma-lobes spreading; nectary modified, widened and angular at base. Fruit oval, thin-walled, dry shortly after ripening, with long porrect bristles from scale axils; dehiscent by a basal pore, wind dispersed; attachment area of perianth remnants medium sized. Seed small. 0.6-0.9 mm, often shiny black; hilum large, wide keyholeshaped, sunken; micropyle partly surrounded by modified hilum rim. DISTRIBUTION (MAP 11). On the coast of Chile, c. 25°45'-29 30'S. The northern populations are found from near sea level to 300 m, but toward the southern end of the range they can be found further inland at elevations of 300-600 m. This distribution matches that of subsect. Eriosyce and subsect. Neoporteria. Plants of these subsections often inhabit the same areas. n

Diagnostic bristles.

feature: Pericarpel and tube with dense wool and many porrect, brownish

24. Eriosyce esmcraldana (Ritter) Kattermann Stem 3—4 cm diam., dark green to red brown, sometimes branching by forming plantlets from outer areoles; ribs 13-22, dissolved into tubercles 2-5 mm diam. and 2-3 mm high; areoles sunken, 1-2 x 1 mm, 3-4 mm apart; spines thin, needlelike, radials 5-7, 3-5 x 0.125 mm; rootstock a short to long conical tap root. Flowers borne from young areoles, 2-3 cm long and wide; pericarpel and tube bract-scale axils with long, black bristles; nectary widened at base. Fruit clavate. Seed |SEM Plate XIII] 0.85 x 0.85 mm, asymmetric ratio 0.6; testa blackish to brownish, deeply ribbed; hilum 0.30 x 0.16 mm; micropyle in groove, hilum rim enlarged and raised at micropyle. DISTRIBUTION (MAP 11). Chile, along the coast from N of Planta Esmeralda to N of Pan de Azucar, Prov. Antofagasta.

97

Kattermann:

Eriosyce

SPECIMENS EXAMINED. CHILE. Antofa­ gasta: Quebrada de Guanillos, A. Hoffmann 132 in FK-437, FK-506; Cerro Morado, FK-795; Planta Esmeralda, A. Hoffmann 207 in FK-436. FK-504.

71

/t

The more porrect bristles on the fruit and flower of E, esmeraldana indicate its correct placement in subsect. Chileosyce. 25. Eriosyce aerocarpa (Ritter I Kattermann Stem subglobose to elongated, 2-3 cm diam., not branching; ribs many, to 24, dissolved into tubercles, these 1.5-4 mm diam. and 2-4 mm high; areoles not sunken, 3 x 1.5 mm, 2-4 mm apart, woolly, wool covering stem apex; spines short acicular, red-brown, straight to recurved, radials 6-14, somewhat pectinate, 2—3 mm, centrals 0—2, much thicker, 10 mm, porrect; rootstock a turbiniform taproot with a neck at apex. Flowers born from young areoles, funnelform, 3-5 cm; pericarpel and tube bract-scale axils with long floccose wool and many porrect bristles; perianth-segments 5-7 mm wide, pale reddish with darker midstripes; style 18—25 x 2 mm, reddish; ovary locule apex updrawn; stigma-lobes 11-14, 3-8 mm. spreading, reddish; nectary simple, not widened at base, 3 mm. Fruit elongating, bract-scale axils with long floccose wool and many porrect, brown bristles (wind-dispersed fruit), thin-walled, hidden by wool and bristles, dry when ripe, dehiscent by a basal pore, seed contained within sac, fruit attach­ ment loose (removed by wind); perianth remnant attachment area wide. Seed [SEM Plate XIIII 1.12 x 1.07 mm, asymmetric ratio 0.6; testa tuberculate, brownish; hilum narrow keyhole-shape, not deeply sunken, 0.40 x 0.25 mm; micropyle at end of narrow groove, hilum rim enlarged at micropyle, not raised. DISTRIBUTION (MAP 11). Chile, from E of Carrizal Bajo to Canto del Agua, Huasco. SPECIMENS EXAMINED. CHILE. Huasco; Canto del Agua, FK-63, 153.

69

<» Tocop на A

An

tofagast.

1

}

25° Talla

t I

•

I

L

С

i

F

i

1 1

Г

I

у

1

If iJ

.

о Copiapo

1

Г

\ Huasco J

?9

\

J

f \ Г 1

T

1

;

E Е.

aerocarpa

E. Е.

tenehrica tenebhca

E. Е.

MAP

11.

Chileosyce.

Distribution

of

Eriosyce

subsect.

La Serena Y

• • 0

E. Е.

esmeraldana krausii

E. laui Е. _i t

Kattermann:

Eriosyce

E. aerocarpa is placed in subsect. Chileosyce because of the mode of fruit dehiscence. Seed characters point to a closer relationship with E. napina (subsect. Horridocactus). Further observations are needed to determine this relationship more clearly. 26. Eriosyce krausii (Ritter) Kattermann Stem subglobular to flattened, 2-4 cm diam., grey-green to reddish brown, sometimes branching from the root-stem neck, offsets then forming their own roots; ribs 12-14, dissolved into tubercles, these 1.5-3 mm diam. and 2-3 mm high; areoles sunken, 2 x 1 mm, 2-3 mm apart; spines very thin, needJelike, straight to slightly curved, radiating to erect, radials 3—8, 1-2 mm, centrals 0—1, 2 mm; rootstock a long tapering turbiniform taproot, without neck, epidermis easily damaged. Flowers born from young areoles, funnelform, 3-4 cm; pericarpel and tube bract-scale axils with long floccose wool and many porrect bristles; perianth-segments 5-6 mm wide, yellow, sometimes with pale reddish midstripes; style 20 x 1.8 mm, pale yellow to reddish; ovary locule apex updrawn; stigma-lobes 7-12, 3—4 mm, spreading, yellowish; nectary tubular, widened at base, 3 mm. Fruit elongating, bract-scale axils with long floccose wool and many porrect brown bristles (wind-dispersed fruit), thin-walled, epidermis completely hidden by wool and bristles, seed loose within, dry when ripe, dehiscent by a basal pore, fruit attachment loose (removed by wind); perianth remnant attachment area 50% of fruit diam., transition from fruit to perianth remnant hidden underneath wool and bristles. Seed [SEM Plate XII] very small, 0,83 x 0.75 mm, asymmetric ratio 0.5; testa shiny black, cells more or less elongated and often not domed but rather flattened, not ribbed; hilum wide keyhole-shaped, 0.23 x 0.21 mm, deeply sunken; micropyle at narrow end of hilum, hilum rim barely enlarged and not raised at micropyle. DISTRIBUTION (MAP 11>. Chile, from N of Caldera to Cifuncho (Prov. Antofagasta). S P E C I M E N S E X A M I N E D . CHILE. Antofagasta: N of Pan de Azucar, FK-790. Chanaral: Pan de Azucar, FK-127; S of Barquito, FK-328, 392, 395, 773. Copiapo: Caleta Obispo, FK-135; N of Caldera, FK-438, A. Hoffmann & R. Ferryman (seed), cult. FK-845, loc. cit., FK-519. 27. Eriosyce tenehrica (Ritter) Kattermann Stem subglobular to flattened, 2-4 cm diam., not branching, blackish brown; ribs 12, dissolved into tubercles, these 3—6 mm diam. and 2-3 mm high; areoles not sunken, 4 x 1.5-2 mm, 3—5 mm apart; spines all radial, 3-6, 1-2 mm, thin; rootstock a long turbiniform taproot, sometimes with a neck. Flowers born from young areoles, funnelform, 3-5 cm; pericarpel and tube bractscale axils with long Ooccose wool and many porrect bristles; perianth-segments 5—6 mm wide, yellow to pale reddish brown; style c. 20 mm, cream-yellow; stigma-lobes 8-14, 3-8 mm, spreading, cream-yellow; nectary tubular, not widened at base, 3 mm. Fruit elongating, bractscale axils with long floccose wool and many porrect brown bristles (wind-dispersed fruit), thinwalled, epidermis completely hidden by wool and bristles, dry when ripe, dehiscent by a basal pore, seed contained within sac, fruit attachment loose (removed by wind); perianth remnant attachment area 40% of fruit diam., transition from fruit to perianth hidden by wool and bristles. Seed |SEM PI. XIII very small, 0.81 x 0.94 mm, asymmetric ratio 0.7; testa dull black, not ribbed, testa-cells high domed; hilum wide keyhole-shaped, 0.4 x 0.33 mm, deeply sunken; micropyle at narrow end of hilum, not groove like, hilum rim barely enlarged and not raised at micropyle. DISTRIBUTION (MAP l l ) . Chile, from west of Domeyko to the Rio Choros valley. S P E C I M E N S E X A M I N E D . CHILE. Huasco: W of Domeyko, FK-531. Elqui: Trapiche, FK-402. 28. Eriosyce laui Luthy: see Appendix II (p. 121). Chile, known only from the type locality south of Tocopilla. CHILE. Tocopilla: S of Tocopilla, FK-774.

DISTRIBUTION (MAP I D . SPECIMENS EXAMINED.

99

Kattermann;

Eriosyce

:

* *

•У/ 2

If

Iff

la

3

V-^>

F I G U R E 2 3 labove)

1 Ч Л Т Е 2 0 <pa*e

101)

1

E. krauaii

FK-438

1

E. aerocarpa

2

E. krauaii

FK-135

2

E. krauaii Hoffmann

3

E. krauaii

4

£.

5

Я . chilenaia FK-3 (bottom left)

6

E. chilenaia

.'] E. tenehrica

100

FK-402

F K - 6 3 f/op

left)

s.n.

top

rigA/)

F K - 3 9 2 (ста/re fe/W

tenehrica

F K - 4 0 2 <
FK-193

(bottom right)

Kattermann:

Erioayce

Sect. II. Neoporteria subsect. 3. Neoporteria (spp. nos. 29-33) Stem globular to elongated, sometimes to 100 cm, 5—18 cm diam.; ribs many, 15—25, deeply depressed or notched between the areoles, 0.5-2 cm high; areoles large, woolly; spines numerous, 20—60, 1-4 cm; roots fibrous or large and tuberous. Flowers from young areoles, narrowly funnelform to almost tubular, carmine red, inner perianth-segments directed inward, outer segments often strongly recurved; pericarpel and tube mostly 1 naked, only sometimes with long white wool and bristles; stigma-lobes upright to spreading, mostly cream-coloured, hut sometimes reddish; nectary modified with widened base, rounded like a bowl, sometimes t angular. Fruit ovoid, thin-walled fleshy, red, elongating by 2-5x before ripening, dehiscent by a basal pore; area of attachment of perianth remnants mostly small. Seeds as in subsect. Neochilenia. D I S T R I B U T I O N . Chile, 27-37°S. Most populations are found at low elevations near the sea, more rarely inland at higher altitudes. The highest population recorded is at almost 2000 m. Diagnostic feature: carmine red, narrow funnelform to tubular flower I adapted to hummingbird pollination). 29. Eriosyce chilensis (Hildmann ex Schumann) Kattermann Stem globular to elongate, sometimes reaching 100 x 6—12 cm, when greatly elongated 1 procumbent, yellowish to grass green; ribs 14—16, 5 mm high, deeply notched or depressed between the areoles, tuberculate; spines numerous, mostly yellowish to brownish, stiffly acicular, often not easily divisible into radial and central spines, radials 16-24, thinly bristly, 10-15 mm, centrals 6-8, thicker, 15-25 mm; roots fibrous. Flowers from young areoles, sometimes > 1 flower per areole; pericarpel and tube bract-scale axils with tufts of wool and glassy-white bristles; perianth-segments curved outward, carmine red or apical one-third carmine red; style 20 x 1.6 mm, cream-white; ovary locule apex updrawn; stigma-lobes 5-10, 4-6 mm, cream, erect to spreading; nectary modified, widened at base, base ± rounded, 2-3 mm. Fruit elongating, bright red, 3 x 1.5 cm, dehiscent by a basal pore, attachment firm; perianth remnant attachment area 50% of fruit diam. Seed [SEM Plate XV| 1.14 x 1.24 mm, asymmetric ratio 0.5; testa tuberculate, slightly ribbed, brownish; hilum keyhole-shaped; micropyle in raised portion of hilum rim, sometimes enclosed by surrounding cells. DISTRIBUTION (MAP 13, page 115). Chile, coastal region of border between Prov. Choapa and Prov. Petorca. K E Y TO VARIETIES OP E. CHILENSIS

Flowers fuchsia red, sometimes with white centre Flowers yellow tinged with red

a. var. chilensis b. var. albidiflora

a. var. chilensis D I S T R I B U T I O N . Chile, Pichidangui to Punta Molles. SPECIMEN EXAMINED. CHILE. Petorca: La Ligua, Punta Molles, FK-3 (see Figure 6.5. page 49); the same, FK-193.

b. var. albidiflora (Ritter) Kattermann Flowers yellow tinged with red. DISTRIBUTION. Chile, known only from the type locality at Pichidangui, Prov. Choapa. SPECIMENS EXAMINED. CHILE. Choapa: Pichidangui, FK-192. The funnelform flower appears to represent a reversal from hummingbird pollination to bee pollination syndrome. At the southern end of its distribution, the flowers are much as in the rest

102

Kattermann:

Eriosyce

of subsect. Neoporteria, except that the inner perianth-segments are spreading outward. Within the next few 100 metres flowers are more funnelform. Over the next 11 km the flower colour changes to yellow, and these plants were described by Ritter as var. albidiflora. They grow in an area no larger than 100 m", completely surrounded by housing construction and are thus highly endangered. When not in flower E. chilensis is indistinguishable from E. subgibbosa. The habitat of E. chilensis is completely surrounded by populations of E. subgibbosa.

30. Eriosyce subgibbosa (Haworth) Kattermann Stem globular to elongated, sometimes to 100 x 6-25 cm diam., procumbent when very elongated, yellowish to grass green, or glaucous dark green; ribs numerous, 16-22, 5-20 mm high, deeply notched or depressed between the areoles, tuberculate, tubercles often very large; areoles large, woolly; spines numerous, yellowish to brownish or black, stiffly acicular, often not easily divisable into radials and centrals, mostly straight, but sometimes strongly curved, radials 8-30, thin, needlelike, 10-20 mm, centrals 1-16, thicker, to 40 mm; roots fibrous or with a short turbinate taproot. Flowers from young areoles, sometimes > 1 flower per areole, buds red, opening carmine red in various shades, variable in size from population to population; pericarpel and tube bractscale axils with tufts of long or short wool, bristles when present only at top of tube, or extending as far as pericarpel, glassy white; tube narrow funnelform to almost tubular; inner perianthsegments curved inward, narrowly lanceolate, carmine red; style 15—30 x ±1.3 mm, cream to sometimes purplish-reddish; ovary locule apex updrawn; stigma-lobes 5-12, 2—6 mm, cream, orange or fuchsia-red, erect to sometimes spreading; nectary modified, widened and ± rounded at base, 5-8 mm. Fruit elongating, bright red. dehiscent by a basal pore, attachment firm; perianth remnant attachment area small, 27% of fruit diam. Seed [SEM Pis XIV, XV] 1.09 x 1.07 mm, asymmetric ratio 0.5; testa finely tuberculate, brownish, slightly ribbed; hilum keyholeshaped, flat, 0.44 x 0.21 mm; micropyle in narrow part of hilum, rim enlarged, not raised. DISTRIBUTION (MAP 12, page 112). Chile, from N of La Serena along the coast S to Concepcion. Most populations are found amongst coastal rocks, but some enter river valleys and inhabit lower coastal mountains. K E Y TO INFRASPECIFIC TAXA OF E. SUBGIBBOSA

1. Stem yellowish green (subsp. subgibbosa) . 1. Stem dark glaucous green Isubsp. clavata) 2. Radial spines thinly acicular, 8-30, 15—40 mm; flower 3-5 cm 2. Radial spines hair-like, 30-40, 10-15 mm; flower 2.5-3 cm .

2

4 3 30.1b. var. litoralis

3. Stem elongating; stigma-lobes included within perianth-segments 30.1a. subsp. & var. subgibbosa 3. Stem rarely elongating; stigma-lobes exserted beyond perianth-segments 30.1.C. var. castanea 4. 4. 5. 5.

Stem globular to slightly elongated Stem slender, erect, columnar . . . Flowers 3-5 cm Flower 2.2-2.75 cm

5 6 . 30.2b. var. nigrihorrida 30.2c. var. wagenknechtii

6. Spines 11-13, often curved to tortuous, often thickly acicular 6. Spines 9-24, straight, thinly acicular

30.2a. subsp. & var. clavata . 30.2d. var. vallenarensis

103

Kattermann: P I A T E 21 (page

1

1041

1

2

3

4

5

6

/-.'.

chilensis

var. albidiflora 2-6

Б.

FK-192

subgibbosa

2

var. subgibbosa

FK-207

3

var. subgibboaa

FK-250

4

var. nigrihorrida

5

var. wugenknechtii

6

var. litoratis

FK-474 FK-183

30.1b. var. litoralis (Ritter) Kattermann Radial spines 30-40, 10-15 mm, centrals to 8, 15 mm. DISTRIBUTION. Chile, Coquimbo and N along the coast. SPECIMEN EXAMINED. CHILE. Elqui: Coquimbo, coastal rocks, FK-475. 30.1c. var. castanea (Ritter) Kattermann Stem globular, not usually elongated, tubercles more pronounced; spines often shorter. Flowers with stigma-lobes exserted several mm beyond the longest peri a nth-segments. DISTRIBUTION. Chile, near Villa Prat (Talca) and north to Santa Cruz (Colchagua). SPECIMENS EXAMINED. CHILE. Colchagua: near Santa Cruz, FK-202; near Lolol, FK-201. Talca: Villa Prat, FK-204.

2

3

/'.'.

30.1. E. subgibbosa subsp. subgibbosa 30.1a. var. subgibbosa Stem yellowish to grass green; spines yellowish to dark horncoloured; style cream, 25-30 mm; stigma-lobes not exserted beyond perianth, cream. DISTRIBUTION. Chile, extending from N of Los Vilos to Concepcion at 37°S. Most populations grow along the coast with some of them so close to the ocean that spray from the surf keeps the plants continuously wet. SPECIMENS EXAMINED. CHILE. Choapa: Rio Choapa, FK-414; N, S & E of Pichidangui, FK-5, 7, 324; Los Vilos, FK-8, 191, 410. Valparaiso: S of Valparaiso, FK-250; N of Valparaiso, FK-1 (neotype), 207; Pta Curaumilla, FK-301; Quintay, FK-253. 255. Melipilla: Naltagua, FK-282; Cardenal Caro: Navidad, FK-195;Pta Topocalma, FK-196; Bucalemu, FK-200; Quirigua, FK-761. Curico: Lloca, FK-758; Llico, FK-759. Talca: Cabo Carranza, FK-754; Constitucitin, Pta Valdes, FK-756. Concepcion: Concepci6n, FK-751, 753.

FK-475

P L A T E 2 2 Г/Мф 10Ю

1

Eriosyce

4

subgibbosa

1

var. subgibbosa

2

var. castanea

FK-8

3

var. clavata

4

var. nigrihorrida

FK-204 FK-27 FK-22

30.2. E. subgibbosa subsp. clavata (Sbhrens) Kattermann 30.2a. var. clavata (Sbhrens! Kattermann Stem globular to sometimes much elongated, but not procumbent like var. subgibbosa, 6-10 cm diam., glaucous green; ribs 10-12, 5 mm high; areoles 7 x 5 mm; spines mostly blackish, often curved, radials 10, 15-25 mm, centrals 1-3, 30-35 mm; rootstock a turbinate taproot, sometimes narrowed and necklike at apex. Flower 5.5 cm; style greenish yellow, carmine towards apex; stigma-lobes 7, 3-5 mm, reddish; nectary modified, widened at base, edges angular. DISTRIBUTION. Chile, mainly in the Elqui valley and hills to the south. SPECIMENS EXAMINED. CHILE. Elqui: E of La Serena. FK-184; Las Rojas, FK-27.

Kattermann:

Eriosyce

30.2b. var. nigrihorrida (Backeberg) Kattermann Stem shorter, rarely over 25 cm high; ribs 11-16; spines more numerous, straight to slightly curved, black to horn-coloured, radials 11-16, centrals 4-6, to 50 mm. DISTRIBUTION. Chile, replacing var. clavata

P L A T E 2 3 (page

108)

1

2

3

4

5

6

in a W and S

direction. In the S, N of Los Vilos, populations merge with subsp.

subgibbosa.

SPECIMENS

EXAMINED.

CHILE.

Elqui:

20 km

N of La

Serena, FK-182; N of La Serena, FK-326; E of La Serena, FK-96; Herradura, FK-477; E of Tongoy, FK-22; S of Quebrada Las Palmas, FK-10; Tanque las Palomas, FK-15, 188. Limari: Estancia Frai Jorge, FK-21; Pta Lengua de Vaca, FK-24; Rio Limari, FK-84; near Higueritas, FK-185; Panuelas, FK-474; Pta Teniente, FK-481. 1

The northern populations often form thickened taproots. 30.2d. var. vallenarensis (Ritter) Kattermann Stem elongated, sometimes reaching 30 x 4-6 cm, grey-green; ribs 11-12, 7-10 mm high, obtuse, deeply notched or depressed between the areoles, tuberculate, with a chin below each areole; areoles 6-10 x 4-6 mm, 6-12 mm apart, woolly; spines black, later grey, radials 8-14, 15-30 mm, centrals 1-4, 20-40 mm; rootstock a thick taproot narrowed at apex into a neck. Flowers carmine red, 3.5 cm; pericarpel and tube bract-scale axils with long wool and white bristles; inner perianthsegments 14 x 2.5 mm; style 23 x ±1 mm; stigma-lobes 6. 4 mm, cream, erect to sometimes spreading; nectary 3-5 mm. Fruit bright red. Seeds 0.81 x 0.90 mm, asymmetric ratio 0.6; testa tuberculate. not ribbed; hilum keyhole-shaped, 0.36 x 0.24 mm; hilum rim strongly enlarged at micropyle, not raised. DISTRIBUTION. Chile, from E of Vallenar to Freirina (Huasco). SPECIMEN EXAMINED. CHILE. Huasco: Freirina, FK-79.

E.

subgibbosa

var. vallenarensis

30.2c. var. wagenknechtii (Ritter) Kattermann Stem at first semiglobular, elongating only with age. Flowers very small, 2.2-3 cm; style whitish, 18 mm; stigma-lobes 6, 3 mm, cream-white. Seed 0.83 x 0.90 mm, asymmetric ratio 0.5; testa smoothly tuberculate, not ribbed; hilum keyhole-shaped, 0.33 x 0.21 mm; micropyle in a narrow groove, hilum rim enlarged at micropyle, not raised. DISTRIBUTION. Chile, from Juan Soldado to Los Choros (Elqui). SPECIMENS EXAMINED, CHILE. Elqui: Cruz Grande, FK-178, 179; E of Los Choros, FK-471; Juan Soldado, FK-183.

E.

senilis

FK-418

E. senilis

FK-422

E.

senilis

FK-423

E. senilis

FK-425

£. senilis

FK-452

P L A T E 2 4 /page

FK-79

109)

1

2

3

4

5

6

1-3

E.

1-2

subsp. elquiensis

senilis FK-29

3

subsp. coimasensis

KK-86

4

£ . villosa

5

E. villosa

FK-467

6

E. villosa

FK-814

FK-71

107

FIGURE 24 E. 1

(above)

F I G U R E 2 5 (right)

subgibbosa

1 - 4 E.

var. subgibbosa

FK 207.

Л. 4 . 5 cm, s t i g m a 6 m m . 2

var. castanea

3

var. clavata

1 2

var.

FK-22

3

nigrihorrida,

var. wagenknechfii

4

var. vatlenarensis

5

E. sociabilis

FK-79

FK-147,

fl. 2.5 c m , s t i g m a 3 m m .

form from P a n u e l a s . F K - 4 7 4

FK-204 FK 27

subgibbosa

var. nigrihorrida

FK-183

Kattermann:

Eriosyce

III

Kattermann:

Eriosyce 31. Eriosyce sociabilis (Ritter) Kattermann Stem globular to elongated, 3-10 cm diam.; ribs 13-16, notched into tubercles with a chin below each areole; areoles 10 x 7 mm, 3 mm apart, sunken; spines irregular, mostly straight, dark brown to blackish, radials 6-20, 3-20 x 0.25 mm, centrals 6-12, 20-40 x 0.55 mm; rootstock a short, turbiniform taproot. Flowers borne at young areoles, narrow funnelform. 2-3 cm; pericarpel and tube bractscale axils with white floccose wool and white, straight bristles; perianth-segments fuchsiared; style 12-15 mm, purplish red; ovary locule apex updrawn; stigma-lobes 4—8, 3-5 mm, purplish red, erect to spreading; nectary widened ut base, 1-2 mm. Fruit elongating, seed loose within, dehiscent hy a basal pore, attachment firm; perianth remnant attachment area small, 28% of fruit diam. Seed [SEM Plate XV| 1 x 1 mm, asymmetric ratio 0.6; testa slightly ribbed; hilum 0.31 x 0.21 mm; micropyle in a fairly wide groove, hilumrimnot raised at micropyle. DISTRIBUTION (MAP 13). Chile, S of Totoral Bajo (Huasco) to S of Caldera (Copiap6). SPECIMEN

EXAMINED.

CHILE.

Copiapo/

Huasco: S of Totoral Bajo, FK-147. On Morro Copiapo E. sociabilis merges with E. taltalensis var. transiens, to which it appears closely related. Transitional flower forms between the two species have been observed and Ritter reports that the flowers of E. sociabilis are more open than other species here included in subsect. Neoporteria.

MAP

1 2 . Distribution of E. subgibbosa

component taxa.

112

a n d its

32. Eriosyce senilis I Backeberg) Kattermann Stem subglobular to globular or elongated. 4-12 cm diam.; ribs 13-21, 3-8 mm high, deeply depressed or notched between the areoles; areoles woolly, 6-10 x 4-6 mm; spines all or some hairlike, white, greyish, yellow, brown or sometimes black, straight to strongly contorted, thin, stiffly needlelike to soft, flexible and bristlelike, radials 15-40, 20-60 mm, centrals 4-20, 30-60 mm; rootstock a large tuberous taproot. Flowers from young areoles, buds red, opening carmine red in various shades; pericarpel and tube bract-

Kattermann:

Eriosyce

scale axils with tufts of wool, in some populations with long wool, bristles always present near apex of tube, sometimes extending to pericarpel; tube narrow funnelform to almost tubular; inner perianth-segments curved inward; style thin, ±1 mm diam.; ovary locule apex updrawn; stigmalobes 6-12, 4—6 mm, cream, erect to sometimes spreading; nectary widened at the t rounded base, 5—8 mm. Fruit elongating, bright red dehiscent by a basal pore, attachment firm; perianth remnant attachment area small. Seed [SEM Plate XVI] 1.09 x 1.19 mm, asymmetric ratio 0.5-0.6; testa ribbed, brownish; hilum keyhole-shaped, 0.36 x 0.23 mm; micropyle in a groove, hilum rim modified at micropyle, barely raised. DISTRIBUTION (MAP 13). Chile, from the Elqui Valley to Ovalle, S to Cuesta Las Chilcas, almost always inland and at higher elevation (to 2000 m). KEY TO SUBSPECIES OF E. SENILIS

1. Spines tortuous 1. Spines i straight 2. Flower with scant wool on pericarpel and tube, bristles few

2 32.2. subsp. coimasensis 32.1. subsp. senilis

2. Flower with dense wool on pericarpel and tube, bristles many , . 32.3. subsp. elquiensis 32.1. E. senilis subsp. senilis Stem semiglobular to rarely elongated; spines soft tortuous, in some populations glassy-white, in others straw-yellow or dark brown, or in combination. Flowers variable in size, from 5-7 cm. DISTRIBUTION. Chile, throughout the Rio Choapa valley and adjoining areas (Prov. Choapa/Limari). SPECIMENS EXAMINED. CHILE. Choapa: Cuncumen, FK-418; Quelen, FK-423; Salamanca, FK-425; Coyton, FK-422 (white form). Limari: E of Ovalle, FK-452. The type locality originally cited by Ritter was the region of Coquimbo, but in Kakteen in Sudamerika (1980) he gives it as E of Salamanca. It is the latter which is assumed to be correct, since the pictures published show plants from Salamanca. 32.2. E. senilis subsp. coimasensis (Ritter) Kattermann Stem 5-18 x 8-12 cm; ribs 15-21, deeply notched between the areoles; areoles 5-10 ram, 5-10 mm apart; spines straight to curved, thin, needlelike, greyish to blackish, radials 15-30, 10-40 mm, centrals 8-20, 20-40 mm. Flower 5.5 cm; style 40 mm, whitish, pale pink towards apex; stigma-lobes 7, 5-9 mm, cream; nectary modified, very large, 10 mm. DISTRIBUTION. Chile, from Las Coimas S to Monte Negro (N of Santiago). All the populations grow quite a distance from the coast. SPECIMENS EXAMINED. CHILE. San Felipe de Aconcagua: Las Coimas, FK-266. Chacabuco: near Monte Negro, A. Hoffmann in FK-406. Santiago: Cuesta Las Chilcas, FK-86. Flower size and stem spination place these plants with E. senilis. 32.3. E. senilis subsp. elquiensis Kattermann Stem globular to slightly elongated; spines whitish, often mixed with stiffer, blackish spines. Pericarpel and tube with more wool and bristles; stigma-lobes often carmine red. DISTRIBUTION. Chile, throughout the central Elqui Valley. SPECIMENS EXAMINED. CHILE. Elqui: W of Vicuna, FK-29; El Tambo, FK-462 (holotype).

113

Kattermann:

114

Eriosyce

Kattermann:

MAP

13. Distribution of E. chilensis.

E. senilis

a n d E.

E.

Eriosyce

stxiabilis,

villosa.

F I G U R E 2 6 flefh E.

senilis

1

subsp. senilis

F K - 4 1 H , fl. 6 c m .

2

subsp. senilis

F K - 4 5 2 , fl. 5 . 5 cm.

3

н и Ь н р . coimasensis

4

s u b s p . senilis

5

subsp. elquiensis

FK-86

FK-425 FK-29

F I G U R E 2 7 (above) E. villosa

FK-71

115

Kattermann:

Erioayce

33. Eriosyce villosa (Monville) Kattermann Stem semiglobular to globular, later elongated, 4—10 cm diam.; ribs 13-15, 3-8 mm high, deeply depressed or notched between the areoles; areoles 6-7 mm diam., 5-10 mm apart; spines all or some hairlike, some stiffly needlelike, brownish to black, radials 12-20,15-25 mm, centrals 4-12, often stiffly needlelike, but sometimes soft and flexible; rootstock a large tuberous taproot. Flowers from young areoles, small, 2.2-3 cm, buds red, opening to carmine red in various shades; pericarpel and tube bract-scale axils with tufts of long wool and many glassy-white bristles; inner perianth-segments curved inward; style thin, ±1.5 mm diam.; ovary locule apex updrawn; stigmalobes 6-12, 2-4 mm, cream, erect to sometimes spreading; nectary widened at the ± rounded base, 2-4 mm. Fruit elongating, bright red, dehiscent by a basal pore, attachment firm; perianth remnant attachment area 37% of fruit diam. Seed |SEM Plate XVI] 0.90 x 0.81 mm, asymmetric ratio 0.5; testa tuberculate, not ribbed, brownish; hilum keyhole-shaped, 0.24 x 0.16 mm; micropyle in a groove, hilumrimenlarged at micropyle, not raised. DISTRIBUTION (MAP 13). Chile, from Totoral Bajo (Prov. Copiapo) S to Carrizallilo (Huasco). SPECIMENS EXAMINED. CHILE. Huasco: N of Huasco, FK-71 (neotype); Huasco, FK-164 (this is probably what Backeberg described as Neoporteria

atrispinosa

cephalophora);

Sarco, FK-467

(Neoporteria

Backeberg?); Carrizal Bajo, FK-814.

Plants collected by W. Jung from Carrizallilo are intermediate between the form from Sarco and E. subgibbosa

var. wagenknechtii.

Indeed E. villosa

appears to be closely related to E.

subgibbosa

var. wagenknechtii on flower and seed characters. Over a distance of about 60 km the seed testa changes from ribbed to smooth iE. villosa having the smooth seed).

116

Kattermann:

Eriosyce

APPENDIX I Validation of new names In the following list all new names used in the above revision are validated. Supplementary details of typification, synonymy and bibliography not essential for valid publication can be found in Appendix IV. E r i o s y c e I sect. E r i o s y c e (Berger)

Kattermann

Pyrrhocactus

subsect.

comb,

Pyrrhocactus

et

stat.

nov.

6. E r i o s y c e Neochilenia Sukk.

Berger, K a k t e e n , .345 ( 1 9 2 9 ) .

andreaeana andreaeana

10: 3 8 ( 1 9 5 9 ) ,

Holotype: Kattermann Eriosyce

Isect.

Eriosyce]

subsect.

Blatt.

Kakteenf.

1934(10):

Islaya

[unpaged,

sect.

Kattermann

Neoporteria

comb,

et

stat.

(Britton nov.

&

Rose)

Neoporteria

E r i o s y c e |sect. N e o p o r t e r i a I subsect. H o r r i d o ­ nov.

593 (DBG).

7. E r i o s y c e v i l l i c u m e n s i s

(Rausch) K a t t e r m a n n

comb.

villicumensis

nov.

Pyrrhocactus

Rausch,

(Backeberg) K a t t e r m a n n c o m b , e t

Horridocactus

Backeberg.

8.

Eriosyce

comb.

islayensis

nov.

(Forster)

Echinocactus

Kattermann

islayensis

Forster,

H a m b u r g . G a r t . Blumenzeit. 17: 1 6 0 ( 1 8 6 1 ) .

Britton & Rose. Cact. 3: 9 4 ( 1 9 2 2 ) .

cactus

Blatt.

stat.

Kakteenf.

9. E r i o s y c e r e c o n d i t a (Ritter) K a t t e r m a n n c o m b , nov.

Pyrrhocactus

reconditus

Ritter.

Succulents

1962: 2 7 (1962). 9.2.

1938(6): [unpaged, p. 2 1 Ц 1 9 3 8 ) .

Eriosyce

recondita

subsp.

iquiquensis

(Ritter) K a t t e r m a n n c o m b , e t s t a t . n o v . E r i o s y c e [sect. N e o p o r t e r i a l subsect. C h i l e o s y c e Kattermann

subsect.

n o v . Chileorebutia

Ritter,

C a c t u s (Parisl 14(64»: suppl. [unpaged, p. 5 | ( 1 9 5 9 ) , пот.

illegit.,

Holotype:

typ.

& C. napina

Eriosyce

I Chileorebutia

krausii

krausii

(Phil.)

(Ritter)

Ritter,

ibid.

Ritter

excl.

Kattermann 15(66):

6-7.

1960).

cactus iquiquensis 10

(Britton

Neoporteria

&

Rosel

Kattermann

Neoport­ stat.

nov.

Britton & Rose, Cact. 3 : 9 4 ( 1 9 2 2 ) .

Pyrrho­

Ritter, T a x o n 12: 3 2 ( 1 9 6 3 ) .

Eriosyce

curvispina

(Bertero

ex

K a t t e r m a n n c o m b . n o v . Cactus curvispinus

Colla) Bertero

ex Colla. M e m . Acad. Sci. Torino 3 7 : 7 5 - 7 6 ( 1 8 3 3 ) . 10b. E r i o s y c e (Ritter)

E r i o s y c e [sect. N e o p o r t e r i a l subsect. eria

and. 37.1).

Kakt. and. Sukk. 2 5 : 2 6 8 (1974).

p. 3 ] ( 1 9 3 4 ) . Eriosyce

nov.

Kakt.

(Arts 9.5 &

Islaya

(Backeberg) K a t t e r m a n n c o m b , e t s t a t . nov. Backeberg.

Kattermann sp. Backeberg,

invalid

curvispina

Kattermann

aconcaguensis

var.

comb.

Pyrrhocactus

Ritter, Succulenta 1 9 6 0 : 1 0 8 ( 1 9 6 0 ) .

10c. E r i o s y c e c u r v i s p i n a Kattermann

aconcaguensis

nov.

comb.

nov.

var. a r m a t a (Ritter) Pyrrhocactus

armatus

Ritter, Succulenta 1 9 6 0 : 4 9 ( 1 9 6 0 ) . lb.

Eriosyce

aurata

var. s p i n i b a r b i s

(Ritter)

K a t t e r m a n n c o m b , e t s t a t . n o v . Eriosyce barbis

spini­

Ritter, K a k t . S u d a m e r . 3: 9 1 5 ( 1 9 8 0 ) .

lOd.

Eriosyce

8.

Eriosyce

mann

bulbocalyx

comb.

nov.

( W e r d e r m a n n i Katter­

Echinocactus

choapensis

10c.

strausianus

Schumann.

M o n a t s s c h r . K a k t . - K u n d e 11: 1 1 2 ( 1 9 0 1 ) . Eriosyce

strausiana

var.

Eriosyce

Pyrrho­

R a u s c h , K a k t . and. S u k k . 2 6 :

73 (1975).

choapensis Pyrrho­

1960:

133

var.

comb.

tuberisulcata

nov.

Echinocactus

Jacobi, Allg. G a r t e n z . 2 4 : 1 0 8 ( 1 8 5 6 ) .

lOf. E r i o s y c e c u r v i s p i n a var. m u t a b i l i s (Ritter)

mutabilis

horridus

var.

Ritter, K a k t . S u d a m e r . 3 : 9 4 6 ( 1 9 8 0 ) .

lOg. E r i o s y c e c u r v i s p i n a Kattermann

comb.

nov.

var. r o b u s t a Pyrrhocactus

(Ritter) robustus

Ritter, Succulenta 1 9 6 0 : 6 5 ( I 9 6 0 ) .

5. E r i o s y c e u m a d e a v e ( W e r d e r m a n n ) K a t t e r m a n n c o m b . n o v . Echinocactus

curvispina

Kattermann

tuberisulcatus

K a t t e r m a n n c o m b . n o v . Pyrrhocactus

pachacoensis

(Rausch) Kattermann c o m b , et stat. n o v . cactus pachacoensis

var.

Succulenta

(1960).

(Jacobi) •1. E r i o s y c e s t r a u s i a n a ( S c h u m a n n ) K a t t e r m a n n

ih.

Ritter,

bulbocalyx

W e r d e r m a n n , Bliih. K a k t . , t. 1 3 6 ( 1 9 3 7 ) .

c o m b . n o v . Echinocactus

curvispina

(Ritter) K a t t e r m a n n c o m b , e t s t a t . n o v . cactus

umadeave

Werdermann,

M o n a t s s c h r . Deutsch. K a k t . - G e s . 3: 2 2 3 ( 1 9 3 1 ) .

11. E r i o s y c e k u n z e i (Forster) K a t t e r m a n n c o m b , n o v . Echinocactus

kunzei

Forster, H a n d b . Cact. 2 9 3

(1845M'kunzii').

117

Kattermann:

lib

E r i o s y c e k u n z e i var. t r a n s i t e n s i s (Ritteri

Kattermann transitensis 12.

comb,

et

stat.

nov.

Pyrrhocactus

Eriosyce

limariensis

i Hitter)

limariensis

19

Eriosyce heinrichiana

mann

comb.

nov.

I Backeberg) Katter­

Horridocactus

heinnchianus

Backeberg, K a k t . - K u n d e 1 9 4 2 : 8 ( 1 9 4 2 ) .

Ritter, T a x o n 12: 3 3 (19631.

c o m b . n o v . Pyrrhocactus

Eriosyce

KattcrmHnn Ritter, Kakt.

S u d a m e r . 3 : 9 5 6 (1980».

1 9 . 2 . E r i o s y c e h e i n r i c h i a n a subsp. i n t e r m e d i a I Ritter) K a t t e r m a n n c o m b , e t s t a t . n o v .

Pyrrho­

cactus

Ritter,

setosiflorus

var.

intermedius

Succulenta 1 9 6 2 : 7 0 ( 1 9 6 2 ) . 13. E r i o s y c e c o n f i n i s (Ritter) K a t t e r m a n n c o m b , n o v . Pyrrhocactus

confinis

Ritter. Succulenta 1 9 6 1 :

4 (1961). 14. E r i o s y c e c r i s p a (Ritteri K a t t e r m a n n c o m b , n o v . Pyrrhocactus

crispus

Ritter, Succulenta 1 9 5 9 :

137 ( 1 9 5 9 ) .

Kattermann

comb,

et

stat.

nov.

Pyrrhocactus

Ritter, Succulenta 1 9 6 0 : 8 9 ( I 9 6 0 ) .

14.2a. E r i o s y c e c r i s p a var. a t r o v i r i d i s (Ritter) Kattermann

stat.

n o v . Pyrrhocactus

atroviridis

Ritter. Succulenta 1 9 6 0 : 8 9 | I 9 6 0 ) . 14.2b. E r i o s y c e c r i s p a var. h u a s c e n s i s (Ritter) Kattermann huascensis

comb,

et

stat.

nov.

Pyrrhocactus

Ritter, Succulenta 1 9 6 1 : 5 7 ( 1 9 6 1 ) .

14.2c. E r i o s y c e c r i s p a var. c a r r i z a l e n s i s (Ritter) K a t t e r m a n n c o m b . n o v . Pyrrhocactus

carrizalensis

Ritter, T a x o n 12: 3 3 ( 1 9 6 3 ) .

totoralensis

Ritter. Succulenta 1 9 6 1 : 1 3 1 ( 1 9 6 1 ) . 15.

Eriosyce nov.

garaventae Horridocactus

Eriosyce aspillagae

comb.

nov.

(Ritter) K a t t e r m a n n garaventae

Ritter,

aspillagae

Sohrens,

Monatsschr. Deutsch. K a k t . - G e s . 1: 1 2 5 ( 1 9 2 9 ) . 17.

Eriosyce

comb.

nov.

marksiana Pyrrhocactus

• Ritter) K a t t e r m a n n marksianus

var. i n t e r m e d i a

nov.

Pyrrhocactus

Ritter,

Succulenta

Eriosyce heinrichiana

var.

setosiflora

(Ritteri K a t t e r m a n n c o m b , e t s t a t . n o v . cactus

setosiflorus

Ritter,

Succulenta

Pyrrho­

1962: 70

(1962). 19.3.

Eriosyce

heinrichiana

вир.

simulans

(Ritter) K a t t e r m a n n c o m b , et s t a t . n o v . cactus simulans

Pyrrho­

Ritter. Succulenta 1 9 6 1 : 3 5 ( 1 9 6 1 ) .

20. E r i o s y c e n a p i n a I Philippi I K a t t e r m a n n c o m b , n o v . Echinocactus

napinus

Philippi. l)escr. nuev.

pi.: 6 2 - 6 3 ( 1 8 7 2 ) . 2 0 . 2 . E r i o s y c e n a p i n a subsp. l e m b e k e i Katterm a n n s u b s p . n o v . Ne
Backeberg,

Die Cact. 3 : 1 8 2 2 ( 1 9 5 9 ) , i n v a h d ( A r t s 9 . 5 & 3 7 . l l .

20.2a.

77 (DBG).

Eriosyce napina

Wt,

m a n n v a r . n o v . Ncncfulenia Holotype: Kattermann

lembekei lembekei

Katter­

Backeberg,

Ritter.

Succulenta 1 9 6 0 : 2 119601.

77 (DBG).

2 0 . 2 b . E r i o s y c e n a p i n a var. d u r i p u l p a (Ritter) Kattermann

(Siihrens) K a t t e r m a n n

Echimxactus

intermedius

D i e Cact. 3 : 1 8 2 2 11959), invalid (Arts 9 . 5 & 3 7 . 1 ) .

Succulenta 1 9 5 9 : 4 1 ( 1 9 5 9 ) . lfi.

var.

Holotype: Kattermann

14.2d. E r i o s y c e c r i s p a var. t o t o r a l e n s i s (Ritter) K a t t e r m a n n c o m b . n o v . Pyrrhocactus

comb.

Kattermann c o m b .

1962: 7 0 ( 1 9 6 2 ) . 19.2b.

1 4 . 2 . E r i o s y c e c r i s p a subsp. a t r o v i r i d i s (Ritter) atroviridis

19.2a. E r i o s y c e h e i n r i c h i a n a (Ritteri

setosiflorus

duripulpa 21.

comb,

et

stat.

nov.

Chileorebutia

Ritter, T a x o n 12: 1 2 3 ( 1 9 6 3 ) .

Eriosyce

Kattermann

odieri comb.

( L e m a i r e ex nov.

Salm-Dyck)

Echinocactus

odieri

Ix-maire ex S a l m - D y c k . Cact. Hort. Dyck. 1 8 4 9 : 1 7 4 (1850). 2 1 . 1 b . E r i o s y c e o d i e r i var. m o n t e - a m a r g e n s i s

17b. E r i o s y c e m a r k s i a n a var. l i s s o c a r p a (Ritter)

Kattermann

K a t t e r m a n n c o m b . n o v . Pyrrhocactus

amargensis

lissocarpus

var.

nov.

Backeberg,

Neochilenia

Descr.

Cact.

monteNov.

( 1 9 6 3 ) , invalid (Art. 9.51. Holotype: Kattermann

Ritter. Succulenta 1 9 6 0 : 17 ( 1 9 6 0 ) .

3:

9

522

(DBG). 17c. E r i o s y c e m a r k s i a n a var. g r a c i l i s (Ritteri K a t t e r m a n n c o m b . n o v . Pyrrhocactus var. gracilis

lissocarpus

Ritter, Succulenta I 9 6 0 ; 17 ( I 9 6 0 ) .

IS. E r i o s y c e e n g l e r i (Ritteri K a t t e r m a n n c o m b , n o v . Horridocactus 76 (1959).

engleri

Ritter. Succulenta 1 9 5 9 :

2 1 . 2 . E r i o s y c e o d i e r i subsp. g l a b r e s c e n s i Ritter I Kattermann glabrescens

comb,

et

stat.

nov.

Chileorebutia

Ritter, C a c t u s (Paris) 15(66): 9 (196ЧК

2 1 . 3 . E r i o s y c e o d i e r i subsp. f u l v a (Ritter) Katter­ mann

comb,

et

stat.

n o v . Chileorebutia

Ritter, C a c t u s (Paris) 15(66): 10 ( I 9 6 0 ) .

118

fulva

Kattermann:

Eriosyce

2 2 . E r i o s y c e t a l t a l e n s i s iHutchison) K a t t e r m a n n

.40. E r i o s y c e s u b g i b b o s a ( H a w o r t h ) K a t t e r m a n n

c o m b . n o v . Neoporteria

comb.

taltalensis

Hutchison, Cact.

Succ. J. ( U S ) 2 7 : 181 ( 1 9 5 5 ) . 2 2 . 1 b . E r i o s y c e t a l t a l e n s i s var. p y g m a e a I Ritter) K a t t e r m a n n c o m b . n o v . Pyrrhocactus

pygmaeus

Hitter, T a x o n 12: 3 2 ( 1 9 6 3 ) . 22.2. E r i o s y c e t a l t a l e n s i s

paucicostatus

Echinocactus

subgibbosus

Haworth,

30.1 b. E r i o s y c e s u b g i b b o s a var. l i t o r a l i s (Hitter i Kattermann litoralis

subsp.

paucicostata

(Ritter) K a t t e r m a n n c o m b , e t s t a t . n o v . cactus

nov.

Phil. M a g . 10: 4 1 9 ( 1 8 3 1 ) .

Ritter. Succulenta

Horrido­ 1959: 1 1 3

(1959).

comb,

et

stat,

Ritter. Succulenta

nov.

Neoporteria

1959: 2 8 < 1 9 5 9 ) .

30.1 c. E r i o s y c e s u b g i b b o s a var. c a s t a n e a (Ritter) Kattermann castanea 30.2.

comb,

et

stat.

nov.

Neoporteria

Ritter. T a x o n 12: 3 4 (19631.

Eriosyce

subgibbosa

subsp.

clavata

2 2 . 3 . E r i o s y c e t a l t a l e n s i s subsp. e c h i n u s (Ritteri

(Sohrens ex S c h u m a n n I K a t t e r m a n n c o m b , e t s t a t .

Kattermann

n o v . Echinocactus

echinus

comb,

et

stat.

nov.

Pxrrhtxractus

Ritter, T a x o n 12: 3 3 ( 1 9 6 3 ) .

clavatus

Sohrens ex S c h u m a n n ,

M o n a t s s c h r . K a k t . - K u n d e 10: 2 7 ( 1 9 0 0 ) .

2 2 . 3 a . E r i o s y c e t a l t a l e n s i s var e c h i n u s (Ritter)

3 0 . 2 a . E r i o s y c e s u b g i b b o s a var. c l a v a t a (Sohrens

Kattermann

ex S c h u m a n n ) K a t t e r m a n n s t a t . n o v .

comb.

nov.

Pyrrhocactus

echinus

clavatus

Ritter, T a x o n 12: 3 3 ( 1 9 6 3 ) . 2 2 . 3 b . E r i o s y c e t a l t a l e n s i s var. f l o c c o s a (Hitter) Kattermann floccosus 22.4.

comb,

et

stat.

nov,

Pyrrhocactus

Ritter, T a x o n 12: 3 2 ( 1 9 6 3 ) .

Eriosyce

taltalensis

pilispina

(Ritter) K a t t e r m a n n c o m b , e t s t a t . n o v . cactus pilispinus

Pyrrho­

Ritter. Succulenta 1 9 6 2 : 4 2 ( 1 9 6 2 ) .

2 3 . E r i o s y c e o c c u l t a K a t t e r m a n n s p . n o v . ab K. odieri M M t O tuberculorum taltalense subsp. evolutis mann

et ab

E.

taltalen.se

E . recondita spinis

vix

albidis

differ!. Holotype: Katter­

et floribus

caulis

acuta

391 (DBG). I Ritter) K a t t e r m a n n

comb.

esmeraldana

Chileorebutia

Ritter,

T a x o n 12: 123 ( 1 9 6 3 ) . 26.

Eriosyce

aerocarpa

c o m b . n o v . Chileorebutia

(Hitter)

аепнагра

Kattermann Ritter. C a c t u s

(Paris) 15(66): 8 ( I 9 6 0 ) . 26

E r i o s y c e k r a u s i i (Ritter) K a t t e r m a n n c o m b ,

n o v . Chileorebutia

krausii

Ritter, C a c t u s (Paris)

15(66): 6 - 7 ( I 9 6 0 ) . 27.

Eriosyce

comb.

Neoporteria

nigrihorrida

tenebrica

n o v . Thelocephala

(Ritter) tenebrica

Kattermann Ritter, K a k t .

2 8 . E r i o s y c e l a u i J. Luthy: see A p p e n d i x II.

30.2c. E r i o s y c e s u b g i b b o s a var. (Ritteri

Kattermann

wagenknechtii

(Ritter)

comb.

Kattermann

wagenknechtii

nov.

Gesamtb.

423

comb,

vallenarensis

vallenarensis

et

stat.

nov.

Ritter. Kakt. S u d a m e r . 3:

n o v . Neoporteria

sociabilis

Ritter, Succulenta 1 9 6 3 :

3(1963). 32.

Eriosyce

comb.

nov.

senilis

(Backeberg)

Neoporteria

senilis

Kattermann

Backeberg

Backeb. & K n u t h , K a k t u s - A B C , 2 6 2 ( 1 9 3 5 ) ;

Kattermann

differt.

in

Echino­

Philippi ( 1 8 8 6 ) non Beaton < 1839).

3 2 . 2 . E r i o s y c e s e n i l i s ssp. c o i m a s e n s i s

32.2b.

chilensis Kakt.

Neoporteria

3 1 . E r i o s y c e s o c i a b i l i s (Hitter) K a t t e r m a n n c o m b ,

comb,

et

stat.

nov.

(Ritteri

Neoporteria

Ritter, T a x o n 12: 3 4 ( 1 9 6 3 ) .

Eriosyce

pericarpello

Kchinin-actus

cum

1032 (1980).

Kattermann

nov.

nov.

Backeb. in

Ritter, Succulenta 1 9 6 3 : 5 (19631.

Neoporteria

Kattermann

Schumann.

stat.

nigrihorrida

lat., пот. inval. ( A r t . 4 3 . 1 ) .

29. E r i o s y c e c h i l e n s i s ( H U d m a n n ex S c h u m a n n ) comb.

nigrihorrida et

Backeb. & K n u t h , K a k t u s - A B C , 4 1 7 ( 1 9 3 5 ) , diagn.

coimasensis

ex

var.

comb,

Backeberg, K a k t . - K u n d e

1939: 8 1 ( 1 9 3 9 ) ; Chilenia

cactus senilis

S u d a m e r . 3 : 1001 ( 1 9 8 0 ) .

Hildmann

subgibbosa

Kattermann

3 0 . 2 d . E r i o s y c e s u b g i b b o s a var.

24. E r i o s y c e e s m e r a l d a n a nov.

K u n d e 10: 2 7 ( 1 9 0 0 ) . 30.2b. E r i o s y c e (Backeberg)

subsp.

Echinocactus

Sohrens ex S c h u m a n n . Monatsschr. Kakt.-

senilis

subsp.

subsp.

nov.

et tuba florum

a

magis

Holotype: Kattermann

elquiensis

subsp. lanatis

senile setosisque

462 (DBG).

(1898). 3 3 E r i o s y c e v i l l o s a (Monville) K a t t e r m a n n c o m b , 2 9 b . E r i o s y c e c h i l e n s i s var. a l b i d i f l o r a (Ritter)

n o v . Cactus

K a t t e r m a n n c o m b . n o v . Pyrrhocactus

(1839).

albidtflorus

chilensis

var.

villosus

Monville, Hort. Univ. 1: 2 2 3

Ritter. K a k t . S u d a m e r . 3 : 9 2 7 ( 1 9 8 0 ) .

119

Kattermann:

Eriosyce

APPENDIX II Eriosyce laui J. Liithy sp. nov.

by Jonas M. Luthy Botanische Institute, Universitat Bern Altenbergrain 21, CH-3013 Bern, Switzerland Introduction During a visit to Alfred Lau's home in Mexico, in 1989, he showed me one of his discoveries, a curious cactus he had first seen in Chile in October 1971, many years before he eventually collected it in 1986 (after the IOS Congress in Salta, Argentina). It was a dwarf plant covered with wool, flowering at the time of my visit (see Plate 19.6) and showing such a strange combination of characters, that I was not able to tell even the genus it might belong to. After study and discussions with experts, I now feel sure enough about the taxon's place in the phylogenetic system to describe it. In line with Kattermann's amplification and revision of Eriosyce I am including this new taxon within that genus as now understood, which comprises a phylad characterized by the synapomorphy of basally opening, non-fleshy fruits. If a narrower generic circum­ scription were to be adopted, I would place this species in Islaya Backeberg, which Kattermann reduces to infrageneric rank. So, in my opinion (contrary to that of Kattermann) Eriosyce laui sp. nov. belongs to subsect. Islaya, where it is admittedly a rather aberrant species, showing remarkable morphological differences to all other species of Eriosyce s.l. During studies of flower morphology of Eriosyce s.l. and Copiapoa Britton & Rose a new chemical character, seemingly specific for the genus Copiapoa, was found, which serves to exclude the new species from the latter genus, as discussed below. Eriosyce laui J. Liithy sp. nov. reactione stigmae поп nigricantis in liquore alcoholico-formaldehydico-glycerinico et fructu basi refringens perspicue a genero Copiapoa separanda; fructu post elongatione et inflatione basi abrumpente generi Eriosyce sens. lat. vel Islaya sens. str. pertinens sed axillis inflorescentiae denudatis ab omnibus speciebus praesenti tempore cognitis separanda; testa seminis leviter colliculosa, cuticula sublaevi quoque notabilis; hylum lateraliter subterminale, anguste lageniforme, micropyle constrictione hyli fere separata; podaria ad extremitatem appendicem foliiformam in plantis juvenilibus adultisque ferentia. Holotypus: Chile, S of Tocopilla, 5 km from the coast, in rocky outcrops of mountain slopes, 400 m, 1986, Lau 1541 (ZSS). Stem to 3 cm diam., globose to somewhat elongated towards the rootstock, from which it is distinctly separated; taproot to 15 cm, napiform, succulent, brittle and rapidly drying when taken out of the substrate, strongly compressed within rock fissures; epi­ dermis whitish green to tinged red, depending on exposure, relatively unstructured and

120

Erioayce laui J. Liithy

FIGURE 28 Erioayce

laui.),

Luthy. A , seedling, 6 months' old, showing leaflike podarin a p p e n d a g e s (scale = В m m ) . B,

flower (scale = 5 m m ) . 0 , bract-scale with papillate epidermis (scale = 1 m m ) . D , longitudinal section of flower (scale • 5 m m ) . E , longitudinal section of ovary at anthesis (scale = 1 m m ) . F, fruit before elongation (scale = 10 m m ) . G, fruit after elongation 'scale = 10 m m ) . H , areole (scale = 1 m m ) . Prepared by D.C. Zappi after d r a w i n g s by the author.

weakly coated with wax (Barthlott in litt.)\ ribs dissolved into 5 x 5 mm, nipple-shaped podaria, which have a narrowly lanceolate, 1.0 x 0.7 mm, leaflike structure at apex below the areole, both in juvenile and adult stages; areoles with white wool, consisting of hyaline, slightly twisted hairs to 6 mm, Spines 9-13, not differentiated into radials and centrals, thin, needlelike, spreading, brittle, translucent white with a yellowish, bulbous base, 9 x c. 0.15 mm (at midpoint). Flowers up to six together at stem apex, opening simultaneously, c. 20 x 15 mm, narrowly funnelshaped, actinomorphic; inner

121

Kattermann:

Eriosyce

perianth-segments whitish yellow at base, sulphur yellow above, with an irregularly toothed, undulate tip; outer perianth-segments strongly curved outwards, abaxially with a rusty violet midstripe on the outermost segments, this broadening to occupy the whole of the inner segments; outermost segments or transitional bract-scales with a strongly papillate epidermis and a sharply porrect, mucronate, hyaline tip, covering the flower-buds prior to anthesis; axils of all perianth-segments naked, without hairs or bristles; pericarpel with a maximum diameter of c. 3.5 mm, naked or with a single narrowly lanceolate bract-scale, tube constricted at the nectar-chamber, which is c. 1.8 x 1 mm, narrowly cylindric, constricted toward base where there is some inconspicuous, probable nectar-secreting tissue (to be confirmed); remainder of tube narrow, inversely conic, c. 5 x 5 mm, c. 1.5 mm diam. at junction with nectar-chamber; stamens few, to 40, filaments 2-5 mm, white, anthers narrowly oval, c. 0.7 mm, yellow; style slender, c. 10 mm, greenish-white; stigma-lobes white, free but not spreading, c. 2.7 mm, not darkening in alcohol-formaldehyde-glycerine mixture. Fruit completly naked, red, fading to white at base, elongating to 35 mm (excl. perianth remnants) before basal dehiscence, then becoming pale pink, forming a thin-walled, balloon-shaped, winddispersed unit with the seeds occupying the apical part; funiculi with 'Arillushutchen' (Buxbaum). Seeds (see SEM Plate, facing page) up to 40 per fruit, pyriform, laterally slightly compressed, 2.0—2.2 x 1.5 x 1.2 mm; hilum lateral-subterminal, narrowly keyhole-shaped, micropyle nearly separate through a constriction; testa shiny black, slightly colliculate, cells 5—7-sided, slightly convex, cuticle weakly ruminate, nearly smooth. DISTRIBUTION. Known only from the type locality, at c. 400 m altitude, c. 5 km from the coast, south of Tocopilla, region of Antofagasta, northern Chile. The above described taxon shows a unique combination of characters, having a basallydehiscent, thin-walled fruit and at the same time perianth-segments naked in their axils, a completely naked fruit and seeds with a nearly smooth testa. Its genericplacement was therefore difficult to determine at first. Should it be Eriosyce s.l. in view of the thin-walled, nasally-dehiscent fruit, or should the fruit be looked at as having evolved in parallel for adaption to wind dispersal and in that case the complete lack of bristles and hairs on the flower and fruit and the testa structure regarded as synapomorphies with Copiapoa? The only known locality of the taxon lies at the southern edge of the range of Eriosyce subsect. Islaya on one hand, and on the other is at the northern limit of the range of Copiapoa, so biogeography gives no clues. During comparative studies of flower structures, an interesting new character was found, which seems to be specific for the genus Copiapoa. It consists of a rapid darkening of the stigma tissue when coming in contact with an alcohol-formaldehydeglycerine mixture. This consistent change of colour, from yellowish to a dark brown, is caused by an as yet uninvestigated chemical reaction. However, since chemical characters are believed to have great significance in systematics, the absence of this reaction in the new taxon from Tocopilla is a strong argument for its exclusion from Copiapoa. Such rapid darkening of the stigma in this mixture has so far not been observed in any member of Eriosyce.

122

Eriosyce laui J. Liithy S c a n n i n g electron micrographs of seed of

Kriosyce

laui J. Liithy (courtesy of Prof. D r W . Barthlott. Univ. Bonn). Above, ventral view of seed, showing the n a i r o w l y keyhole-shaped hilum (scale = 1 m m ) . Centre, testa-cells, showing the slightly r u m i n a t e cuticle (scale = 60 p m ) . Below, testa-cell periclinal wall (scale = 2 0 p m ) .

Systematic placement of the new taxon in Eriosyce requires justification in view of its aberrant flower and fruit characters. The absence of hairs and bristles in the axils of the floral bract-scales, which is in contrast to all other known taxa of Eriosyce s.l., is tentatively regarded as an autapomorphic state. Other flower characters also suggest that a process of reduction has taken place: the low number of stamens and perianthsegments and the low number of scale-bracts in the proximal part of the flower-tube, which leads to the naked fruit unique within Eriosyce. The following characters

123

Kattermann:

Erioayce

indicate the new species to be a highly derived member of the Islaya phylad: 1) the narrow nectar-chamber, 2) the very inconspicuous nectary glands, 3) the narrowly keyhole-shaped hilum, 4) the fruit adapted to wind-dispersal, 5) the habitat lying at the edge of the range of subsect. Islaya. The high systematic value of the basal fruit dehiscence as a synapomorphy of Eriosyce s.l. is supported by this interpretation. Once again, we have an example of the difficulties encountered in systematics of Cactaceae because of parallelisms in evolution through reduction of morphological structures representing a derived character state. The characters which have evolved convergently in this case are the naked axils of the perianth-segments and the nearly smooth structure of the seedcoat, both shared with Copiapoa. Another notable morphological phenomenon is the leaflike tips of the stem podaria, that in adult plants are covered by wool and, therefore, are not readily observed. It has been found in all individuals of Eriosyce laui studied so far, in both juvenile as well as adult specimens. It will need histological investigation to determine whether this structure on the lower side of the areole is a rudimentary leaf or a prolonged podarium apex, comparable with the podarium of a Neowerdermannia. Acknowledgments I am very obliged to the following persons for their valuable contributions to this article: Dr Alfred Lau (Mexico), for having entrusted to me one of his most interesting discoveries for study and description; Fred Kattermann (New Jersey, USA) for placing his unpublished manuscript at my disposal; Prof. Dr Wilhelm Barthlott (Bonn, Germany) for helping me with many SEM pictures of microstructures, of which only a few can be published here; and Dr David Hunt and Nigel Taylor (Kew) for their editorial improvements. Bibliography AX, P. (1988). Systematik in der Biologie. Gustav Fischer Verlag, Stuttgart. B R I T T O N , N.L., ROSE, J.N. (1919-23). The Cactaceae. Carnegie Institute, Washington, D.C. H O F F M A N N , A. (1989). Cactaceas en la flora silvestre de Chile. Ediciones Fundacion Claudio Gay, Santiago de Chile. H U N T , D.R., & T A Y L O R , N.P., eds (1986). The genera of the Cactaceae: towards a new consensus. In Bradleya 4: 65-78.

121

Kattermann:

Eriosyce

APPENDIX III Phylogenetic analysis of Eriosyce

Robert S. Wallace Department of Botany, Iowa State University 353 Bessey Hall, Ames, Iowa 50011 1020, U.SA. Data derived from vegetative and floral morphological characters, along with those from seed micromorphology as elucidated with scanning electron microscopy (characters described in Table 1), were encoded for cladistic analysis, as summarized in the data matrix shown in Figure 29. Character state polarities were assigned relative to the condition observed in Corryocactus brevistylus (K. Schum.) Britton & Rose, which served as the outgroup for this analysis. Other presumptively-related taxa (e.g. Notocactus s. str.) were also observed to assist in the establishment of character polarities. However, these were not included in the analysis due to several problems associated with the consistent scoring of the same characters for all potential outgroup and ingroup species. Multistats characters were coded using an hypothesized directional character transition series, which in all cases was unbranching and unidirectional. The data were subjected to cladistic parsimony analysis using the microcomputer program Phylogenetic Analysis Using Parsimony (PAUP, developed by David L. Swofford, Illinois Natural History Survey, USA), Version 3.1. The analyses were run on a Macintosh IIFX computer; both heuristic and branch and bound tree searching algorithms were used (MULPARS and TBR options were chosen). Parsimony analysis of the 34 taxa with 58 characters produced six equally parsimonious trees, each with 262 steps, and a consistency index (CI) of 0.344. The 50% majority-rule consensus tree of these six trees is shown in Figure Y. All six trees are nearly completely congruent, differing only in the placement of two clades: the clade leading to E. crispa; and, the clade comprising E. occulta—E. taltalensis, both differences relative to the remaining terminal branches (i.e. E. odieri to E. laui), whose topology remains the same in all six trees. Four of the six trees have the same topology shown for the nodes subtending these clades. The remaining portions (all taxa below E. heinrichiana—E. chilensis) of all six trees are identical in topology. The infrageneric taxonomic circumscription by the author of this revision (FK) for species groups whose inferred phylogeny (Figure 30) is hypothesized by the tree presented, defines a monophyletic section [sect. Neoporteria\ and a paraphyletic (polyphyletic?) section (sect. Eriosyce] within the genus Eriosyce. Additionally, the recognition of both monophyletic (subsects Chileosyce, Neoporteria, Eriosyce, and Islaya) and para/polyphyletic subsections (subsects Horridocactus and Pyrrhocactus) is made for pragmatic reasons.

125

Kattermann:

Eriosyce

TABLE 1. Description of characters and character state definitions for phylogenetic parsimony analysis of the genus Eriosyce. 1

S T E M E L O N G A T I O N . 0 • Columnar; higher than diameter, without apparent limit to maximum elongation; 1 = Globular; plants more or less spherical, s o m e t i m e s slightly elongated, elongation never more than 2 x plant diam.; 2 = Subglobular; plants rarely reaching spherical shape, s o m e plants may b e flattened (lush with the ground but may b e almost globular under cultivation.

2

STEM DIAMETER.

Represents average diameter of field-collected flower-bearing plants. In the miniature

plants of this g e n u s cultivation can often substantially increase the size of a plant; 0 = > 5 c m but < 2 0 cm: 1 = > 2 0 cm; 2 = < 5 cm. (It is a s s u m e d that the large globular plants evolved from the tree-like forms of Corryocactus 3

and that the small plants evolved from the large globular plants.)

S T E M F I R M N E S S . 0 = Firm; surface not easily d e p r e s s e d ; 1 = Soft; surface easily d e p r e s s e d . (Soft stem tissue has been observed in E. aspillagae.

4 5

E. crispa

and E.

villosa.)

R I B N U M B E R . 0 = < 1 3 ; 1 = > 1 3 . In many populations the number of ribs increases with a g e RIBS N O T C H E D O RD I S S O L V E D

INTO TUBERCLES

0 = Not notched; 1 = Notched; 2 = Dissolved into

elongated tubercles: 3 = Dissolved into isodiametric tubercles. 6

CHIN-LIKE PROTRUSION B E L O W AREOLE

7

AREOLES SUNKEN

0 = No chin; 1 = Chin weakly developed; 2 - Strong chin.

0 = Not sunken; the areole appears to be slightly raised to almost flat with the top of

the rib or tubercle; 1 = Sunken; spine b a s e and wool clearly below rib or tubercle. 8

AREOLE

I N D U M E N T U M : length of wool at areole. 0 = Short; wool 1 - 2 mm long; 1 = Long: wool 3 mm or

more, s o m e t i m e s to 10 mm. 9

APEX

COVER

0 = Naked; wool barely visible; 1 = Barely covered: wool covering stem epidermis; 2 =

Densely woolly: wool long floccose. 10

R A D I A L S P I N E S : T H I C K N E S S . 0 = Thin acicular; 1 = Thick acicular; stiff, often very thick, not flexible.

11

R A D I A L S P I N E : A T T I T U D E . 0 = Radiating; 1 = Upcurved; 2 = Appressed: 3 -

Pectinate.

12

R A D I A L S P I N E S T O R T U O U S ; all or s o m e radial spines strongly twisted. 0 - No; 1 = Y e s .

13

C E N T R A L S P I N E S : P R E S E N T / A B S E N T . 0 = Present; 1 = Absent.

14

C E N T R A L S P I N E S : T H I C K N E S S . 0 = Thin acicular; 1 = Thick acicular.

15

C E N T R A L S P I N E S : A T T I T U D E . 0 = Deflexed: 1 - Barely upcurved to straight porrect: 2 = Strongly upcurved

16

C E N T R A L S P I N E S T O R T U O U S : all or s o m e central spines strongly twisted. 0 • None; 1 = S o m e : 2 = Ai

17

R O O T S . 0 = Fibrous; 1 = Short-turbinate; 2 = Elongate-turbinate; 3 - Tuberous.

18

N E C K L I K E R E S T R I C T I O N where root joins plant-body. 0 = Absent; 1 и Present.

19

ROOT EPIDERMIS

0 - Firm, not easily d a m a g e d : 1 = Very soft, easily d a m a g e d ( £ krausii).

occurs easily in E. krausii

126

and E.

laui.)

(Root d a m a g e

Phylogenetic 20

FLOWER PRODUCED

FROM AREOLES

analysis

(Wallace)

0 = O n side of s t e m ; 1 = O l d a r e o l e s , in a circle a r o u n d a p e x ; 2

= Y o u n g a r e o l e s . in a circle a t a p e x . 21

NUMBER

22

WOOL

O F F L O W E R S P E RA R E O L E . 0 = One, 1 = Several

L E N G T H . 0 = S h o r t ; a f r i n g e o f w o o l is visible a t e d g e of b r a c t - s c a l e , n e v e r p r o j e c t i n g > l e n g t h of

b r a c t s c a l e , t u b e a p p e a r s ± n a k e d : 1 = L o n g ; w o o l o f t e n e x t e n d s t o n e x t h i g h e r b r a c t - s c a l e , partially o r fully obscuring tube or pericarpel epidermis. 23

P E R I C A R P E L B R I S T L E S P H E S E N T . 0 = Yes; 1 = No.

24

P E R I C A R P E L B R I S T L E N U M B E R . 0 = C l u s t e r s of 3 o r m o r e ; 1 - C l u s t e r s of 1 - 2 ;2 = 0 - 1 ; 3 = A l w a y s 0 .

25

PERICARPEL >10

26

BRISTLE

L E N G T H . M e a s u r e d from b a s e of bract-scale. 0 = Short, 5 - 1 0 m m ; 1 = Long,

mm

PERICARPEL BRISTLE SHAPE

27

PERICARPEL

28

OVARY

0 - Straight: 1 = C u r v e d or tortuous.

B R I S T L E A T T I T U D E . 0 = N o t porrect; 1 = Porrect.

LOCULE

A P E X . T h e i n s e r t i o n o f t h e s t y l e i n t o t h e t o p o f t h e o v a r y l o c u l e v a r i e s little

between

a d j o i n i n g p o p u l a t i o n s , b u t o f t e n c l e a r l y v a r i e s b e t w e e n s p e c i e s . 0 = D r a w n u p t o flat: 1 = D r a w n d o w n . 29

T U B E B R I S T L E N U M B E R . 0 = Clusters of 3 o r m o r e ; 1 = Clusters of 1 - 2 ;2 = Bristles solitary.

30

T U B E B R I S T L E L E N G T H . M e a s u r e d from b a s e of scale. 0 = Short. 5 - 1 0m m long; 1 = Long. > 1 0 m m , 2 =

31

T U B E B R I S T L E S H A P E . 0 = C u r v e d or tortuous: 1 = Straight.

Absent.

32

TUBE BRISTLE THICKNESS

33

T U B E B R I S T L E A T T I T U D E . 0 = N o t porrect: 1 = Porrect.

34

P E R I A N T H A T T I T U D E . 0 = Erect to curved outward; 1 = C u r v e d inward (see subsect

35

NECTARY

0 = T h m . f l e x i b l e : 1 = T h i c k , stiff, s p i n e - l i k e ( s u b s e c t .

Eriosyce).

Neoporteria).

S H A P E . 0 = S i m p l e ; t h e w a l l s of t h e n e c t a r y a r e p a r a l l e l t o e a c h o t h e r a n d its b a s e m a y b e

r o u n d e d o r a n g u l a r : 1 = M o d i f i e d I; t h e u p p e r p o r t i o n o f t h e n e c t a r y is n a r r o w e r t h a n t h e b a s e (this m a y b e by a t h i c k e n e d t u b e wall, n a r r o w i n g of t u b e just a b o v e t h e p e r i c a r p e l , o r b y w i d e n i n g a t t h e b a s e of t h e n e c t a r y } a n d t h e b a s e o f t h e n e c t a r y is ± f l a t t e n e d ; 2 = M o d i f i e d I I . b a s e of n e c t a r y m u c h w i d e n e d , r o u n d , almost pear-shaped, sometimes more

36

STIGMA-LOBE

ATTITUDE

angular

0 • S p r e a d i n g t o r e c u r v e d ; 1 = U p r i g h t t o s p r e a d i n g (tips of s t i g m a - l o b e s n o t

i n c u r v e d , to s p r e a d i n g less t h a n 1 0 ° from vertical); 2 = C l a s p e d 37

STIGMA-BASE

together.

I n s e v e r a l s p e c i e s t h e r e is a c l e a r e n l a r g e m e n t of t h e t o p of t h e style, w h i l e in o t h e r s t h e r e

is a s m o o t h u n i n t e r r u p t e d t r a n s i t i o n f r o m s t y l e i n t o s t i g m a - l o b e s . 0 = N o t s t e p p e d , 1 = S t e p p e d . 38

FRUIT DEHISCENCE

0 = I n d e h i s c e n t ; 1 = D e h i s c e n t b y partial circumscissile b a s a l splitting; 2 = D e h i s c e n t

by c o m p l e t e circumscissile b a s a l splitting. 39

F R U I T A B S C I S S I O N . 0 = A b s e n t ; 1 = I n c o m p l e t e ; fruit r e m a i n s a t t a c h e d t o p l a n t until r e m o v e d b y w i n d , r o d e n t s , birds etc.; 2 • C o m p l e t e , shortly after r e a c h i n g

maturity.

127

Kattermann:

Eriosyce

40

F R U I T W A L L T H I C K N E S S a t time of maturity. 0 = 2 m m o r m o r e ; 1 = 1 - 2

41

F R U I T W A L L , two w e e k s after reaching maturity. 0 m Fleshy/juicy; 1 = Dry.

42

FRUIT ELONGATION

mm; 2 = <1 m m .

0 = Not e l o n g a t i n g , or only slightly, n e v e r m o r e t h a n 1.5x d i a m . ; 1 = E l o n g a t i n g , 1 . 5 - 3 x

d i a m . ; 2 = Elongating 3 or > 3 x diam., then often enlarged, balloon-like. 43

SEED

R E T E N T I O N w i t h i n fruit. 0 = I n fruit p u l p ; 1 = L o o s e / r e s t r i c t e d ; s e e d h e l d i n s i d e fruit b y a n e t - l i k e

s t r u c t u r e w h i c h r e l e a s e s s e e d o v e r a l o n g p e r i o d ; 2 * L o o s e ; s e e d falls o u t f r e e l y u p o n fruit d e h i s c e n c e . 44

W R I N K L E S / R I D G E S O N T E S T A O F S E E D . V a r i o u s a r r a n g e m e n t s of testa-cells in rows a n d o r g r o u p s . 0 Not wrinkled; 1 = Wrinkled.

45

R I B T Y P E S . 0 = N o n e ; 1 = S i n g l e rows of testa-cells; 2 = Multiple celts in r a n d o m r o w s .

46

K E E L o n dorsal side of s e e d . 0 = A b s e n t : 1 = P r e s e n t .

47

T E S T A - C E L L S I Z E , m e a s u r e d in m m ' . 0 = L a r g e . 0 . 0 0 4 9 - 0 . 0 1 0 0 ; 1 - M e d i u m , 0 . 0 0 3 0 - 0 . 0 0 4 8 ; 2 = S m a l l .

48

A N T I C L I N A L B O U N D A R Y O F T E S T A - C E L L S . 0 = E x p o s e d ; clearly visible; 1 * C o v e r e d b y striation o r other

49

T E S T A - C E L L I N T E R S T I C E S (cell corner intersections). 0 = N o t s u n k e n or pitted; 1 = S u n k e n o r pitted.

0.0015-0.0028.

structures.

50

PERICLINAL TESTA-CELL WALL-SHAPE

0 = Moderately convex, 0.5-1x diam.; 1 = Low convex.

<0.5x

d i a m . ; 2 = Flat. 51

SECONDARY

SCULPTURE

O FTESTA-CELL

PERICLINAL WALL

g r o o v e s , pits, microropapillae etc.). 0 = A b s e n t , s m o o t h ; 1 52

DISTRIBUTION

O F SECONDARY

PERICLINAL

(the s u r f a c e relief of individual cells,

Present.

SCULPTURING

over surface of testa-cells. 0 =

Total

s u r f a c e : 1 - C e n t r e only. 3 3 - 6 7 % ; 2 = C e n t r e only, < 0 . 3 3 % of cell-surface. 53

T E R T I A R Y S C U L P T U R E O F T E S T A - C E L L P E R I C L I N A L W A L L ( a p p e n d a g e s which a r e additional to surface relief. 0 = A b s e n t ; 1 = P r e s e n t .

54

D I S T R I B U T I O N O F T E R T I A R Y P E R I C L I N A L S C U L P T U R I N G . 0 = E d g e only, 3 3 % o r less; 1 = E d g e only 3 3 - 6 7 % ; 2 = Total surface.

55

P O S I T I O N of hilum tissue. 0 = Superficial: tissue e v e n with r i mo r slightly e x s e r t e d ; 1 = I m p r e s s e d ;

hilum

deeply sunken. 56

POSITION O FMICROPYLE

R E L A T I V E T O H I L U M R I M . 0 = Flat; micropyle o n s a m e p l a n e a s hilum; 1 =

R a i s e d ; micropyle raised a b o v e hilum plane; 2 = B e n e a t h . 57 58

S H A P E O F H I L U M . 0 - Oval/ovate; 1 = Narrow oval; 2 = W i d e keyhole; 3 = Narrow

keyhole.

H I L U M R I M M O D I F I C A T I O N A T M I C R O P Y L E . 0 = N o t m o d i f i e d ; t e s t a - c e l l s e n d a b r u p t l y a t h i l u m rim b y t h e micropyle; 1 = Slightly modified: additional rim cells f o r m a n e n l a r g e d structure; 2 = Strongly modified; m a n y a d d i t i o n a l r i m c e l l s f o r m a s e m i - r i n g a r o u n d m i c r o p y l e , s o m e t i m e s a full r i n g , t h e n s e p a r a t i n g t h e from the hilum.

128

micropyle

Phylogenetic

analysis

(Wallace)

C H A R A C T E R S 5 Corryo.

1

1

2

2

3

3

0

5

0

5

0

5

brevistylus 0 1 0 0 0 bulbocalyx 11011

00000 00000 00001 10012

aerocarpa

10013

tenebrica

10003

krausii

20013

rodentiophila

12011

aurata

12011

islayensis

11011

limariensis

11011

napina

10003

crispa

11011

garaventae

11011

01001

10100

10222 11100

03102 01001

01001

10100 10101

10222 11200 00222 11200 002?0 00211

00021 10012 00100 00000

00001 0113? ?0021 00002 0102? 00001

102?0 00200 00212 12200

02001 11071 021 02??1 01111 0 0 0 01111 10071 0 0 0 12010 00071 0 2 0

10001 10012 11000 201?? 11001 01001 10001 10012

00002 0013? ??111 10001 2 0 2 1 1 01212 03102 01001 00001 00000 10211 01112 23002 01011 10101 00000 00211 01212 00002 00011 10001 00000 10211 01200

71100 12110 032 01100 02100 132 01100 11100 132 02101 12710 032

01002 01011 01002 00121

10111

11001

10001

01111

10001

01100 11100 132 00100 12100 132 01100 12110 032 00102 12100 132 02101 10110 232

aspillagae

21101

00000

marksiana

11011

10001

curvispina

11011

10000

odieri chilensis villosa subgibbosa senilis strausiana villicumensis andreaeana umadeave laui kunzei

11011

2?00?

21013

11000

21001

2100?

21012

10000

11011

10001

20013 11011 11111 01011 11011 11011 11001 11001 11011 10003 11011

8

07000 700?? 0 0 0

03102 01001

sociabilis

engleri

5

5

02012 01001 00001 00001 0113? ?0021

10000

heinrichiana

5

0

11020 00001 00111 10012

21001 00011 1000? 10012

occulta

5

10000 301?? 11010 201??

11011

esmeraldana

4

10000 00000

11011

recondita

11001

4

0 5 00000 00070 20002 10100

00001 00000

confinis

taltalensis

00000 00000 00000 00000

11000

00001

01002

00001

03002

10012

00002

10012

00002

00001

01002

00001

02002

?0???

03002

00001

03102

10012

00002

001?? 00001

1000?

00001 17211 01212 10211 01200 00011 00211 01212 00000 00111 01200 00000

00137 ??110

10000

0013? ??111

10000

0013? 77011

70001

01001

10101

01001

01001

10001

01011

10101

0013? 77110

01002 01010 00002 01011 10000 00001 1 3 1 0 ? 11001 10000 00001 0000? 1013? 10000 01002 13002 7012? 10001 10012 00002 01001 10000 10002 03002 01001 10000 00012 00002 01001 10001 10012 00002 00121 ?0??0 00001 0301? 0713? 10001 10012 00002 0113? 10000

10701

11?00 11700

20211 00200 21211 00200 ?02?1 01700 21222 11112

00001 2 1 2 1 1 01112 20211 01212 10000 ??211 00200

00000

10001

00000

10101

00000

00211

10001 00012

00211

77011

10211

00012

00211

00011 00012

00211

00001

10000

10212

00001

10000

20212

10000 00000

20212

10000 00000

00212

?????

??700

00212

??111

10000

20211

01112 01212 01200 01212

00001 00120 0 ? 0 02100 11100 132 00100 11101 0 2 1

00100 00110

131

17100 1???? ? 2 1 01100 11100 132 00100 11100 132 12100 11100 132 02100 12110 01100 11100 11100 11120 12100 12100

01212

02100 12100 12100 12120

10200

01101

10200

00001

10200

00001

10212

00001

12200

07001

01200

00100

232 232 032 032 132 032

00121 0 0 0 00121 0 0 0 00121 0 0 0 10121 0 0 0 10070 0 1 0 11100

132

FIGURE 29 D a t a m a t r i x of oh" morphological characters scored for 3 3 infrroup taxa and (or Corryocactus

hrvvistylus

which

serves as the outgroup for these analyses.

129

Kattermann:

Eriosyce laui krausii tenebrica аегосафа esmeraldana napina odieri taltalensis occulta crispa heinrichiana chilensis sociabilis villosa subgibbosa senilis aspillagae recondita confinis kunzei marksiana engleh curvispina limariensis garaventae rodentiophila aurata strausiana bulbocalyx umadeave andreaeana villicumensis islayensis

rd

Chileosyce

Horridocactus

Neoporteria

Horridocactus

Eriosyce

Pyrrhocactus

Islaya

Corryocactus brevistylus i'ii;i UK :«i T h e 50'S majority-rule tree derived from nix equally-parsimonious trees of 2 6 2 steps, C I = 0 . 3 4 4 . All trees were nearly identical with the exception of the placement of two clades: that of crispa. taltalensis

and that of occulta i

with respect to the terminal clade iislieri to laui I. Taxonomic circumscription of the ingroup into

infrageneric groups includes both monophyletic and paraphyletic sections a n d subsections. Subsection group n a m e s are shown to the right of the ingroup taxa. Section Eriosyce

= subsects. Islaya

Eriosyce.

+

130

Section Neoporteria

= subsects. Horridocactus

* Neoporteria

Chileosyce.

+ Pyrrhocactus

+

Kattermann:

Eriosyce

APPENDIX IV Documentation of accepted names and synonyms

Compiled by David Hunt, Nigel Taylor and Daniela Zappi The Herbarium, Royal Botanic Gardens Kew Richmond, Surrey TW9 3AE, U.K. The compilers have attempted to verify the author citations, places of publication and typification of all names relevant to the preceding account, including some names considered relevant by previous authors. The list does not include all names published under invalid or illegitimate generic names such as Hildmannia. The citations of specific and infraspecific names are followed by a serial number in | I for ease of reference from the index of names on pp. 170-176. The synonymy given, and the choice of neotypes, are the responsibility of Mr Kattermann and do not necessarily reflect the views of the compilers. References E G G L I , U. (1987). A Type Specimen Register of Cactaceae in Swiss Herbaria. In Trop. subtrop. Pflanzenw. 59: 1-124. KlESLING, R. (1984). Recopilacidn, en edicidn facsimilar, de todos los trahajos о refercncias sohre cactaceas publicadas por el Dr. Carlos Spegazzini. Librosur Ediciones, Buenos Aires. M u n o z PiZAKKO, C. 11960). Las especies de Plantas Descritas por R.A. Philippi en el siglo XIX. Ed. Univ. Chile. Abbreviations A r t . = Article of I C B N : G r e u t e r , W . e t ЛЬ. < 1 9 8 8 ) . International C o d e of Botanical N o m e n c l a t u r e . R e g n u m Veg.

vol.

1 1 8 . O b s . : Observationis); T : N o m e n c l a t u r a l type (with

HOLMfiREN. P . K . ET AU, Iruiex Uerbariarum G E N K R I C S Y N O N Y M S O F ERIOSYCE Brulgesia

herbarium

acronyms as

listed

by

ed. 8 ( 1 9 9 0 ) ; T L : T y p e locality.

(listed in alphabetical/date order)

Backeberg ( 1 9 3 5 ) , Blatter Kakloenforschung 1 9 3 5 : 111 1 1 9 3 5 ) . illegit. (Art. 6 4 ) . T : Cactus

villosus

Monville. Ceratistes

lattrib. to Labourot ( 1 8 5 3 ) byl Byles, Diet. G e n . S u b g e n . Cact., 8 ( 1 9 5 7 ) . O b e . : T h e n a m e s

'Ceratistes

verus

Echimx-actus

hort. Paris' and 'Ceratistes

ceratistes,

copiapen sis',

mentioned by Labouret as s y n o n y m s of

m a y h a v e been abbreviations omitting the generic n a m e

Echinocactus.

Chilenia

Backeberg & K n u t h . K a k t u s - A B C . 2 9 9 ( 1 9 3 5 ) , invalid (Art. 3 6 . 1 ) .

Chilenia

Backeberg* 1 9 3 8 1 . Rlatter K a k t e e n f o r s c h u n g 1 9 3 8 : [211 (July 1 9 3 8 ) . T : Echinocactus

?Chilenia

senilis

Backeberg. K a k l e e n k u n d e 1 9 3 9 : 8 2 ( 1 9 3 9 ) , illegit. (Arts 6 3 . 1 , 6 4 ) . T : Echinocactusjussieut

Chileniapsis

Backeberg in Backeberg & K n u t h , K a k t u s - A B C , 4 1 6 ( 1 9 3 5 ) . T : Cactus

Ckileocactus

Fric, K a k t u s a r 2 : 21 ( 1 9 3 1 ) . invalid (Art. 3 2 . 1 ) .

IChileorebutia

villosus

Philippi. Monville.

Monville.

Ritter, C a c t u s (Paris) no. 14: Suppl. 151 ( 1 9 5 9 ) , illegit. (Art. 6 3 . 1 1 . T : Echinocactus

reichei

Schumann. Chiliorebutia Delaetta

Frit, Succulenta 2 0 : 5 5 ( 1 9 3 8 1 , invalid ( A r t . 3 6 . 1 ) .

Backeberg, Die Cact. 6: 3 7 8 8 ( 1 9 6 2 ) , invalid (Art. 3 7 . 1 ) . T : Delaetia

ivoutersiana

Backeberg. invalid

(Arts 9 . 5 , 3 7 . 1 ) . Dracocactus Euporteria

Y . Ito, Bull. T a k a r a z u k a Insectarium 7 1 : 1 3 - 2 0 ( 1 9 5 0 ) , invalid (Art. 3 6 . 1 ) . K r e u z m g e r & Burning. Fedde's Rcpert. Sp. N o v . 5 0 : 2 9 9 1 1 9 4 1 ) , illegit. (Art. 6 3 . 1 1 . T :

subgibbosus

Echinocactus

Haworth.

131

Kattermann: 'tHildmannia

K r e u z i n g e r & Buining, Fedde's Repert. Sp. N o v . 5 0 : 1 9 9 ( 1 9 4 1 ) , invalid (Art. 3 7 . 1 ) . Intended

T: Echinocactus Horridocactus Islaya

Eriosyce

ehcnacanthus

Monville ex I*abouret, invalid (Art. 4 3 . 1 ) .

Backeberg. Blatter Kakteenforschung 1938(6): |7I ( 1 9 3 8 ) . T : Cactus

Backeberg, Blatter Kakteenforschung 1934< 10): [ 3 ] ( 1 9 3 4 ) . T : Islaya

INeochilenia

minor

homdus

Colla.

Backeberg.

Backeberg ex Dolz. Fedde's Repert. S p . N o v . 5 1 : 6 0 ( 3 0 April 1942). T : Echinocactus

jussieui

Monville. Neoporteria

Britton & Rose, Cact. 3: 9 4 ( 1 9 2 2 ) . T : Echinocactus

?Neoporteria INeotanahashia Wtchelia

subgibbosus

Haworth.

Backeberg, Blatter Kakteenforschung 1935(6): I 5 | ( 1 9 3 5 ) and in Backeberg & K n u t h , K a k t u s -

A B C , 2 5 8 ( 1 9 3 5 ) . illegit. (Art. 6 4 . 1 ) . T : Echinocactus

jussieui

Monville.

Y . I to, Kxpl. Diagr., 1 1 3 , 2 9 0 ( 1 9 5 7 ) . T : Echimx-actus

reichei

Schumann.

Bullock in K e w Bull. 1 9 3 8 : 2 9 7 (Oct. 1 9 3 8 ) , illegit. (Art. 6 3 . 1 ) . T : Nichelia

Bullock. O b s . : Norn. nov. fur Neoporteria

jussieui

(Monville)

Backeberg non Britton & Rose, which though illegitimate w a s

validly published (sec Art. 4 8 ) . Pyrrhtx-actus

(Berger) Berger, K a k t e e n , 3 4 5 ( 1 9 2 9 ) . L e c t o t y p e I Backeberg, Die Cact. 3: 1 5 6 3 . 1959):

Echintxactus

strausianus

S c h u m a n n . B a s i o n y m : Echinocactus

subg. Pyrrhocactus

Berger, K a k t e e n . 2 1 5

(1929). Rodentiophila Thelocephala

Backeberg, Die Cact. 3 : 1 7 9 9 ( 1 9 5 9 ) , invalid ( A r t s 3 6 , 371. Y . I to. Kxpl. Diagr.. 2 9 2 ( 1 9 5 7 ) . T : Neoporteria

парта

(Philippi! Britton & Rose.

SPECIFIC A N D LNFRASPECIFIC N A M E S l a . E r i o s y c e a u r a t a iPfeilTer) Backeberg, Cact. J. (Croydon) 5: 9 ( 1 9 3 6 ) I var. a u r a t a ) . f l ] Echinocactus

auratus

Pfeiffer, Abbild. C a c t . 2 ( 3 ) : s u b t. 14 ( 1 8 4 7 ) . T L : Chile, Bellavista (valley of the the

A c o n c a g u a river I, T : not designated, a s s u m e d to h a v e been collected by T. Bridges

(see I . M . Johnston,

Contr. G r a y H e r b . no. 8 1 : 9 8 - 1 0 6 . 1 9 2 8 ) . [ 2 | Echintxactus

sandillon

G a y . Fl. Chil. 3: 1 4 ( 1 8 4 8 ) . T L : Chile, 'cerros d e las provincias de A c o n c a g u a у

Coquimbo... h a s t a 8 6 3 7 pies'. S y n t y p e s : Gay

7 4 and Gay 3 2 2 IP, not seen-. See Kiesling, R., Tipos de

C a c t a c e a s s u d a m e r i c a n a s en herbarios extranjeros. In Bol. Soc. A r g e n t . Bol. 2413—4): 3 8 1 - 3 8 6 ( 1 9 8 6 ) . [31 Eriosyce

sandillon

( G a y ) R.A. Philippi, Descr. pi. nuev., 6 3 ( 1 8 7 2 ) . | 4 |

Eriosyce

ausseliana

Eriosyce

algarrobensis

T : Ritter Eriosyce

Ritter. Kakt. S u d a m e r . 3; 9 1 3 ( 1 9 8 0 ) . invalid (Art. 3 6 . 1 ) . Based on Ritter

ihotzkyanoe

2 5 4 ( U ) . Г51

Ritter. Kakt. S u d a m e r . 3 : 9 1 4 ( 1 9 8 0 ) . T L : Chile. H u a s c o . S of Algarrobo m i n e s , 1 9 6 9 ,

& Ruining

1 4 5 6 ( U ) . 161

Ritter. Kakt. S u d a m e r . 3: 9 1 2 ( 1 9 8 0 ) . T L : Chile, El Dorado, near O v a l l e . 1 9 5 4 . T : Ritter

2 5 3 ( Z S S ) . [7] Eriosyce

lapampaensis

Ritter. Kakt. S u d a m e r . 3: 9 1 4 ( 1 9 8 0 ) . T L : Chile, prov. A t a c a m a , La P a m p a . Rio

Conay, S E of T r a n s i t o , 1 3 0 0 - 2 4 0 0 m. 1 9 5 4 , T : Ritter 2 5 5 ( Z S S ) . | 8 | [Eriosyce Eriosyce

ceratistes ceratistes

Britton & Rose, Cact. 3: 1 8 6 ( 1 9 2 2 ) , misapplied.! [91 var. conibarbalensis

Backeberg, Descr. Cact. Nov. I: 3 3 ( 1 9 5 7 ) . T L : Chile. M a n q u e h u a .

near C o m b a r b a l a , T : not designated. 110J Eriosyce

ceratistes

var. coquimbensis

Backel>erg, Descr. Cact. Nov. 1: 3 2 11957). T L : Chile, prov. CoquimlHi.

Fray Jorge, T : not designated. ( I l l Eriosyce

wratistes

var. jorgensis

Backeberg, Descr. Cact. N o v . 1: 3 3 ( 1 9 5 7 ) . T L : C h i l e , prov. C o q u i m b o .

Estancia Fray Jorge, T : not designated. [ 121 Eriosyce

ceratistes

var. molfensis

Backeberg, Descr. Cact. N o v . 1: 3 2 ( 1 9 5 7 ) . T L : Chile, prov. C o q u i m b o , Rio

Molles, 2 3 0 0 m, T : Jiles 5 (not preserved). [ 1 3 ] [ S y n o n y m s continued on page I34J

SEM PLATE I Rows 1 - 3 : Eriosyce

aurata.

Row 1, F K - 4 4 9 . Row 2 . left, F K - 4 8 6 ; centre.

F K - 4 4 9 ; right. F K - 5 3 5 . Row 3. l e a . Xoellncr centre, F K - 4 8 6 ;

right,

FK-454.

F K - 8 0 8 . R o w 5 . E. rodentiophila,

13^

s.n. (Santiago. Rio Blanco,

Row 4. E. aurata E.F. Anderson

var.

in F K - 8 5 1 .

spinibarbis,

Kattermann: I Further s y n o n y m s of E. Eriosyce

ceratistes

T : Kraus Eriosyce

Eriosyce

aurata\

var. vallenarensis

Backeberg, Descr. Cact. N o v . 1: 3 2 (1957». T L : Chile, A t a c a m a , V a l l e n a r .

17 (not preserved). [ 1 4 ]

ceratistes

var. zorillaensis

Backeberg. Descr. Cact. Nov. 1: 3 2 11957 >. T L : Chile. O v a l l e , near Zorilla,

4 0 0 m, T : not designated. [151 Eriosyce

ceratistes

var. tranquillaensis

Ritter, K a k t . S u d a m e r . 3: 9 1 2 ( 1 9 8 0 ) , invalid (Art. 3 6 . 1 ) . T L : Chile,

iilapel. Tranquilla. [ 1 6 ] l b . E r i o s y c e a u r a t a var. s p i n i b a r b i s (Ritter) K a t t e r m a n n , supra. Eriosyi-e spinibarbis

117. f 17.]

Ritter, Kakt. S u d a m e r . 3 : 9 1 5 ( 1 9 8 0 ) . T L : C h i l e , Copiapo. W from Can-era Pinto. 1 9 5 4 .

T : Ritter 5 3 4 (U). U 8 | 2. E r i o s y c e r o d e n t i o p h i l a Ritler, Kakt. S u d a m e r . 3 : 9 1 6 ( 1 9 8 0 ) . T L : Chile, C h a n a r a l , W from S a l a d o . 1 9 5 4 . T : Ritter

2 6 4 ( U , holo., Z S S . iho.). O b e . : Priority of E. rodentiophila

Ritter ( 1 9 8 0 ) over E.

megacarpa

Ritter ( 1 9 8 0 ) w a s established by A . H o f f m a n n . Cact. Fl. Chile, 1 4 4 , 1 4 6 ( 1 9 8 9 ) ; cf. I C B N Art. 5 7 . 2 . [191 Eriosyce

rodentiophila

var. lanata Ritter. K a k t . S u d a m e r . 3 : 9 1 7 ( 1 9 8 0 ) . T L : Chile, Taltal. between Cifuncho

and E s m e r a l d a , T : Ritter 5 1 5 (Щ Rodentiophila Eriosyce

lanata

megacarpa

[20]

Ritter, Kakt. S u d a m e r . 3 : 9 1 7 ( 1 9 8 0 1 . pro syn. [211 Ritter. Kakt. S u d a m e r . 3: 9 1 7 ( 1 9 8 0 ) . T L : Chile. C h a n a r a l , 1 9 5 4 . T : Ritter 5 1 4 ( U . holo.,

Z S S , iso.). | 2 2 | Rodentiophila

megacarpa

(Ritter) Y . Ito, Cact.. 4 2 9 ( 1 9 8 1 ) . invalid (Art. 4 3 . 1 ) ; Ritter, Katalog H . W i n t e r

( 1 9 5 7 ) . invalid (Art. 3 6 . 1 . 3 7 . 1 ) . 123] Rtnientiophila

atacamensis

Ritter ex Ito. Cact., 4 2 9 ( 1 9 8 1 ) . invalid (Art. 4 3 . 1 ) . [241

3. E r i o s y c e b u l b o c a l y x ( W e r d e r m a n n ) K a t t e r m a n n , supra. Echinocactus

bulbocalyx

1 1 7 . 125]

W e r d e r m a n n . B l u h . K a k t . , t. 1 3 6 ( 1 9 3 7 ) . T L : N Argentina, 8tum*T(Ufa Backeberg

1959). L e c t o t y p e (designated here): W e r d e r m a n n , I.e.. t. 1 3 6 ( 1 9 3 7 ) . |261 Pyrrhocactus Neoporteria Pyrrhocactus Rausch Pyrrhocactus

bulbocalyx

( W e r d e r m a n n ) Backeberg. Cact. 3: 1 5 6 5 ( 1 9 5 9 ) . [ 2 7 ]

bulbocalyx megliolii

( W e r d e r m a n n ) Donald & Rowley. Cact. Succ. J. O r . Brit. 2 8 : 5 5 ( 1 9 6 6 ) . [ 2 8 | R a u s c h . K a k t . and. S u k k . 2 5 : 2 2 0 ( 1 9 7 4 ) . T L : A r g e n t i n a , San J u a n , near M a r a y e s , T :

559 (Wl. |29l umadeave

var. marayesensis

Backeberg, Descr. Cact. N o v . 3: 13 ( 1 9 6 3 ) , invalid (Art. 9 . 5 ) . T L :

A r g e n t i n a , N A r g e n t i n a , near M a r a y e s . T : Uhlig Neoporteria

umadeave

var. marayesensis

137 Inot preserved). [ 3 0 |

(Backebergl Donald & Rowley. Cact. Succ. J. O r . Brit. 2 8 : 5 8 ( 1 9 6 6 ) ,

invalid. [311 'IPyrrhocactus

dubius

Backeberg, K a k t u s - A B C , 4 1 5 ( 1 9 3 5 ) . T L : ' N W Argentina'. [Identification is tentative;

plants in cultivation s h o w this affinity - F K ] . [ 3 2 | Weoporteria

dubia

iBackeberg) Donald & Rowley, Cact. Succ. J. O r . Brit. 2 8 : 5 6 ( 1 9 6 6 ) . [ 3 3 ]

4 a . E r i o s y c e s t r a u s i a n a ( S c h u m a n n ) K a t t e r m a n n , supra. Echinocactus

strausianus

1 1 7 . |var. s t r a u s i a n a | 3 4 |

S c h u m a n n , M o n a t s s c h r . K a k t . - K u n d e 11: 1 1 2 ( 1 9 0 1 ) . T L : A r g e n t i n a , Rio N e g r o .

N of the [Rio] Colorado. T : llammerbacher

iBi"). L e c t o t y p e (designated here): S c h u m a n n , I.e. 1 0 7

(illustration). [351 Pyrrhocactus Neoporteria

strausianus strausiana

( S c h u m a n n ) Berger, K a k t e e n , 3 4 5 ( 1 9 2 9 ) . ] 3 6 | ( S c h u m a n n ) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 8 ( 1 9 6 6 ) . ( 3 7 1

I S y n o n y m s continued on page 1361

S E M P L A T E 11 Row 1, Eriosyce

umadeave,

Row 3 , E. strausiana, E. strausiana,

134

F K - 8 4 3 . Row 2 . E. andreaeana,

F K - 5 8 3 . R o w 4 , E. villicumensis,

FK-576.

FK-593

F K - 5 8 4 . Row 5.

Kattermann: IFurther s y n o n y m s of Eriosyce Echinocactus

catamarcensis

Eriosyce

strausiana]

Spegazzini, Cact. Plat. Tent. 11: 5 0 0 (19051. T L : Argentina, collinis

aridissimis pracandinis in M e n d o z a , S a n J u a n et C a t o m a r c a ' , 2 2 Jan. Spegazzini Malacocarpus

saxosis

19871. T :

LPS 2 3 1 3 0 (LPS). | 3 8 ] catamarcensis

Pyrrhocactus

1 9 0 5 |cf. Riesling

catamarcensis

(Spegazzinii Britton & Rose, Cact. 3 : 197 ( 1 9 2 2 ) . [39J I Spegazzini) Ritter, Succulenta 1959: 1 3 0 ( 1 9 5 9 ) , illegitimate n a m e (Art. 6 4 . 1 ) .

140] Echinmactus

sanjuanensis

Spegazzini, Cact. Plat. Tent. 1 1 : 5 0 1 ( 1 9 0 5 ) . T L : A r g e n t i n a , prov. S a n Juan,

'montuosis aridissimis', T : ? ( L P S , photo.). [411 Pyrrhocactus Neoporteria

sanjuanensis

(Spegazzinii Backeberg, Cactaceae ( D K G ) 1939(2): 5 4 ( 1 9 3 9 ) . [ 4 2 |

sanjuanensis

Pyrrhocactus

iSpegazzini) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 7 ( 1 9 6 6 ) . | 4 3 ]

volliaiius

Backeberg, Descr. Cact. N o v . , 3 1 ( 1 9 5 7 ) . T L : A r g e n t i n a , S A r g e n t i n a , T : not

designated, ex cult. R B . [441 Neoporteria

vnlliana

Pyrrhocactus

(Backeberg) Donald & Rowley, Cact. Succ. J. Gr. Brit. 2 8 : 5 8 < 1 9 6 6 ) . [451

setosiflorus

Backeberg, Cact. 6: 3 7 4 5 < 1 9 6 2 ) , mm Ritter. invalid ( A r t . 9 . 5 , 3 7 . 1 ) . T L : Based on

living pi. from A r g e n t i n a and Chile. ] 4 6 ] Pyrrhixactus

setiflorus

setosiflorus Neoporterui

Backeherg, Descr. Cact. Nov. 3 : 13 ( 1 9 6 3 ) . nom. nov., invalid (Art. 3 7 . 1 ) .

(P.

Backeberg поп Ritter). [471

setiflora

(Backeberg) Donald & Rowley. Cact. Succ. J. G r . Brit. 2 8 : 5 8 ( 1 9 6 6 ) , invalid (Art. 37.11.

[481 Pyrrhocactus

atrospinosus

Backeberg, Descr. Cact. N o v . 3: 13 ( 1 9 6 3 ) , invalid ( A r t 9 . 5 ) . T L : Argentina.

M e n d o z a , 4 0 k m W of M e n d o z a , T : Uhlig PyrrhiH-actus

platyacanthus

1 9 5 5 , T : Ritter

141 (not preserved). [ 4 9 ]

Ritter. Kakt. S u d a m e r . 2: 4 2 0 11980i. T L : A r g e n t i n a , prov. M e n d o z a , M a l a r g u e .

4 4 8 (ZSS).

[60]

4 b . E r i o s y c e s t r a u s i a n a var. p a c h a c o e n s i s iRauschi K a t t e r m a n n . supra, PyrrhtK-actus

pachacoensis

1 7 0 0 m, T : Rausch

5 5 6 ( Z S S (orig. cited at W » . [ 5 2 ]

5 . E r i o s y c e u m a d e a v e ( W e r d e r m a n n ! K a t t e r m a n n , supra, Echinocactus

117. | 5 1 |

R a u s c h . Kakt. and. S u k k . 2 6 : 7 3 ( 1 9 7 5 ) . T L : A r g e n t i n a , S a n J u a n , near Pacheco.

umadeave

117. [ 5 3 |

W e r d e r m a n n , Monatsschr. Deutsche K a k t . - G e s . 3 : 2 2 3 ( 1 9 3 1 ) . T L : A r g e n t i n a , prov

S a l t a , near Puerto Tastil. 2 5 0 0 - 3 0 0 0 m, T : Ritter (lost). L e c t o t y p e (designated herel: prov. S a l t a and Jujuy, Stumer Friesia

umadeave

s.n. ( B ) . f 5 4 | Fric ex Kreuzinger, Moller's D e u t s c h e Gart.-Zeit. 4 4 : 2 5 3 ( 1 9 2 9 ) , invalid (Art. 32.1.c). Based

on a photo, from Kreuzinger. | 5 5 | Pyrrhocactus Neoporteria

umadeave umadeave

( W e r d e r m a n n ) Backeberg. K a k t u s - A B C , 2 6 4 ( 1 9 3 5 ) . | 5 6 | ( W e r d e r m a n n ) Donald & Rowley. Cact. Succ. J. G r . Brit. 2 8 : 5 8 ( 1 9 6 6 ) . [ 5 7 ]

6 . E r i o a y c e a n d r e a e a n a K a t t e r m a n n , supra, Neochilenia

andreaeana

Pyrrhocactus Neoporteria

andreaeanus andreaeana

AcanthiK-alyeium

117. T : Kattermann

5 9 3 ( D B G . hob.). [58]

Backeberg, Kakt. and. S u k k . 10: 3 8 ( 1 9 5 9 ) . invalid (Art. 9 . 5 , 3 7 . 1 ) . T L : Chile, loc

unknown. Based on Andreae

s.n. (not preserved). [ 5 9 |

(Backeberg) Ritter, Succulenta 1 9 5 9 : 131 ( 1 9 5 9 ) , invalid. [601 (Backeberg! Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 5 (19661, invalid. | 6 1 ]

andreaeanum

iBackeberg) Donald, A s h i n g t o n i a 1: 1 2 5 ( 1 9 7 5 ) . invalid. | 6 2 ]

S E M P L A T E III Rows

1-4,

1 (Islaya

Eriosyce

islayensis

malendensis);

2

4 (/. chalaensis).

136

(/.

(seeds

from

divaricatiflora);

Row 5 , E. bulbocalyx,

FK-706.

commercial 3

(/.

souimr

copiapotdemz

Kattermann:

Eriosyce

7 . E r i o s y c e v i l l i c u m e n s i s I Rausch) K a t t e r m a n n , supra, Pyrrhocactus

villicumensis

Neoporteria

villicumensis

117. | 6 3 |

R a u s c h , Kakt. a n d . S u k k . 2 5 ( 1 2 ) : 2 6 8 ( 1 9 7 4 ) . T L : A r g e n t i n a , prov. San J u a n ,

Sierra Villicum, T : Rausch

5 5 5 ( Z S S . [ 6 4 ]

(Rausch (Donald, Ashingtonia 2(4): suppl. lAshingtonia Species C a t . Cact. (part 12],

without pagination] ( 1 9 7 6 ) . { 6 5 ] Pyrrhocactus

melanacanthus

Backeberg, Descr. Cact. N o v . 3: 13 ( 1 9 6 3 ) , invalid (Art. 9 . 5 ) . T L : Argentina,

prov. S a n J u a n , T : Uhtig Neoporteria

melanacantha

1 5 2 (not preserved). [ 6 6 ]

(Backeberg) D o n a l d & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 7 ( 1 9 6 6 ) . invalid. (671

8. E r i o s y c e i s l a y e n s i s (Forster) K a t t e r m a n n , supra, Echinocactus

islayensis

117. |68]

Forster, H a m b u r g . G a r t . Blumenzeit. 17: 1 6 0 ( 1 8 6 1 ) . T L : [Peru:l prov. Islay. T : not

designated. [ 6 9 | Islaya

islayensis

Neoporteria Echinocactus

(Forster) Backeberg, Blatt. Kakteenf. 1934(3): [unpaged] < 19341. [ 7 0 ]

islayensis

T: Weherhauer Islaya

(Vaupel) Backeberg. Blatt. Kakteenf. 1 9 3 5 ( 7 ) : lunpagedl ( 1 9 3 5 ) . [ 7 3 ]

islayensis

'mnllendensis'). Islaya

V a u p e l . Bot. J a h r b . 5 0 : Beibl. I l l : 2 4 ( 1 9 1 3 ) . T L : S Peru, Mollendo, 3 Oct 1 9 0 2 ,

1506 (B). | 7 2 |

malendensis

Neoporteria

(Forster) Donald & Rowley. Cact. Succ. J. G r . Brit. 2 8 : 5 6 ( 1 9 6 6 ) . f711

malendensis

f. malendensis

i V a u p e l ) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 6 11966) (as

[74]

minor Backeberg, Blatt. Kakteenf. 1934110): ] unpaged I ( 1 9 3 4 ) . T L : Peru. M o l l e n d o . 9 0 0 m. L e c t o t y p e

(designated here): Backeberg, I.e. (illustration). [751 Islaya

islayensis

Neoporteria

var. minor

islavensis

(Backeberg) Ritter, K a k t . S u d a m e r . 4: 1 3 0 0 ( 1 9 8 1 ) . [ 7 6 ]

f. minor

( R a u h & Backeberg) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 6 ( 1 9 6 6 ) .

[77] Islaya

bicolor A k c r s & Buining, Succulenta 1 9 5 1 : 3 8 ( 1 9 5 1 ) . T L : Cent. Peru, m e s a between Nazca and L a m a .

T : Akers Neoporteria Islaya

s.n. ( D S ) . [ 7 8 ] bicolor

( A k c r s & Buiningi Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 5 ( 1 9 6 6 ) . [791

brcvicylindrica

1 9 5 6 , T : Rauh Neoporteria

R a u h & Backeberg, Descr. Cact. Nov,, 3 3 11957). T L : S Peru, near C a n a m a n a , 4 0 0 m,

K . 1 3 6 (HELD?). | 8 0 |

islayensis

f. brevicytindrica

( R a u h & Backeberg) Donald & Rowley. Cact. Succ. J. G r . Brit. 2 8 :

56 (1966). [ 8 1 | Islaya

copiapoides

Rauh & Backeberg, Descr. Cact. Nov., 3 3 11957). T L : S Peru, km 5 2 V i a P a n a m e r i c a n a .

1 9 5 4 . T : Rauh Islaya

islayensis

Neoporteria

К 54 (НЕЮ?). [82|

var. copiapoides

islayensis

I Rauh & Backeberg) Ritter, Backeberg's Descr. Erbrt. [unpaged] ( 1 9 5 8 ) . [ 8 3 ]

var. copiapoides

( R a u h & Backeberg) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 6

(1966). [84] Islaya

copiapoides

T: Ritter Islaya

var. chalaensis

Ritter, Kakt. S u d a m e r . 4: 1 2 9 5 ( 1 9 8 1 ) . T L : Peru, depto A r e q u i p a , C h a l a .

1 2 8 ( U . holo., Z S S . iso.). [ 8 5 |

copiapoides

var. pseudoniollendensis

Ritter, Kakt. S u d a m e r . 4: 1 2 9 7 ( 1 9 8 1 ) . T L : Peru, depto Arequipa,

C a l a v e r i t a s . between Atico and Ocona, T : Ritter Islaya grandiflorens 1 9 5 6 . T : Rauh Neoporteria

128a ( U , holo,). 186)

R a u h & Backeberg, Descr. Cact. Nov., 3 3 ( 1 9 5 7 ) . T L : S Peru, k m 6 9 7 V i a P a n a m e r i c a n a . К

islayensis

130a ( H E I D ? ) . (87J f. grandiflorens

( R a u h & Backeberg) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 6

(1966). [88) [ S y n o n y m s continued on page 1 4 0 |

SEM PLATE Р/ Eriosyce armata

curvispina.

138

Row

1, var. curvispina.

F K - 2 8 1 . Row 3 , var. mutabtlts,

Kattermann

FK-279.

Row 2 .

w .

F K - 3 2 3 . Row 4 , var. гоЬимШ.

2 6 2 . Row 5, var. tuberisulcata,

FK-251.

Kattermann: (Further s y n o n y m s of Eriosyce

Itlaya grandiflorens

Erioayce

islayensis]

var. spinosior

R a u h & Backeberg in Backeberg, Die Cact. 3: 1 8 9 4 119591, invalid

(Art.

3 7 . 1 ) . [891 Neoporteria

islayensis

f. spinosinr

(Rauh & Backeberg) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 6

( 1 9 6 6 ) , invalid. 1901 Islaya grandiflorens

var. tenuispina

invalid (Art. 2 6 . 1 ) . T : Rauh Islaya grandis

R a u h & Backeberg in R a u h . Beitr. K e n n t n . Peruan. K a k t . . 5 0 0 ( 1 9 5 8 » ,

K. 130a ( H E I D ? ) . [ 9 1 ]

R a u h & Backeberg, Descr. Cact. Nov., 3 3 11957 (. T L : S Peru, valley of Rio M a j e s , Hacienda

O n g o r o , k m 1 8 0 . 1 0 0 0 m, 1 9 5 6 . T : Rauh Neoporteria

islayensis

var. grandis

K. 1 5 0 ( H E I D ? ) . [ 9 2 |

(Rauh & Backeberg) Donald & Rowley. C H C I . S U C C . J . G r . Brit. 2 8 : 5 6

(1966). [93] Islaya grandis

var. hrevispina

Rauh & Backeberg in R a u h . Beitr. Kenntn. Peruan. K a k t . . 4 9 4 119581. T L : S

Peru. Rio M a j e s . Hacienda Ongoro. 1 9 5 6 T : Rauh Neoporteria

islayensis

f. hrevispina

K . 150a ( Z S S ) . 1941

(Rauh & Backeberg) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 6

11966). | 9 5 ] Islayagrandis

var. neglecta

S i m o , K a k t . and. Sukk. 9(7): 103 (19581, inval. (Art. 2 6 . 1 ) . T L : S Peru, Hacienda

Ongoro, km 1 8 0 , 1 0 0 0 m, 1 9 5 6 (living pi.?), T : Rauh Islaya

paucispinosa

Rauh

&

Backeberg,

P a n a m e r i c a n a , 1 9 5 4 , T : Rauh Islaya раш-ispina

Descr.

K. 150. [ 9 6 ]

Cact. N o v .

Ill, 3 3 (1957).

T L : S Peru,

km

52 Via

K . 4 2 ( Z S S ) . 1971

R a u h & Backeberg in R a u h . Peruan. KakU?enveg., 5 0 0 ( 1 9 5 8 ) . O b s . : error for

paucispinosa.

198] Islava paucispinosa TL:

var. runispina

Peru, C h a l a , 1 9 5 6 , T : Rauh

Neoporteria

lindlevi

1. curvispina

R a u h & Backeberg in Rauh. Beitr. K e n n t n . Peruan. K a k t . . 5 0 2 ( 1 9 5 8 ) . K. 127b (HEID?). [99]

(Rauh & Backeberg) Donald & Rowlev, Cact. Succ. J. G r . Brit. 2 8 : 5 7 11966).

11001 Islaya krainziuna Neoporteria

Ritter. S u k k u l e n t c n k . 7-8: 31 11963). T L : Chile. Poconchile, T : Ritter 2 0 0 p.p. ( Z S S ) . [ 1 0 1 ]

krainziana

Islaya divarieatiflora

(Ritter) Donald & Rowley. Cact. Succ. J. G r . Brit. 2 8 : 5 7 ( 1 9 6 6 ) . [1021 Ritter. Taxon 13: 144 (19641. T L : Peru, depto Arequipa, C a n a m a n a , T : Ritter 5 8 8 ( U ,

holo., Z S S , too.). 11031 Neoporteria

islayensis

var. divarieatiflora

(Ritter) Donald & Rowley. Cact. Succ. J. G r . Brit. 2 8 : 5 6 ( 1 9 6 6 ) .

[1041 Islaya flavida Ritter. K a k t . S u d a m e r . 4 : 1 2 9 8 ( 1 9 8 1 ) . T L : Peru. prov. Caraveli. Convento, 1 9 5 3 . T : Ritter 1 8 6 ( U , holo., Z S S , iso.l. [1051 Islaya

maritime

Ritter, K a k t . S u d a m e r . 4: 1 3 0 1 ( 1 9 8 1 ) . T L : Peru. Atico, 1 9 5 6 . T : Ritter

nov. for / . grandiflorens Islaya

minuseula

1462 Islaya (U).

(U).

5 9 0 ( U l . O b a . : Sp.

R a u h & Backeberg 1 9 5 8 non 1 9 5 7 . 11061

Ritter, K a k t . S u d a m e r . 4: 1 3 0 3 ( 1 9 8 1 ) . T L : Peru, depto Arequipa. Atico, 1 9 6 9 , T : Ritter

11071

ungnispina

Ritter. K a k t . S u d a m e r . 4: 1 3 0 4 ( 1 9 8 1 ) . T L : Peru, depto Tacna, Y a r a d a , 1 9 5 6 , T : Ritter 5 9 1

1108]

Islaya omasensis

Ostolaza & Mischler, K a k t . a n d . S u k k . 3 4 ( 3 1 : 5 4 ( 1 9 8 3 ) . T L : Peru, prov. C a n e t e . 9 0 km S E

of L i m a , between Cnnyllo and O m a s , without date or collector, T : [Cactaceae] 5 4 0 7 ( M l . 11091 Neoporteria

omasensis

I O s t o l a z a & Mischler) F e r r y m a n in P r e s t o n - M a f h a m . Cact. 111. Diet., 1 4 7 ( 1 9 9 1 ) . 1110]

SEM

PLATE V

Rows 1 - 3 , Eriosyce

kunzei.

F K - 8 2 0 . 3 , var. transitensis. 5, E. confinis.

140

FK-141.

1, var. kunzei,

F K - 4 5 9 . 2 , var.

F K - 8 2 0 . R o w 4 , E. limariensis.

transitensis. FK-20. Row

Kattermann: 9 . 1 . E r i o s y c e r e c o n d i t a (Ritter) K a t t e r m a n n . supra, Pyrrhocactus

reconditus

Eriosyce 1 1 7 . [subsp. r e c o n d i t a l [1111

Ritter. Succulenta 1 9 6 2 : 2 7 ( 1 9 6 2 ) . T L : Chile, N A n t o f a g a s t a . 1 9 5 4 . T : Ritter

204

(ZSS). | 1 1 2 | Netxhilenia

remndita

Neoporteria

recondita

Pyrrhocactus T : Ritter

(Ritter) Backeberg. Descr. Cact. N o v . 3: 1 0 ( 1 9 6 3 ) . [1131 (Ritter) D o n a l d & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 7 ( 1 9 6 6 ) . [1141

vexatux

Ritter, K a k t . S u d a m e r . 3: 9 8 1 ( 1 9 8 0 ) . T L : Chile. N W A n t o f a g a s t a , Morro Moreno, 1 9 6 0 .

1 0 5 5 ( U ) . [1151

9 . 2 . E r i o s y c e r e c o n d i t a subsp. i q u i q u e n s i s (Ritteri K a t t e r m a n n , supra, Pyrrhocactus

iquiquensis

1 1 7 . [1161

Ritter, T a x o n 12: 3 2 ( 1 9 6 3 ) . T L : N Chile, Iquique, T : Ritter

201 (ZSS). [117]

Neochilenia

iquiquensis

I Ritter) Backeberg, Descr. Cact. Nov. 3: 9 1 1 9 6 3 ) . 11181

Neoporteria

iquiquensis

(Ritter) D o n a l d & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 6 ( 1 9 6 6 ) . [1191

Pyrrhocactus

aricensis

Ritter, T a x o n 12: 3 2 ( 1 9 6 3 ) . T L : N Chile, coastal region of A r i c a . T : RUter 2 6 8 ( Z S S ) .

1120] Neochilenia

aricensis

(Ritter) Backeberg, Descr. Cact. N o v . 3: 9 ( 1 9 6 3 ) . [ 1 3 1 ]

Neoporteria

aricensis

(Ritter) D o n a l d & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 5 ( 1 9 6 6 ) . [ 1 2 2 ]

Pyrrhocactus

residuus

Ritter & Buining, Taxon 12: 3 3 11963). T L : N Chile, coastal region of A n t o f a g a s a t a .

T : Ritter 2 0 3 ( U ) . [ 1 2 3 ] Nem:hUenia

residua

(Ritter & Buiningl Backeberg, Descr. Cact. N o v . 3 : 10 ( 1 9 6 3 ) . | 1 2 4 |

Neoporteria

residua

(Ritter) D o n a l d & Rowley. Cact. Succ. J. G r . Brit. 2 8 : 5 7 ( 1 9 6 6 ) . [1251

Pyrrhocactus

saxifraga

Ritter, K a k t . S u d a m e r . 3: 9 7 0 ( 1 9 8 0 ) . T L : Chile, Iquique, Guanillos. 1 9 5 8 , T : Ritter

7 1 2 < U ) . [1261 Chileorebutia

saxifraga

Ritter, C a c t u s (Parisl 14(63): Suppl. ( 1 9 5 9 ) and I.e. 15(66): 10 ( I 9 6 0 ) , invalid (Art.

36.1). [127] Neochilenia

nuda Backeberg, Cact. 6 : 3 7 7 7 1 1 9 6 2 ) , invalid ( A r t . 3 3 . 2 ) . O b s . : B a s e d on Chileorebutia

saxifraga.

1128] Reicheocactus

floribundus

Backeberg, Cact. 6: 3 8 0 2 1 1 9 6 2 ) . invalid (Art. 3 7 . 1 ) . [ 1 2 9 ]

Pyrrhocactus

floribundus

(Backeberg) Ritter, K a k t . S u d a m e r . 3 : 9 4 0 ( 1 9 8 0 1 , invalid (Art. 3 7 . 1 ) . O b s . :

B a s i o n y m invalid. [ 1 3 0 |

1 0 a . E r i o s y c e c u r v i s p i n a (Bertero ex Colla) K a t t e r m a n n , supra, Cactus

curvispinus

1 1 7 . (var. c u r v i s p i n a ! 11311

Bertero ex Colla. M e m . A c a d . Sci. Torino 3 7 : 7 5 - 7 6 , t. 1 6 . 2 ( M a y 1 8 3 3 , preprint; 1 8 3 4 ) .

Bertero, Merc. Chil. 13: 5 9 8 ( 1 8 2 9 ) , nom. nud. T L : C h i l e , C a e h a p u a l , T : Bertero ( T O , not found i. N e o t y p e (designated here): Colla, I.e. t. 1 6 . 2 . [1321 Echimxactus Malacocarpus Pyrrhocactus

curvispinus

(Bertero ex C o l l a ! G a y , Fl. Chil. 3 : 16 ( 1 8 4 8 ) . (1331

curvispinus

(Bertero ex Colla) Britton & Rose, Cact. 3: 2 0 3 ( 1 9 2 2 ) . 1134] (Bertero ex Colla I Berger, K a k t e e n . 3 4 5 1 1 9 2 9 ) . O b e . : C o m b , also m a d e by Backeberg

curvispinus

in Backeberg & K n u t h . K a k t u s - A B C , 2 6 2 (19351. | 1 3 5 | Horridixactus Neoporteria Horridocactus

curvispinus curvispina

(Bertero ex Colla) Backeberg, K a k t e e n k u n d e 1 9 4 0 : 5 1 ( 1 9 4 0 ) . [1361

(Bertero ex Colla) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 5 ( 1 9 6 6 ) . 1137]

andicola

Ritter. Succulenta 1959(7): 9 7 ( 1 9 5 9 ) . T L : Chile, prov. A c o n c a g u a . 2 0 0 0 m, T : Ritter

4 8 6 icited at Z S S but not in Eggli 1987). I ^ e c t o t y p c (designated here): Ritter, I.e. 9 8 (illustration!. 1138] Pyrrhtxactus Horridixactus

andicola

(Ritter) Ritter, Succulenta 1 9 5 9 ( 1 0 ) : 1 3 1 ( 1 9 5 9 ) . 1139]

andicola

m, T : Ritter

var. descendens

Ritter. Succulenta 1 9 5 9 1 7 1 : 9 7 ( 1 9 5 9 ) . T L : Chile, Cordillera 33"S. 1 0 0 0

4 6 8 b (cited at Z S S but not in Eggli 1 9 8 7 ) . L e c t o t y p e (designated here): Ritter, I.e. 1 0 0

(illustration). 1140] lyrrhocactus

andicola

var. descendens

(Ritter) Ritter, Succulenta 1 9 5 9 ( 1 0 ) : 131 ( 1 9 5 9 ) . 1141]

I S y n o n y m s continued on page 1 4 4 ]

S E M PLATE VI Eriosyce

crispa.

R o w 1, subsp. crispa,

F K - 6 7 . Row 3 . var. totoralensis. Row 5, var. carrizalensis,

142

F K - 7 8 . Row 2 , var.

atroviridis.

F K - 5 2 . Row 4 , var. huascensis,

FK-58.

FK-76

Kattermann: I Further s y n o n y m s of Eriosyce Horridocactus

andicola

curvispina

var. robustus

var.

Eriosyce

curvispina}

Ritter, Succulenta

1959(7): 9 7 ( 1 9 5 9 ) . T L : Chile, 111 a pel, T i l a m a , T :

Hitter 4 6 8 d (cited at Z S S but not in Eggli 19871. L e c t o t y p e (designated here): Ritter, I.e. 9 9 (illustration). 1142) Pyrrhocactus

andicola

Pyrrhocactus

grandiflorus

T: Ritter

var. robustus

I Ritter) Ritter, Succulenta 1959( 10): 131 ( 1 9 5 9 ) . [ 1 4 3 ]

Ritter. Succulenta 1 9 6 0 ( 4 ) : 4 1 ( 1 9 6 0 ) . T L : Chile, Santiago, Cerro R a m o n , 2 0 0 0 m,

4 6 9 (cited

at Z S S b u t not in

Eggli

1987». L e c t o t y p e

(designated

here): Ritter, I.e. 4 2

(illustration). [ 1 4 4 | Horridocactus Pyrrhocactus Ritter

grandiflorus

(Ritteri Backeberg. D i e Cact. 6: 3 7 9 6 (19621. 1145)

coliguayensis

Ritter. K a k t . S u d a m e r . 3 : 9 2 9 11980). T L : Chile, Valparaiso, C o l y g u a y , 1 9 6 7 , T :

1 4 3 9 ( U ) . [1461

Pyrrhocactus

pamaensis

Ritter. K a k t . S u d a m e r . 3: 9 6 2 < 1 9 8 0 ) . T L : Chile. C o m b a r b a l a , P a m a . 1 9 5 5 , T : Ritter

2 2 2 a ( U ) . 11471 1 0 b . E r i o s y c e c u r v i s p i n a var. a c o n c a g u e n s i s (Ritter) K a t t e r m a n n . supra, Pyrrhocactus

aconcaguensis

117. [148]

Ritter. Succulenta 1 9 6 0 ( 9 ) : 1 0 8 ( 1 9 6 0 ) . T L : Chile. Chacres, T : Ritter 5 4 2 (cited

at Z S S b u t not in Eggli 1987). L e c t o t y p e (designated here): Ritter, I.e. (illustration). 1149) Horridocactus Neoporteria

aconcaguensis curvispina

(Ritteri Backeberg, Die Cact. 6: 3 7 9 1 ( 1 9 6 2 ) . [1501

var. aconcaguensis

(Ritter) Donald & Rowley, Cact. Succ. J. G r . Brit.: 2 8 : 5 5 ( 1 9 6 6 ) .

11511 lyrrhocactus

horridus

Pyrrhocactus Ritter

var. aconcaguensis

aconcaguensis

var. orientalis

(Ritteri Ritter, K a k t . S u d a m e r . 3: 9 4 8 ( 1 9 8 0 ) . 11521 Ritter, Succulenta 1 9 6 0 ( 9 ) : 1 0 9 ( 1 9 8 0 ) . T L : Chile, S a n Felipe. T :

5 4 2 a (cited at Z S S but not in Eggli 1 9 8 7 ; not illustrated). 11531

Horridocactus Neoporteria Pyrrhocactus

aconcaguensis curvispina horridus

var. orientalis

f. orientalis

(Ritter) Backeberg, Die Cact. 6: 2 7 9 1 ( 1 9 6 2 ) . [1541

(Ritteri Donald & Rowley. Cact. Succ. J. G r . Brit. 2 8 : 5 5 ( 1 9 6 6 ) . 1155)

var. orientalis

(Ritter) Ritter, K a k t . S u d a m e r . 3: 9 4 9 ( 1 9 8 0 ) . ( 1 5 6 )

1 0 c . E r i o s y c e c u r v i s p i n a var. a r m a t a (Ritter) K a t t e r m a n n , supra, Pyrrhocactus

armatus

117. [ 1 5 7 |

Ritter, Succulenta 1 9 6 0 ( 9 ) : 4 9 ( I 9 6 0 ) . T L : Chile, near S a n t i a g o . T : Ritter

4 9 9 (ZSS).

1158] Horridocactus

armatus

Neoporteria

armata

Neoporteria

tuberisulcata

(Ritter) Backeberg. D i e C a c t . 6: 3 7 9 2 ( 1 9 6 2 ) . ] 1 5 9 ]

(Ritter) K r a i n z . K a t a l o g S S Z . ed. 2 . 8 6 ( 1 9 6 7 ) . [1601 var. armata

(Ritter) Donald & Rowley. Cact. Succ. J. G r . Brit. 2 8 : 5 8 ( 1 9 6 6 ) . [ 1 6 1 ]

! 0 d . E r i o s y c e c u r v i s p i n a var. c h o a p e n s i s i Ritter) K a t t e r m a n n , supra, Pyrrhocactus

choapensis

1 1 7 . [1621

Ritter, Succulenta 1 9 6 0 ( 1 2 ) : 133 1 1 9 6 0 ) . T L : Chile, Illapel, 1 9 5 4 . T : Ritter 2 3 8 ( Z S S ) .

1163] Horridocactus Neoporteria

choapensis choapensis

(Ritter) Backeberg, Die Cact. 6: 3 7 9 3 ( 1 9 6 2 ) . [ 1 6 4 ] (Ritter) D o n a l d & Rowley. Cact. Succ. J. G r . Brit. 2 8 : 5 5 ( 1 9 6 6 ) . [1651

lOe. E r i o s y c e c u r v i s p i n a tuberisulcata

var. t u b e r i s u l c a t a

Echinocactus

iJacobi) K a t t e r m a n n . supra,

w a s established by publication of Neoporteria

Rowley (see 10c) and N. tuberisulcata tuberisulcatus

var. vegasana

Horrid(x-actus

tuberisulcatus tuberisulcatus

1 1 7 . < His.

var. armata

Priority for var.

(Ritter) D o n a l d &

Jacobi. Allg. G a r t e n z . 2 4 : 1 0 8 ( 1 8 5 6 ) . T L : Chile, Valparaiso, T : not extant or

illustrated. N e o t y p e (designated here): Kattermann Pyrrhoiactus

tuberisulcata

(Ritter) Donald & Rowley isee 10g). [ 1 6 6 ]

2 5 1 ( D B G ) . J167]

(Jacobi) Berger, K a k t e e n , 3 4 5 ( 1 9 2 9 ) . [1681 (Jacobi) Ito. Cacti, 8 0 ( 1 9 5 2 ) . [1691

I S y n o n y m s continued on page 1 4 6 ]

S E M P L A T E VII Rows 1 - 2 , Eriosyce lissocarpa. garaventae.

144

marksiana.

1, var. marksiana.

F K - 2 0 6 . Row 3 , E. aspillagae, F K - 1 0 2 . Row 5 . E. engleri,

Kattermann

FK-567.

F K - 2 0 3 . 2 , var. 1 9 7 . Row 4. E

Kattermann:

Eriosyce

Index of accepted taxa TAXON

Eriosyce aerocarpa E. andreaeana E. aspillagae E. aurata var. aurata var. spinibarbis E. bulbocalyx E. chilensis var. albidiflora var. chilensis E. confinis E. crispa s u b s p . atroviridis var. atroviridis var. carrizalensis var. huascensis var. totoralensis s u b s p . crispa E. curvispina var. aconcaguensis var. armata var. choapensis var. curvispina var. mutabilis var. robusta var. tuberisulcata E. engleri E. esmeraldana E. garaventae E. heinrichiana s u b s p . heinrichiana s u b s p . intermedia var. intermedia var. setosiflora s u b s p . simulans E. islayensis E. krausii E. kunzei var. kunzei var. transitensis E. laui E. limariensis E. marksiana var. gracilis var. lissocarpa var. marksiana

174

No.

25 6 16 1 la lb 3 29 29b 29a 13 14 14.2 14.2a 14.2c 14.2b 14.2d 14.1 10 10b 10c lOd 10a lOf lOg lOe 18 24 15 19 19.1 19.2 19.2a 19.2b 19.3 8 26 11 11a lib 28 12 17 17c 17b 17a

Page

98 89 73 30 31 31 34 102 102 102 64 66 68

68 69 68 69 66

60 51 54 54 51 54 54 54 78 97 69 79 79 82

82 82 82

42 99 62 62 63 120 63 76 76 76 76

TAXON

E. napina

No.

20 lembekei 20.2 var. lembekei 20.2a var. duripulpa 20.2b s u b s p . napina 20.1 E. occulta 23 E. odieri 21 s u b s p . fulva 21.3 s u b s p . glabrescens 21.2 s u b s p . odieri 21.1 var. odieri 21.1a var. monte-amargensis . 21.1b E. recondita 9 s u b s p . iquiquensis 9.2 s u b s p . recondita 9.1 E. rodentiophila 2 E. senilis 32 s u b s p . coimasensis 32.2 s u b s p . elquiensis 32.3 s u b s p . senilis 32.1 E. sociabilis 31 E. strausiana 4 var. pachacoensis 4b var. strausiana 4a E. subgibbosa 30 s u b s p . clavata 30.2 var. clavata 30.2a var. nigrihorrida 30.2b var. vallenarensis 30.2d var. wagenknechtii . . . . 30.2c s u b s p . subgibbosa 30.1 var. subgibbosa 30.1a var. castanea 30.1c var. litoralis 30.1b E. taltalensis 22 s u b s p . echinus 22.3 var. echinus 22.3a var. floccosa 22.3b s u b s p . paucicostata 22.2 s u b s p . pilispina 22.4 s u b s p . taltalensis 22.1 var. taltalensis 22.1a var. pygmaea 22.1b E. tenebrica 27 E. umadeave 5 E. villicumensis 7 E. villosa 33 subsp.

Page

82 83

83 83 83

97 89 90 90 89

89 89 47 48 47

32 112 113 113 113

112 34 38 38 103 106

106 107 107 107 106

106 106 106 90 91

91 94 91 94 91

91 91 99 38 40 116

Kattermann: IFurther s y n o n y m s of Eriosyce Neoporteria Cactus

tuberisulcata

horridus

curvispina

var.

Eriosyce

tuberisulcata]

(Jacob.* Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 8 (19661. | 1 7 0 j

Colla, M e m . Лес. Torin. 3 7 : 7 6 t . I 7 f . I ( M a y 1 8 3 3 , preprint; 1 8 3 4 ) , nom. i l l e g . ( A r t . 6 4 . 1 ) . T L :

"rupestribus maritimiR V a l p a r a i s o , 1830", L e c t o t y p e (designated here): Bertero Echinocactus

horridus

horridus Pyrrhocactus Neoporteria

C . G a y , Fl. Chil. 3; 15 ( 1 8 4 8 ) . T L : Chile. Valparaiso. (His.

( Т О ) . Г171] nom. nov. for

Cactus

Colla. 11721 horridus horrida

( C . G a y ) Backeberg in Backeberg & K n u t h , K a k t u s - A B C , 2 6 4 11935). [1731

( C . G a y ) D. H u n t , Bradleya 5: 9 3 ( 1 9 8 7 ) . [ 1 7 4 )

lOf, E r i o s y c e c u r v i s p i n a var. m u t a b i l i s (Ritter) K a t t e r m a n n , supra, Pyrrhocactus

horridus

var. mutabilis

117. [175]

Ritter, K a k t . S u d a m e r . 3 : 9 4 6 11980). T L : Chile, Illapel, Los Vilos, T :

Hitter 2 2 3 b ( U ) . 11761 Pyrrhocactus

odoriflorus

Ritter, Succulenta 1 9 6 0 : 1 1 6 ( I 9 6 0 ) . T L : Chile, Pichidangui, T : Ritter 4 7 0 (cited at

Z S S but not in Eggli 1 9 8 7 ) . L e c t o t y p e (designated here): Ritter, I.e. (illustration). [1771 Neochilenia

odoriflora

Neoporteria

horrida

Pyrrhocactus

(Ritter) Backeberg. Die Cact. 6: 3 7 7 8 ( 1 9 6 2 ) . [ 1 7 8 ] var. odoriflora

horridus

var. minor

(Ritter) A . H o f f m a n n , Cact. Fl. Chil. 1 9 0 ( 1 9 8 9 ) . [ 1 7 9 ] Ritter, Kakt. S u d a m e r . 3: 9 4 6 ( 1 9 8 0 ) . invalid (Art. 3 6 . 1 ) . T L : Based on

material from M o l l e s - P i c h i d a n g u i , Ritter 2 2 3 a . 11801 IOg. E r i o s y c e c u r v i s p i n a var. r o b u s t a (Ritter) K a t t e r m a n n , supra, Pyrrhocactus

robustus

1987). O b s . : Not the s a m e a s P. andicola Horridocactus

117. [181]

Ritter, Succulenta 1 9 6 0 : 6 5 ( I 9 6 0 ) . T L : Chile, Ocoa, T : Ritter

robustus

var. robustus

2 3 9 a ( Z S S ; see Eggli

(Ritter) Ritter. 1 1 8 2 ]

Ritter, W i n t e r C a t . 1 9 6 2 unpag. ( 1 9 6 2 ) , invalid ( A r t s 3 6 . 1 , 3 7 . 1 ) ; Ito, Cact., 4 1 6

( 1 9 8 1 ) , invalid (Art. 3 3 . 2 ) . [ 1 8 3 ] Pyrrhocactus Pyrrhocactus Neoporteria

horridus

var. robustus

robustus

var. vegasanus

(Ritter) Ritter, Kakt. S u d a m e r . 3 : 9 4 7 ( 1 9 8 0 ) . [ 1 8 4 ] Ritter, Succulenta 1 9 6 0 ( 6 ) : 6 5 ( 1 9 6 0 ) . T L : Chile, Quillota, L a s V e g a s ,

2 3 9 ( Z S S ; see Eggli 1 9 8 7 ) . 11851

T : Ritter

tuberisulcata

var. vegasana

(Ritter) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 8 1 1 9 6 6 ) . 11861

N a m e s of doubtful application, possibly referable to ERIOSYCE Echinocactus

ceratistes

CURVISPINA:

Otto e x Pfeiffer, E n u m . Cact., 5 1 ( 1 8 3 7 ) . T L : Chile, Bellavista, T : not designated.

[1871 Echinmactus

centeterius

L e h m a n n ex Pfeiffer, E n u m . Cact., 6 5 ( 1 8 3 7 ) . T L : [Brazil. M i n a s G e r a i s (in error;

see fig. in Pfeiffer & Otto; the plant appears to be an Eriosyce]. Gymnocalycium Pyrrhocactus Uebelmannia Neoporteria Echimx-actus

centeterium centeteria

centeteria

( L e h m a n n ex Pfeiffer) Schnabel, K a k t . a n d . S u k k . 2 2 : 164 ( 1 9 7 1 ) . [ 1 9 1 ] ( L e h m a n n ex Pfeiffer) Rowley, Excelsa 12: 3 9 ( 1 9 8 6 ) . [ 1 9 2 ]

mammillarioides

Mackie, T : Hitchin Malactxrarpus Pyrrhocactus Echinocactus

[188]

( L e h m a n n ex Pfeiffer) H o s s e u s , A p u n t c s Cact., 2 2 11926). 1189]

( L e h m a n n ex Pfeiffer) Berger, K a k t e e n , 3 4 5 ( 1 9 2 9 ) . 11901

centeterius

Hooker. Bot. M a g . 6 4 : t. 3 5 5 8 ( 1 8 3 7 ) . T L : Chile, without l o c , cult. hort.

s.n. (not preserved?!. L e c t o t y p e (designated here): Bot. M a g . t. 3 5 5 8 . [ 1 9 3 1

mammillarioides mammillarioides superlextus

(Hooker) Britton & Rose, Cact. 3: 2 0 3 ( 1 9 2 2 ) . [ 1 9 4 ] (Hookeri Backeherg in Backeberg & K n u t h , K a k t u s - A B C , 2 6 3 ( 1 9 3 5 ) . [ 1 9 5 ]

Pfeiffer. Abbild. Cact. 2(3): s u b t.

14 ( 1 8 4 7 ) . T L : Chile, A c o n c a g u a . T : not

designated. 1196] Echinocactus

soehrensii

S c h u m a n n , M o n a t s s c h r . K a k t . - K u n d e 1 1 : 7 5 ( 1 9 0 1 ) . T L : Chile. Valparaiso, without

collector's name. L e c t o t y p e (designated here): Illustration, S c h u m a n n I.e. 7 3 . [1971 Echinix-actus

frix'hlichianus

date, T : Sohrens Chileocactus Pyrrhocactus HUdmannia Horridocactus

S c h u m a n n , G e s a m t b . Nachtr., 124 ( 1 9 0 3 ) . T L : Chile, without further data or

s.n. ( B , destroyed). [ 1 9 8 ]

froehlichianus froehlichianus froehluhiana froehlichianus

( S c h u m a n n ) Kreuzinger, Verzeichnis, 3 9 ( 1 9 3 5 ) , invalid (Art. 4 3 ) . 1 1 9 9 | ( S c h u m a n n ) Backeberg in Backeberg & K n u t h , K a k t u s - A B C , 2 6 2 1 1 9 3 5 ) . 1200 ( S c h u m a n n ) K r e u z i n g e r & Buining, Feddes Repert. 5 0 : 2 0 6 ( 1 9 4 1 ) . [ 2 0 1 ] ( S c h u m a n n ) Backeberg, Die Cact. 3: 1 8 3 9 ( 1 9 5 9 ) . [ 2 0 2 ]

fFurther n a m e s of doubtful application on page 1 4 8 ]

146

S E M P L A T E VTII Rows 1 - 4 , Eriosyce setosiflora,

heinrichiana:

1, subsp. heinrichiana.

F K - 1 1 . 4, subsp. simulans,

F K - 4 6 5 . 2 , subsp. intermedia.

F K - 4 7 3 . 3 , var.

FK-82.

147

Kattermann:

Eriosyce

I Further n a m e s of doubtful application, possibly referable to Eriosyce Neoporteria

tuberisulcata

var. froehliekiana

curvispina\

( S c h u m a n n ! Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 8

(1966). [203] Horridocactus

kesselringianus

Dab., K a k t e e n k u n d e 1942( 1): 5 ( 1 9 4 2 ) . T L : Cult. hart. bot. D a r m s t a d t , without

d a t a (illustrated I.e.). ( 2 0 4 ] Neoporteria Pyrrhocactus Neoporteria

kesselringiana

(Dolz) Hutchison, Cact. Succ. J. ( U S ) 2 7 : 1 8 3 ( 1 9 5 5 ) . [ 2 0 5 ]

kesselringianus curvispina

(Dolz) Ritter, Succulenta 1 9 5 9 : 131 ( 1 9 5 9 ) . [ 2 0 6 ]

var. kesselringiana

( D o l z ) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 6 ( 1 9 6 6 ) .

[2071 1 1 a . E r i o s y c e k u n z e i (Fiirster) K a t t e r m a n n . supra, Echinocactus

kunzei

1 1 8 . |var. k u n z e i ] ( 2 0 8 ]

Forster, H a n d b . Cact., 2 9 3 ( 1 8 4 5 ) . T L : Chile, a t high altitudes; not e x t a n t or illustrated.

N e o t y p e : designated

here): Kattermann

4 5 9 ( D B G ) . O b s . : Originally spelt kunzii,

but this is not

acceptable as a n intentional Latinization of K u n z e . [ 2 0 9 ] Neoporteria lyrrhocactus Chilenia

kunzei kunzei

kunzei

Neochilenia

(Forster) Backeberg in Backeberg & K n u t h , K a k t u s - A B C , 2 6 0 ( 1 9 3 5 ) . [ 2 1 0 ]

Horridocactus

(Forster) Borg, Cacti, 2 6 2 ( 1 9 3 7 ) . ( 2 1 1 ]

(Forster) Backeberg, K a k t e e n k u n d e 1939: 8 2 ( 1 9 3 9 ) . | 2 1 2 1

kunzei

(Forster) Backeberg, F e d d e s Repert. 5 1 : 6 0 ( 1 9 4 2 ) . | 2 1 3 |

eriosyzoides

Ritter, Succulenta 1 9 5 9 : 4 9 ( 1 9 5 9 ) . T L : Chile, H u a n t a | G u a n t a | , Ritter 4 8 4 (cited

at Z S S but not in Eggli 1 9 8 7 ) . L e c t o t y p e (designated here): Ritter, I.e. 5 0 (illustration). ( 2 1 4 ] Pyrrhocactus

eriosyzoides

(Ritter) Ritter, Succulenta 1 9 5 9 ( 1 0 ) : 131 ( 1 9 5 9 ) . | 2 1 5 ]

Neochilenia

eriosyzoides

(Ritter) Backeberg, Die Cact. 6: 3 7 7 7 ( 1 9 6 2 1 . | 2 1 6 |

Neoporteria

eriosyzoides

(Ritteri Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 6 ( 1 9 6 6 ) . [2171

Pyrrhocactus

vallenarensis

Ritter, Succulenta 1 9 5 9 ( 1 2 ) : 1 5 7 ( 1 9 5 9 ) . T L : N C h i l e . V i c u n a , T : Ritter 4 8 6 < cited

at Z S S but not in Eggli 1 9 8 7 ) . L e c t o t y p e (designated here): Ritter, I.e. (illustration). [ 2 1 8 ] Horridocactus Neoporteria

vallenarensis curvispina

(Ritter) Backeberg, Die Cact. 6: 3 7 9 8 ( 1 9 6 2 ) . 12191

var. vallenarensis

(Ritter) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 5 ( 1 9 6 6 ) .

[220] l i b . E r i o s y c e k u n z e i var t r a n s i t e n s i s (Ritter) K a t t e r m a n n . supra, Pyrrhocactus

transitensis

118.12211

Ritter, T a x o n 12: 3 3 ( 1 9 6 3 ) . T L : N C h i l e , road to T r a n s i t o , T : Ritter 4 8 5 ( U ) . [ 2 2 2 ]

Neochilenia

transitensis

(Ritteri Backeberg, Descr. Cact. N o v . 3 : 10 ( 1 9 6 3 ) . [ 2 2 3 ]

Neoporteria

transitensis

(Ritter) F e r r y m a n in P r e s t o n - M a f h a m , Cact. 111. Diet., 1 4 9 ( 1 9 9 1 ) . [ 2 2 4 ]

N a m e s of doubtful application, possibly referable to ERIOSYCE Echinocactus

geissei

KUNZEI:

P o s e l g e r e x S c h u m a n n , G e s a m t b . K a k t . , 4 0 6 ( 1 8 9 8 ) . T L : C h i l e or Bolivia, neither extant

nor illustrated. O b s . : T h e description most closely fits plants collected at the upper Rio M o l l e s . 8 5 0 0 ft ( F K ) . [2251 Horridocactus Echinocactus

geissei geissei

(Poselger ex S c h u m a n n ) Dolz, Feddes Repert. 5 1 : 61 ( 1 9 4 2 ) . 12261 var. albicans

S c h u m a n n , G e s a m t b . K a k t . , 4 0 6 ( 1 8 9 8 ) . T L : N o t e x t a n t nor illustrated.

12271 Echinocactus

nidus Sohrens ex S c h u m a n n , M o n a t s s c h r . K a k t . - K u n d e 10: 1 2 2 ( 1 9 0 0 ) . T L : Chile. L e c t o t y p e

(designated here): Illustration, I.e. 1 2 3 . [ 2 2 8 ] Neoporteria

nidus

iSohrens ex S c h u m a n n ) Britton & Rose, C a c t . 3: 9 4 ( 1 9 2 2 ) . O b s . : B. & R.'s description

combines E. senilis Chilenia

nidus

4 3 . 1 ) . (2301

148

Philippi and E. nidus

Sohrens ex S c h u m a n n . 12291

(Sohrens) Backeberg, Blatt. Kakteenf. 1 9 3 8 ( 3 ) A n h a n g 13: [unpaged] 11938), invalid

(Art.

S E M P L A T E IX Rows 1 - 4 , Eriosyce

odieri.

F K - 5 2 9 . 4, subsp. glabrescens.

1, subsp. odieri.

F K - 8 0 2 . 2 , var. monte-amargensis.

F K - 5 2 2 . 3 , subsp.

fulva.

FK-53.

148

Kattermann: 1 2 . E r i o s y c e l i m a r i e n s i s (Ritter) K a t l c r m a n n . supra, Pyrrhocactus Ritter

limariensis

Eriosyce 1 1 8 . [2311

Ritter, K a k t . S u d a m e r . 3: 9 5 6 ( 1 9 8 0 ) . T L : Chile, Fray Jorge, V i a P a n a m e r i c a n a , T :

222b iV\. 1 2 3 2 !

Neoporteria Horridocactus

limariensis

(Ritter) F e r r y m a n in Preston-.Mafham, Cact. 111. Diet., 144 ( 1 9 9 1 ) . [ 2 3 3 ]

andicola

var. mollensis

Ritter, Succulenta 1959(7): 9 7 ( 1 9 5 9 ) . T L : O v a l l e , Molles, Ritter 4 6 8 e

(cited at Z S S but not in Eggli 1 9 8 7 : not illustrated). [ 2 3 4 ] Pyrrhocactus

andicola

var. mollensis

(Ritter) Ritter, Succulenta 1 9 5 9 ( 1 0 ) : 131 ( 1 9 5 9 ) . [2351

N a m e s of doubtful application, possibly referable to ERIOSYCE Echinopsis Chilenia

nigricans nigricans

Pyrrhocactus Neoporteria Echinocactus

LIMARIENSIS:

Linke, Allg. G a r t e n z . 2 5 : 2 3 9 ( 1 8 5 7 ) . T L : Chile; neither extant nor illustrated. [2361

( L i n k e ) Backeberg, K a k t e e n k u n d e 1 9 3 9 : 8 2 < 1939). [ 2 3 7 ]

nigricans

( L i n k e ) Ritter, Succulenta 1 9 5 9 : 1 3 1 ( 1 9 5 9 ) . 1238)

tuberisulcata

var. nigricans

nigricans

( L i n k e ) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 8 ( 1 9 6 6 ) . 12391

A . Dietrich ex S c h u m a n n , G e s a m t b . K a k t . , 4 2 0 ( 1 8 9 8 ) ; S c h u m a n n , B l u h e n d e K a k l .

1: t. 4 5 ( 1 9 0 3 ) . T L : C h i l e or Bolivia; not e x t a n t or illustrated. O b s . : It is not clear if this n a m e w a s b a s e d on the s a m e type a s that of Echinopsis

nigricans,

but the descriptions correspond. N e o t y p e (designated

here): S c h u m a n n , I.e.. t. 4 5 ( 1 9 0 3 ) . [2401 Horridocactus Neoporteria

nigricans nigricans

(A. Dietrich ex S c h u m a n n ) Backeberg & Dolz, Feddes Repert. 5 1 : 6 1 11942). Г2411

( A . Dietrich ex S c h u m a n n ) Britton & Rose, Cact. 3 : 9 5 ( 1 9 2 2 ) . [ 2 4 2 |

13. E r i o s y c e c o n f i n i s I Ritter) K a t t e r m a n n , supra, Pyrrhocactus

confinis

Ritter, Succulenta

118. 1243]

1961(1): 4 ( 1 9 6 1 ) . T L : N Chile, near Copiap6, M o n t e A m a r g o , T :

Ritter 4 9 4 ( U ) . [ 2 4 4 ] . Neochilenia

confinis

(Ritter) Backeberg, Descr. Cact. N o v . 3: 9 11963). [ 2 4 5 ]

Neoporteria

confinis

I Ritter) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 5 ( 1 9 6 6 ) . 12461

1 4 . 1 . E r i o s y c e c r i s p a (Ritter) K a t t e r m a n n , supra. Pyrrhocactus

crispus

1 1 8 | s u b s p . c r i s p a J. [ 2 4 7 ]

Ritter, Succulenta 1 9 5 9 ( Ш : 1 3 7 ( 1 9 5 9 ) . T L : N Chile, Freirina. T : Ritter

4 9 1 (cited at

Z S S but not in Eggli 1 9 8 7 ) . L e c t o t y p e (designated here): Ritter. I.e. 1 3 8 (illustration). | 2 4 8 | Horridocactus

crispus

(Ritter) Backeberg. Die Cact. 6: 3 7 9 5 ( 1 9 6 2 ) . | 2 4 9 |

Neoporteria

crispa

Neochilenia

nigriscaparia

(Ritter) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 5 ( 1 9 6 6 ) . [2501 Backeberg, Die Cact. 6: 3 7 8 4 ( 1 9 6 2 ) . invalid (Art. 3 7 . 1 ) . T L : C h i l e (living plant?).

[251]

1 4 . 2 a . E r i o s y c e c r i s p a subsp. and var. a t r o v i r i d i s (Ritter) K a t t e r m a n n , supra, Pyrrhxractus

atroviridis

1 1 8 . [2521

Ritter, Succulenta 1960(8): 8 9 ( I 9 6 0 ) . T L : Chile, c. 3 0 k m N W of V a l l e n a r , T : Ritter

4 7 5 (cited at Z S S but not in Eggli 1 9 8 7 ) . L e c t o t y p e (designated here): Ritter, I.e. 9 0 (illustration). [ 2 5 3 ] Horridocactus

atroviridis

Neoporteria

atroviridis

Neoporteria

tuberisulcata

(Ritter) Backeberg, Die Cact. 6: 3 7 9 3 ( 1 9 6 2 ) . [ 2 5 4 ]

(Ritter) F e r r y m a n in P r e s t o n - M a f h a m , Cact. III. Diet., 1 4 0 ( 1 9 9 1 ) . [ 2 5 5 ] var. atroviridis

(Ritter) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 8 (1966>.

1256] 1 4 . 2 b . E r i o s y c e c r i s p a var. h u a s c e n s i s (Ritter) K a t t e r m a n n , supra, Pyrrhocactus

huascensis

118. I 2-571

Ritter. Succulenta 1961(6): 5 7 ( 1 9 6 1 ) . T L : Chile. H u a s c o . T : Ritter

260 (U). |258]

Neochilenia

huascensis

(Ritter) Backeberg, Descr. Cact. N o v . 3: 9 ( 1 9 6 3 ) . 12591

Neoporteria

huascensis

(Ritter) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 6 ( 1 9 6 6 ) . [ 2 6 0 ]

1 4 . 2 c . E r i o s y c e c r i s p a var. c a r r i z a l e n s i s (Ritter) K a t t e r m a n n , supra, Pyrrh<xactus

carrizalensis

1 1 8 . [2611

Ritter, T a x o n 12: 3 3 ( 1 9 6 3 ) . T L : N Chile, prov. A t a c a m a , W of Carrizal Alto. T :

Ritter 4 9 3 ( U ) . [ 2 6 2 ] Horridocactus

carrizalensis

Neoporteria

carrizalensis

Neoporteria

totoralensis

[265|

150

(Ritter) Backeberg, Descr. Cact. N o v . 3 : 7 ( 1 9 6 3 ) . [ 2 6 3 ] (Ritter) A . E . H o f f m a n n , Cact. Chile, 1 7 8 ( 1 9 8 9 ) , incorrect n a m e (Art. 5 7 ) . [ 2 6 4 ]

var. carrizalensis

(Ritter) F e r r y m a n m l Y e s t o n - M a f h a m , Cact. 111. Diet., 1 4 9 ( 1 9 9 1 fl

SEM Р1ЛТЕ X Rows 1 - 3 , Eriosyce

napina.

1, subsp. napina.

Kattermann

7 5 . 2 , var. lembekei,

F K - 7 7 , 3 , var.

duripulpa.

FK-398.

151

Kattermann:

Eriosyce

1 4 . 2 d . E r i o s y c e c r i s p a var. t o t o r a J e n s i s (Ritteri K a t t e r m a n n , supra, Pyrrhocactus

totoralensis

Ritter. Succulenta

118. | 2 6 6 ]

1961112): 131 ( 1 9 6 1 ) . T L : Chile, prov. A t a c a m a . Totoral Bajo,

1 9 5 6 . T : Hitter 4 9 5 ( C l . 1 2 6 7 ] Neochilenia

totoralensis

(Ritter) Backeberg, Descr. Cact. N o v . 3: 10 ( 1 9 6 3 ) . 12681

Neoporteria

totoralensis

(Ritter) Donald & Rowley. Cact. Succ. J. G r . Brit. 2 8 : 5 8 ( 1 9 6 6 ) . 12691

1 5 . E r i o s y c e g a r a v e n t a e iRitter) K a t t e r m a n n , supra, Horridocactus

garaventae

118. [270J

Ritter, Succulenta 1959(4): 4 1 ( 1 9 5 9 ) (as 'garaventai').

T L : Chile Meridional, Cerro

de la С а т р а п а , T : Hitter 4 6 7 (cited at Z S S but not in Eggli 19871. L e c t o t y p e (designated here): Ritter. I.e. 4 2 (illustration). [2711 Pyrrhocactus Neoporteria Pyrrhocactus

garaventae

(Ritter) Ritter. Succulenta 1 9 5 9 ( 1 0 ) : 131 ( 1 9 5 9 ) . | 2 7 2 ]

garaventae

(Ritteri F e r r y m a n in P r e s t o n - M a f h a m . Cact. III. Diet., 1 4 3 ( 1 9 9 1 ) . [2731

subaianus

Backeberg, Die Cact. 3: 1 5 7 5 ( 1 9 5 9 ) , invalid (Art. 9 . 5 . 3 7 . 1 ) . T L : Based on [livingl

pi. from Cerro de la C a m p a n a , 2 0 0 0 m, coll. Krauw. [ 2 7 4 ] Neoporteria

subaiana

(Backeberg) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 8 ( 1 9 6 6 ) , invalid. [ 2 7 5 ]

1 6 . E r i o s y c e a s p i l l a g a e ( S o h r e n s ) K a t t e r m a n n , supra, Echinocactus

aspillagae

118. (2761

S o h r e n s . M o n a t s s c h r . D e u t s c h e K a k t . - G e s . 1: 1 2 5 ( 1 9 2 9 ) las aspillagai').

T L : Chile.

Hacienda T a n u m e , 30"S, not extant. L e c t o t y p e (designated here): Sohrens, I.e. 127 (illustration). [ 2 7 7 ] Neoporteria Pyrrhocactus

aspdlagae

( S o h r e n s ) Backeberg in Backeberg & K n u t h , K a k t u s - A B C . 2 5 9 ( 1 9 3 5 ) . [ 2 7 8 ]

aspillagae

( S o h r e n s ) Ritter, Succulenta 1959* 10): 131 ( 1 9 5 9 ) . [ 2 7 9 ]

1 7 a . E r i o s y c e m a r k s i a n a (Ritter) K a t t e r m a n n . supra. Pyrrhocactus

marksianus

1 1 8 [var. m a r k s i a n a ] . [ 2 8 0 |

Ritter, Succulenta 1960( 1): 2 ( 1 9 6 0 ) . T L : C h i l e merid., Villa Prat, S W from Curico,

T : Ritter 2 3 4 ( Z S S ) . [2811 Horridocactus Neoporteria Pyrrhocactus

marksianus marksiana

(Ritter) Backeberg, Die Cact. 6: 3 7 9 7 ( 1 9 6 2 ) . | 2 8 2 ]

(Ritter) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 7 ( 1 9 6 6 ) . [ 2 8 3 [

marksianus

var. tunensis

Ritter, Succulenta

1960(1): 2 ( 1 9 6 0 ) . T L : Chile merid.. T u n a , S a n

Fernando, T : Hitter 2 3 4 a (cited at Z S S but not in Eggli 1 9 8 7 ; not illustrated). [ 2 8 4 ] Neoporteria

marksiana

var. tunensis

(Ritter) Donald & Rowley. Cact. Succ. -). G r . Brit. 2 8 : 5 7 ( 1 9 6 6 ) . | 2 8 5 ]

1 7 b . E r i o s y c e m a r k s i a n a var. l i s s o c a r p a (Ritteri K a t t e r m a n n , supra, Pyrrhocactus

lissocarpus

118. | 2 8 6 ]

Ritter. Succulenta 196(X2): 17 ( 1 9 6 0 ) . T L : C h i l e merid., C o q u e n a , 3 4 ' 1 5 ' S , T : Ritter

4 6 6 (cited at Z S S but not in Eggli 1987). L e c t o t y p e (designated here): Ritter. I.e. 18 (illustration). [2871 Horridocactus Neoporteria Pyrrhocactus

lissocarpus curvispina

(Ritter) Backeberg. Die Cact. 6: 3 7 9 6 ( 1 9 6 2 ) . 1288]

var. lissocarpa

truncatipetalus

7 5 0 m. 1 9 6 8 , T : Ritter

(Ritteri Donald & Rowley. Cact. Succ. J. Gr. Brit. 2 8 : 5 6 ( 1 9 6 6 ) . [ 2 8 9 ]

Ritter. K a k t . S u d a m e r . 3 : 9 8 0 ( 1 9 8 0 ) . T L : Chile, S a n Fernando. RioTinguiririca. 1 4 4 8 ( U ) . (2901

1 7 c . E r i o s y c e m a r k s i a n a var. g r a c i l i s (Ritteri K a t t e r m a n n . supra. Pyrrhocactus

lissocarpus

var. gracilis

de M o s t a z a l , 34"S, T : Ritter

118. [2911

Ritter, Succulenta 196042): 17 ( I 9 6 0 » . T L : Chile. W of S a n Francisco

4 6 6 a loc. 1 (L*, holo.. seed only; S G O , iso.. fragment of ftowerl. P a r a t y p e :

Coliguay. F R 4 6 6 a loc. 2 ( S G O . f r a g m e n t of plant). | 2 9 2 ] Horridocactus

lissocarpus

var. gracilis

(Ritteri Backeberg, Die Cact. 6: 3 7 9 7 ( 1 9 6 2 ) . 12931

SEM PLATE XI Rows 1 - 5 , Eriosy<-e taltalensis. subsp. paucicostata, pilispina.

152

FK-34.

1, var. taltalensis. 3 . var. glaucescens.

F K - 7 7 2 . 5 , subsp. pygmaea

Kattermann FK-380.

(form intermedia),

5 1 1 . 2. 4.

FK-132.

var

Kattermann: 1 8 . E r i o s y c e e n g l e r i (Ritteri K a t t e r m a n n , supra, Horridocactus

engleri

Ritter. Succulenta

V a l p a r a i s o . 1 9 0 0 m, 1 9 5 4 . T : Ritter Pyrrhisaetus Neoporteria Horridocactus Neoporteria

engleri engleri

118. | 2 9 4 ]

1 9 5 9 ( 6 1 : 7 6 ( 1 9 5 9 ) . T L : Chile, Cordillera, between S a n t i a g o and

2 3 5 ( Z S S ) . 12951

(Ritter) Ritter, Succulenta 1 9 5 9 ( 1 0 ) : 131 ( 1 9 5 9 ) . [ 2 9 6 ] (Ritter) D o n a l d & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 6 ( 1 9 6 6 ) . 12971

engleri engleri

Eriosyce

var. krausii

var. krausii

Backeberg. D i e Cact. 6: 3 7 9 6 ( 1 9 6 2 ) . invalid ( A r t s 9 . 5 , 3 7 . 1 ) . [2981

I Backeberg) D o n a l d & Rowley, Cact. Succ. J. Gr. Brit. 2 8 : 5 6 ( 1 9 6 6 ) , invalid.

[2991 19.1. E r i o s y c e h e i n r i c h i a n a Horridocactus

heinrichianus

[Backeberg) K a t t e r m a n n , supra,

1 1 8 [subsp. h e i n r i c h i a n a I. 13001

Backeberg. K a k t e e n k u n d e 1 9 4 2 ( 1 ) : 8 ( 1 9 4 2 ) . T L : Chile, H u a s c o . L e c t o t y p e

(designated here): Backeberg. I.e. (illustration). 13011 Pyrrhocactus Neoporteria Pyrrhocactus

heinrichianus heinrichiana

(Backeberg) Ritter, Succulenta 1 9 5 9 ( 1 0 ) : 131 ( 1 9 5 9 ) . [302J (Backeberg) F e r r y m a n in P r e s t o n - M a f h a m . Cact. 111. Diet., 1 4 3 ( 1 9 9 1 ) . 13031

chorosensis

Ritter, Succulenta 1 9 6 0 : 1 2 1 ( 1 9 6 0 ) . T L : Chile, Rio Chorns, T : Ritter 4 8 9 (cited at

Z S S but not in Eggli 19871. L e c t o t y p e (designated here): Ritter, I.e. 1 2 2 (illustration). 13041 Neochilenia

chorosensis

(Ritter) Backeberg, D i e Cact. 6: 3 7 7 7 ( 1 9 6 2 ) . [ 3 0 5 ]

Neoporteria

chorosensis

(Ritter) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 5 ( 1 9 6 6 ) . 13061

Neochilenia

deherdtiana

Backeberg, Descr. Cact. N o v . 3: 9 ( 1 9 6 3 ) , invalid ( A r t s 9 . 5 . 3 7 . 1 ) . T L : based on living

pi. from hort. Saint-Pie. [3071 Neoporteria

deherdtiana

(Backeberg) D o n a l d & Rowley, C a c t . Succ. J. G r . Brit. 2 8 : 5 6 ( 1 9 6 6 ) , invalid. [3081

Pyrrhocactus

dcherdtianus

Pyrrhocactus

chaniarensis

(Backeberg) K a t t e r m a n n . C a c t . Succ. J. ( U S ) 5 5 : 121 ( 1 9 8 3 ) , invalid. [309J Ritter, K a k t .

S u d a m e r . 3 : 9 2 6 ( 1 9 8 0 ) . T L : Chile, La Serena,

C h a n a r a l de

A c e i t u n e s , 1 9 5 6 , T : Ritter 4 9 0 ( U ) . 1310] Pyrrhocactus

trapichensis

Ritter, K a k t . S u d a m e r . 3 : 9 7 9 ( 1 9 8 0 ) . T L : Chile, Im Serena, Trapiche, Rio Choros,

1 9 5 5 , T : Wagenkneeht Horridocactus Neochilenia

s.n. ( U ) . [3111

trapichensis trapichensis

Ritter, hort. W i n t e r C a t . , lunpagedl (19591. invalid ( A r t s 3 6 . 1 , 3 7 . 1 ) . 13121 Backeberg, Die Cact. 3: 1 8 3 4 1 1 9 5 9 ) , invalid (Art. 3 7 . 1 ) . 13131

1 9 . 2 a . E r i o s y c e h e i n r i c h i a n a subsp. a n d var. i n t e r m e d i a (Ritteri K a t t e r m a n n , supra, Pyrrhocactus

setosiflorus

var. intermedius

Ritter, Succulenta

118. [3141

1 9 6 2 ( 6 ) : 7 0 ( 1 9 6 2 ) . T L : Chile, 6 0 k m S of

Coquimbo, 1 9 5 7 , T : Д / » е г 7 0 8 a ( U ) . 13151 Neochilenia Pyrrhisaetus

setosiflora

var. intermedia

wagenknechtii

(Ritter) Backeberg, Descr. Cact. N o v . 3: 10 ( 1 9 6 3 ) . [3161

Ritter. Succulenta

196047): 8 2 ( 1 9 6 0 ) . T L : N Chile, La Serena, T : Ritter

487

(cited at Z S S b u t not in Eggli 1 9 8 7 ) . L e c t o t y p e (designated here): Ritter, I.e. 8 3 (illustration). 13171 Neochilenia Pyrrhocactus

wagenknechtii dimorphus

(Ritter) Backeberg, D i e Cact. 6: 3 7 8 3 ( 1 9 6 2 ) . 13181 Ritter. Succulenta 1 9 6 2 ( 5 ) : 3 ( 1 9 6 2 ) . T L : Chile, C o q u i m b o , 1 9 5 7 . T : flitter 7 0 7 ( U ) .

[3191 Neochilenia

dimorpha

Neoporteria

dimorpha

Pyrrhocactus T : Ritter Pyrrhocactus

jussieui

(Ritter) Backeberg. Descr. Cact. N o v . 3: 9 ( 1 9 6 3 ) . 13201 (Ritter) D o n a l d & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 6 ( 1 9 6 6 ) . [ 3 2 1 ] var. spinosior

Ritter, K a k t . S u d a m e r . 3 : 9 5 2 < 1980). T L : C h i l e , V i c u n a , Elqui-Tal, 1 9 5 5 .

2 5 2 b ( U l . 13221 jussieui

var. australis

Ritter, K a k t . S u d a m e r . 3 : 9 5 2 ( 1 9 8 0 ) . T L : Chile, O v a l l e , L a s Perdices.

1 9 6 1 . T : Ritter 2 5 2 d ( U ) . [3231

SEM

P L A T E XII

R o w s 1 - 2 , Eriosyce

krausii:

F K - 4 0 2 . Row 4 , E. laui, from Totoral. W. dung

154

1, F K - 3 9 2 ; 2 , F K - 7 9 0 . Row 3 , E.

F K - 7 7 4 . Row 5 , E. sp. (subsect.

s.n.

tenehnea. Chileosyce!

Kattermann:

Eriosyce

1 9 . 2 b . E r i o s y c e h e i n r i c h i a n a var, s e t o s i f l o r a (Ritter) K a t t e r m a n n , supra, Pyrrhocactus

setosiflorus

118, 1324)

Ritter. Succulenta 1962(6): 7 0 ( 1 9 6 2 ) . T L : Chile merid.. 30"30'S, 1 9 5 7 , T : Ritter 7 0 8

CO). [325J Neochilenia

setosiflora

(Ritter) Backeberg, Descr. Cact. N o v . 3: 10 ( 1 9 6 3 ) . 1326]

Neoporteria

setosiflora

(Ritter) Donald & Rowley, C a c t . Succ. J. G r . Brit. 2 8 : 5 8 ( 1 9 6 6 ) . Г3271

19.3. E r i o s y c e h e i n r i c h i a n a subsp. s i m u l a n s (Ritter) K a t t e r m a n n . supra. Pyrrhocactus

simulans

118. | 3 2 8 |

Ritter, Succulenta 1 9 6 1 ( 4 ) : 3 5 ( 1 9 6 1 ) . T L : N C h i l e . Valley of Choro. T : Ritter 4 8 8 (U).

[3291 Neochilenia

simulans

(Ritter) Backeberg, Descr. Cact. N o v . 3 : 1 0 ( 1 9 6 3 ) . [3301

Neoporteria

simulans

(Ritter) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 8 ( 1 9 6 6 ) . 1331]

N a m e s of doubtful application, possibly referable to ERIOSYCE Echinocactus

jussieui

HEINRICHIANA:

Monville ex S a l m - D y c k , Cact. Hort. Dyck. 1 8 4 9 : 1 7 0 ( 1 8 5 0 ) . T L : N o t extant or

illustrated. 1332] Chilenia

jussieui

(Monville ex S a l m - D y c k ) Backeberg, K a k t e e n k u n d e 1 9 3 9 : 8 2 ( 1 9 3 9 ) . [3331

Neochilenia

jussieui

(Monville ex S a l m - D y c k ) Backeberg, Feddes Repert. 5 1 : 6 0 ( 1 9 4 2 ) . 1334]

Neoporteria

jussieui

( M o n v i l l e ex S a l m - D y c k ) Britton & Rose, Cact. 3: 9 6 ( 1 9 2 2 ) . [ 3 3 5 |

Pyrrhocactus Echinocactus

jussieui

(Monville ex S a l m - D y c k ) Ritter, K a k t . S u d a m e r . 3 : 9 5 1 11980). [3361

occultus

Philippi, Fl. A t a c , 2 3 ( 1 8 6 0 ) . T L : 'In toto litoralis a Copiapo usque ad C o b r e eum

inveni', T : Philippi

( S G O 5 2 6 9 9 . not seen; cf. M u n o z

Pizarro 1 9 6 0 : 9 2 ) . O b e , D . W . W h i t e l e y t i n

A s h i n g t o n i a 3 ( 3 - 4 ) : 6 2 - 7 1 . - 1 9 7 8 ) suggested that this n a m e is to be identified with E. odieri taxa here referred to subsect. Chileosyce,

or one of the

but he had not e x a m i n e d Philippi's type, preserved at S G O . A n

herbarium specimen (at K ) collected and identified by Philippi at C o q u i m b o in 1861 is probably to be referred to Eriosyce Chilenia

occulta

heinrichiana.

For Eriosyce

occulta

K a t t e r m a n n , see sp. no. 2 3 . [3371

(Philippi) Backeberg. K a k t e e n k u n d e 1 9 3 9 : 8 2 11939). [ 3 3 8 ]

Neochilenia

wculta

(Philippi) Backeberg. Feddes Repert. 5 1 : 6 0 ( 1 9 4 2 ) . | 3 3 9 ]

Neoporteria

occulta

(Philippi) Britton & Rose, Cact. 3: 9 5 ( 1 9 2 2 ) . [ 3 4 0 |

2 0 . 1 . E r i o s y c e n a p i n a (Philippi) K a t t e r m a n n . supra, Echinocactus

napinus

118. ] 3 4 1 ] [ s u b s p . n a p i n a ]

Philippi in Regel, Gartenflora 2 1 : 129, t. 7 2 1 , fig. 1 ( M a y 1 8 7 2 ) , and in Descr. пиел

pi. 6 2 - 6 3 ( N o v e m b e r 1 8 7 2 ) . T L : Chile. 'Playas arenosas de Huasco', not e x t a n t or illustrated (cf. M u n o i Pizarro 1 9 6 0 : 9 2 ) . L e c t o t y p e (designated here): Philippi, loc. cit.. t. 7 2 1 , fig. 1. [ 3 4 2 | Neoporteria Chilenia

napina

napina

Neochilenia

(Philippi) Backelwrg. Blatt. Kakteenf. 1935(9): [ 4 ] ( 1 9 3 5 ) . f3431

(Philippi) Backeberg, K a k t e e n k u n d e 1 9 3 9 : 8 2 ( 1 9 3 9 ) . [ 3 4 4 ]

napina

(Philippi) Backeberg, Feddes Repert. 5 1 : 6 0 < 1 9 4 2 ) . [ 3 4 5 ]

Thelocephala

napina

(Philippi) Ito, Expl. Diagr., 1 4 9 ( 1 9 5 7 ) . [ 3 4 6 |

Chileorebutia

napina

(Philippi) Ritter, C a c t u s (Paris) 14(63): Suppl.: | 5 I ( 1 9 5 9 ) , incorrect n a m e (Art- 1 I A .

1347] Neoporteria

napina

var. spinosior

Backeberg, Blatt, Kakteenf. 1 9 3 5 ( 9 ) : [41 ( 1 9 3 5 ) , invalid (Art. 3 6 . 1 • TJL

Based on material from Chile, Huasco. I34HI Echinocactus

nutis Philippi, A n . U n i v . Chile 8 5 : 4 9 3 ( 1 8 9 4 ) . T L : Chile, Huasco; not e x t a n t nor illustrated -
Muiioz Pizarro 1 9 6 0 : 9 2 ) . N e o t y p e (designated here): Kattermann Neochilenia

napina

Neochilenia

mitis (Philippi) Backeberg. D i e Cact. 3: 1 8 1 8 ( 1 9 5 9 ) . [ 3 5 1 !

75 (DBG). [ 3 4 9 |

var. mitis (Philippi) Backeberg, Feddes Repert. 5 1 : 6 0 ( 1 9 4 2 ) . [ 8 6 0 ]

S E M P L A T E XI11 Row 1. Eriosyce

occulta,

3 , E. aerocarpa.

F K - 6 3 . Rows 4 - 5 , E. recondita:

subsp. iquiquensis,

156

F K - 3 9 1 . R o w 2 . E. esmeraldana.

FK-366.

FK-5&*.

4 .
Kattermann:

Eriosyce

2 0 . 2 a . E r i o s y c e n a p i n a subsp. and var. l e m b e k e i K a t t e r m a n n , supra,

1 1 8 . H o l o t y p e : Kattermann

11

( D B G ) . 13521 Neochilenia

lembekei

Backeberg, D i e Cact. 3 : 1 8 2 2 ( 1 9 5 9 ) . invalid (Art. 3 7 . 1 ) . T L : Based on living pi. from

Chile, without loc. 1 3 5 3 ] Thelocephala Neochilenia

lembekei imitans

(Backeberg) Ritter, K a k t . S u d a m e r . 3 : 1 0 0 5 ( 1 9 8 0 ) . invalid. 13541

Backeberg, Die Cact. 3: 1821 ( 1 9 5 9 ) . invalid (Art. 3 7 . 1 ) . T L : B a s e d on living material

from C h i l e , without loc. [ 3 5 5 ] Neochilenia

neoreichei

Backeberg, Die Cact. 3: 1 8 2 5 ( 1 9 5 9 ) , invalid (Art. 37.11. [ 3 5 6 |

2 0 . 2 b . E r i o s y c e n a p i n a var. d u r i p u l p a (Ritteri K a t t e r m a n n . supra, Chileorebutia

duripulpa

Netx-hilenia

duripulpa

Neoporteria

reichei

Thetotephala

118. | 3 5 7 ]

Ritter, T a x o n 12: 1 2 3 1 1 9 6 3 ) . T L : Chile ( X ) . H u a s c o . T : Ritter

f. duripulpa

duripulpa

(Ritter) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 7 ( 1 9 6 6 ) . 1360]

(Ritter) Ritter, K a k t . S u d a m e r . 3: 1 0 1 0 ( 1 9 8 0 ) . | 3 6 1 |

2 1 . 1 a . E r i o s y c e o d i e r i I L e m a i r e ex S a l m - D y c k ) K a t t e r m a n n . supra, Echinocactus

odieri

Chilenia

nor

802 (DBG). [363]

odieri ( L e m a i r e ex S a l m - D y c k I Backeberg in Backeberg & K n u t h , K a k t u s - A B C . 2 6 1 1 1 9 3 5 ) . 1364]

odieri

Chileorebutia

118 [subsp. and var. o d i e r i 1. [ 3 6 2 |

L e m a i r e ex S a l m - D y c k , Cact. Hort, Dyck. 1 8 4 9 : 174 ( 1 8 5 0 ) . T L : N o t extant

illustrated. N e o t y p e (designated here): Kattermann Neoporteria

1056 (Ui. | 3 5 8 |

(Ritter) Backeberg, Descr. Cact. N o v . 3 : 9 1 1 9 6 3 ) . 13591

( L e m a i r e ex S a l m - D y c k ) Backeberg, K a k t e e n k u n d e 1 9 3 9 : 8 2 ( 1 9 3 9 ) . [ 3 6 5 | otlieri ( L e m a i r e ex S a l m - D y c k ) Ritter, C a c t u s (Paris) 14(63): Suppl. | u n p a g e d | ( 1 9 5 9 ) , incorrect

(Art. 11.3». [ 3 6 6 ] Neochilenia Thehx-ephala

odieri odieri

( L e m a i r e ex S a l m - D y c k ) Backeberg, F e d d e s Repert. 5 1 : 6 0 ( 1 9 4 2 ) . [3671 ( L e m a i r e ex S a l m - D y c k ) Ritter. K a k t . S u d a m e r . 3 : 1 0 1 2 11980). | 3 6 8 ]

21.1b. Eriosyce odieri

var. m o n t e - a m a r g e n s i s

K a t t e r m a n n , supra,

1 1 8 . H o l o t y p e : Kattermann

522

( D B G ) . [3691 Neix-hilenia

monte-amargensis

Backeberg, Descr. Cact. N o v . 3 : 9 ( 1963), invalid (Art. 9 . 5 ) . T L : B a s e d on living

material from Chile, M o n t e - A m a r g o , Uhlig & Backeberg Neoporteria

monte-amargensis

2 0 3 8 . [3701

F e r r y m a n in P r o s t o n - M a f h a m . Cact. 111. Diet.. 1 4 6 ( 1 9 9 1 1 . invalid (Art. 3 2 . 3 ' .

13711 2 1 . 2 . E r i o s y c e o d i e r i subsp. g l a b r e s c e n s (Ritter) K a t t e r m a n n , supra. Chileorebutia

glabrescens

Thelocephala Neochilenia

glabres
118. [372]

Ritter. C a c t u s (Paris) 1 5 ( 6 6 ) : 9 ( I 9 6 0 ) , incorrect n a m e (Art. 11.3). T L : Chile.

Copiapo, coastal region. T : Ritter

7 1 0 ( U ) . 13731

(Ritter) Ritter. K a k t . S u d a m e r . 3 : 1 0 0 3 ( 1 9 8 0 ) . [3741 KiHan ex Backeberg, K a k t e e n l e x i k o n , 2 8 8 ( 1 9 6 6 ) , invalid I A r t s 9 . 5 , 3 7 . 1 ) . T L : Base*

on living pi. from Chile. [3751 2 1 . 3 . E r i o s y c e o d i e r i subsp. f u l v a i Ritter) K a t t e r m a n n . supra, Chileorebutia

fulva

Copiapo, Totoral, 1 9 5 6 , T : Ritter Thelocephala

fulva

118. [3761

Ritter. C a c t u s (Paris) 15*66): 10 ( 1 9 6 0 ) , valid but incorrect n a m e (Art. 11.3>. T L : Г Ь к 5 0 0 . [ 3 7 7 ]

Ritter, K a k t . S u d a m e r . 3: 1 0 1 1 ( 1 9 8 0 ) . | 3 7 8 |

S E M PI A T E XTV R o w s 1 - 5 , Eriosyce castanea,

o, var. litoralis

158

subgibbosa:

F K - 2 0 4 ; 3 , var. clavata, FK-475.

1, var. subgibbosa.

FK-207; 1 . 1

F K - 2 7 ; 4 . var. nigrihorrida.

I

1

Kattermann:

Eriosyce

N a m e s of doubtful application, possibly referable to ERIOSYCE Echinocactus

odieri

var. mebbesii

ODIERI:

RUmpler in Forster's H a n d b . Cact. ed. 2 , 5 9 6 ( 1 8 8 6 ) . T L : N o t extant or

illuetrated. [379J Neoporteria

odieri

var. mebhcsii

Neochilenia

odieri

var. mebbesii

Neoporteria

napina

Echinocactus

f. mebbesii

reichei

(Riimpler) Backeberg in Backeberg & K n u t h . K a k t u s - A B C . 2 6 1 ( 1 9 3 5 ) . [3801 (RUmpler) Ito, Expl. Diagr. Austroech., 2 7 8 ( 1 9 5 7 ) . [381J I Riimpler) D o n a l d & Rowley, Cact. Succ. J . G r . Brit. 2 8 : 5 7 ( 1 9 6 6 ) . 1382]

S c h u m a n n . G e s a m t b . N a c h t r . . 1 1 0 ( 1 9 0 3 ) . T L : C h i l e , without loc.. T : Reiche

s.n. ( B ) .

[383| Malacocarpus Notocactus

reichei

Neoporteria Chilenia

reichei

( S c h u m a n n ) Berger, K a k t e e n , 2 4 3 1 1 9 2 9 ) . [3851

reichei

reichei

Neochilenia

( S c h u m a n n ) Britton & Rose, Cact. 3 : 1 9 1 ( 1 9 2 2 ) . 13841

( S c h u m a n n I Backeberg in Backeberg & K n u t h , K a k t u s - A B C , 2 1 6 1 1 9 3 5 ) . (3861

( S c h u m a n n ) Backeberg, K a k t e e n k u n d e 1 9 3 9 : 8 2 ( 1 9 3 9 ) . 1387]

reichei

( S c h u m a n n ) Backeberg, Feddes Repert. 5 1 : 6 0 ( 1 9 4 2 ) . | 3 8 8 |

Chileorebutia

reichei

( S c h u m a n n ) Ritter, C a c t u s (Paris) 14(64): suppl. [5] ( 1 9 5 9 ) . [3891

Thelocephala

reichei

( S c h u m a n n ) Ritter, K a k t . S u d a m e r . 3 : 1 0 2 3 ( 1 9 8 0 ) . [ 3 9 0 ]

Neochilenia

pseudoreichei

Lembcke & Backeberg, Die Cact. 3: 1 8 2 7 . t. 1 5 9 upper ( 1 9 5 9 ) , invalid (Art. 3 7 . 1 ) .

[391] Neochilenia

atra Backeberg, Descr. Cact. N o v . 3: 9 ( 1 9 6 3 ) , invalid (Art. 9 . 5 ) . T L : B a s e d on living pi. from

Chile, without loc. in hort. Marnier-Lapostolle M 1 2 1 / 1 . [ 3 9 2 ] 2 2 . 1 a . E r i o s y c e t a l t a l e n s i s (Hutchison) K a t t e r m a n n . supra, Neoporteria

taltalensis

1 1 9 [subsp. and var. t a l t a l e n s i s ] [3931

Hutchison, Cact. Succ. J . ( U S ) 2 7 ( 6 ) : 181 i 1 9 5 5 ) . T L : C h i l e , prov. A n t o f a g a s t a , T a l - T a l ,

Sierra EBmeralda, 3 m i N of Planta E s m e r a l d a , T : Hutchison Neochilenia

taltalensis

420 (UC). [394]

(Hutchison) Backeberg, Die Cact. 3: 1 8 1 2 ( 1 9 5 9 ) . ( 3 9 5 ]

Pyrrhocactus

taltalensis

Weochitenia

neofusca

(Hutchison) Ritter. Succulenta 1 9 5 9 : 131 < 1959). 13961 Backeberg, D i e Cact. 3: 1 8 1 0 ( 1 9 5 9 ) , invalid ( A r t s 9 . 5 , 3 7 . 1 ) . T L : B a s e d on living

material. O b s . : Identification tentative ( F K ) . [3971 Pyrrhocactus Ritter

rupicola

Ritter, T a x o n 12: 3 2 ( 1 9 6 3 ) . T L : Chile ( N ) , Alorro, S W of C a l d e r a and N of Paposo, T :

2 1 3 ( Z S S ) . (3981

Neochilenia

rupiiola

Neoporteria

rupicola

Pyrrhocactus

tenuis

(Ritter) Backeberg, Descr. Cact. N o v . 3 : 1 0 ( 1 9 6 3 ) . [ 3 9 9 ] (Ritter) Donald & Rowley. Cact. Succ. J . Gr. Brit. 2 8 : 5 7 ( 1 9 6 6 ) . ( 4 0 0 ] Ritter. K a k t . S u d a m e r . 3: 9 7 6 ( 1 9 8 0 ) . T L : Chile, Paposo, 1 9 6 9 , T : Ritter

2 2 . 1 b . E r i o s y c e t a l t a l e n s i s var. p y g m a e a I Ritter) K a t t e r m a n n , supra, Pyrrhocactus

pygmaeus

HorridtK-actus Neochilenia

pygmaea pilisina

Pyrrhocactus

1 1 9 . 1402]

Ritter, T a x o n 12: 3 2 ( 1 9 6 3 ) . T L : Chile. 2 0 k m N of C h a n a r a l , T : Witter 9 1 9
pygmaeus

Neoporteria

1 4 5 3 ( U ) . [4011

Ritter, W i n t e r K a t . ( 1 9 5 7 ) , invalid ( A r t . 3 6 . 1 , 3 7 . 1 ) . [ 4 0 4 ]

(Ritter) Backeberg. Descr. Cact. N o v . 3: 9 ( 1 9 6 3 ) . [ 4 0 5 ] var. pygmaea

calderanus

(Ritter) D o n a l d & Rowley, Cact. Succ. J . G r . Brit. 2 8 : 5 7 ( 1 9 6 6 ) . 14061

Ritter, Succulenta 1 9 6 1 : 13 ( 1 9 6 1 ) . T L : N Chile, C a l d e r a , T : Ritter

Neochilenia

calderana

(Ritter) Backeberg, Descr. Cact. Nov. 3 : 9 ( 1 9 6 3 ) . [ 4 0 8 ]

Neoporteria

calderana

(Ritter) Donald & Rowley. Cact. Succ. J . G r . Brit. 2 8 : 5 5 ( 1 9 6 6 ) . [ 4 0 9 ]

Pyrrhocactus

gracilis

4 9 6 (Ul. | 4 0 7 ]

Ritter, Succulenta 1 9 6 1 : 1 2 9 ( 1 9 6 1 ) . T L : N Chile. A t a c a m a , Caldera, at M o r r o Copiapo.

1 9 5 6 . T : Ritter 4 9 5 ( U ) . [ 4 1 0 ] Neochilenia Pyrrhocactus

gracilis

(Ritter) Backeberg, Descr. Cact. N o v . 3 : 9 ( 1 9 6 3 ) . [ 4 1 1 ]

scoparius

Ritter. Succulenta 1 9 6 2 : 5 U 1 9 6 2 ) . T L : Chile, prov. A t a c a m a , N of C h a n a r a l . T : Rater

1 0 8 5 (U). [412] Neochilenia

scoparta

(Ritter) Backeberg. Descr. Cact. N o v . 3 : 10 ( 1 9 6 3 ) . 14131

Neoporteria

scopaha

(Ritter) D o n a l d & Rowley. Cact. Succ. J . G r . Brit. 2 8 : 5 8 ( 1 9 6 6 ) . 1414]

Pyrrhocactus

intermedius

Ritter, T a x o n 12: 3 2 ( 1 9 6 3 ) . T L : N Chile, N оГ C h a n a r a l , T : Ritter 2 1 3 c (Ш

Neochilenia

intermedia

(Ritter) Backeberg, Descr. Cact. Nov. 3: 9 ( 1 9 6 3 ) . [4161

Neoporteria

intermedia

(Ritter) Donald & Rowley, Cact. Succ. J . G r . Brit. 2 8 : 5 6 1 1 9 6 6 ) . ( 4 1 7 ]

( S y n o n y m s continued on page 162]

1(H)

[4Щ

SEM PLATE X V Rows 1 - 2 , Eriosyce sociabilis,

subgibbosa:

1, var. wagenknechtii,

F K - 1 8 3 ; 2 , subsp. vallenarensis,

F K - 7 9 . Row 3 , E.

F K - 1 4 7 . Row 4 , E. chilensis, F K - 3 .

161

Kattermann: I Further s y n o n y m s of Eriosyce Pyrrhocactus

pulchellus

1 9 5 6 , T : Ritter Neoporteria

var. pygmaea

\

Ritter, K a k t . S u d a m e r . 3: 9 6 5 11980). T L : Chile, S of C h a n a r a l . 2 5 k m S of Barquito.

5 2 0 (U). [ 4 1 8 |

pulchella

Pyrrhocactus

taltalensis

Eriosyce

(Ritter) F e r r y m a n in P r e s t o n - M a f h a m , Cact. III. Diet., 1 4 7 ( 1 9 9 1 ) . 14191

transiens

Ritter, K a k t . S u d a m e r . 3: 9 7 7 ( 1 9 8 0 ) . T L : Chile, N of C a l d e r a , 1 9 6 5 . T : Ritter

1432

( U ) . 1420] Neoporteria

taltalensis

Neoporteria

transiens

var. transiens

(Ritter) A . E . H o f f m a n n , Cact. Fl. C h i l . . 2 1 6 ( 1 9 8 9 ) . | 4 2 1 ]

(Ritteri F e r r y m a n in P r e s t o n - M a f h a m . Cact. III. Diet., 1 4 9 ( 1 9 9 1 ) . | 4 2 2 ]

22.2. E r i o s y c e t a l t a l e n s i s subsp. p a u c i c o s t a t a (Ritter) K a t t e r m a n n , supra, Horridocactus

paucicostatus

119. 1423]

Ritter, Succulenta 1959: 1 1 3 ( 1 9 5 9 ) . T L : Chile, 2 0 km N of Paposo. T : Ritter 5 2 1

(cited at Z S S but not in Eggli 1 9 8 7 ) . L e c t o t y p e (designated here): Ritter, I.e. 114 (illustration), [ 4 8 4 ] Pyrrhocactus

paucicostatus

(Ritter) Ritter, Succulenta

1 9 5 9 : 131 ( 1 9 5 9 ) . [ 4 2 5 ]

Neochilenia

paucicostata

(Ritter) Backeberg, Die Cact. 6: 3 7 8 0 < 1 9 6 2 ) . [ 4 2 6 ]

Neoporteria

paucicostata

(Ritter) Donald & Rowley, Cact. Succ, J. G r . Brit. 2 8 : 5 7 ( 1 9 6 6 ) . [ 4 2 7 |

Horridocactus

paucicostatus

var. ciridis

Ritter, Succulenta 1 9 5 9 ( 9 ) : 1 1 3 ( 1 9 5 9 ) . T L : Chile, Paposo, T : Ritter

5 2 1 a (cited at Z S S b u t not in Eggli 1 9 8 7 ; not illustrated). [4281 Iyrrhocactus

paucicostatus

Pyrrhocactus

neohankeanus

var. viridis (Ritter) Ritter, Succulenta

1 9 5 9 ( 1 0 ) : 131 ( 1 9 5 9 ) . [ 4 2 9 ]

Ritter. K a k t . S u d a m e r . 3 : 9 5 9 ( 1 9 8 0 ) . T L : C h i l e , T a l t a l , C a c h i t a , T : Ritter

212

(U). [430] Neoporteria

neohankeana

Pyrrhixactus

(Ritter) F e r r y m a n in P r e s t o n - M a f h a m , Cact. 111. Diet.. 1 4 5 ( 1 9 9 1 1 .

neohankeanus

var. densispinus

[431]

Ritter, K a k t . S u d a m e r . 3: 9 6 0 ( 1 9 8 0 ) . T L : C h i l e , Taltal, E from

Paposo, T : Ritter 2 1 2 d
neohankeanus

Paposo, T : Ritter Pyrrhocactus

var. elongatus

Ritter, K a k t . S u d a m e r . 3 : 9 6 0 ( 1 9 8 0 ) . T L : Chile, T a l t a l , E from

212b (U). | 4 3 3 ]

neohankeanus

var. flaviflorus

Ritter. K a k t . S u d a m e r . 3 : 9 6 0 ( 1 9 8 0 ) . T L : Chile, 15 km N E T a l t a l .

T: Д ( « е г 2 1 2 а (U). [434] Delaetia

woutersiana

Backeberg, D i e Cact. 6: 3 7 8 8 ( 1 9 6 2 ) , invalid (Art. 3 7 . 1 ) . T L : based on material from

?Chile. 14351 Pyrrhocactus

neohankeanus

f. woutersianits

(Backeberg) Ritter, K a k t . S u d a m e r . 3: 9 6 0 ( 1 9 8 0 ) . invalid (Art*

9 . 5 . 3 7 . 1 ) . [4361

N a m e of doubtful application, possibly referable to ERIOSYCE Echinocactus Neoporteria Chilenia

fuscus fusca

fusca

Neochilenia Echinocactus

subsp.

PAUCICOSTATA

(Muehlenpfordt) Backeberg, K a k t e e n k u n d e 1 9 3 9 : 8 2 ( 1 9 3 9 ) . [ 4 3 9 |

fusca

( M u e h l e n p f o r d t ) Backeberg, Cact. Succ. J. ( U S ) 2 3 : 8 8 ( 1 9 5 1 ) . [4401

ehenaeanthus

Monville ex Labouret, M o n g r . Cact., 2 5 3 ( 1 8 5 3 ) , invalid (Art. 4 3 . 1 ) . T L : Chile.

not e x t a n t nor illus.. O b s . : = E. fuscus Chilenia

TALTALENSIS

M u e h l e n p f o r d l . Allg. G a r t e n z . 16: 10 ( 1 8 4 8 ) . T L : N e i t h e r e x t a n t nor illustrated. 1437

( M u e h l e n p f o r d t ) Britton & Rose, Cact. 3: 9 9 ( 1 9 2 2 ) . [ 4 3 8 |

ebenacantha

Muehlenpfordt. [4411

(Monville ex Labouret) Backeberg, K a k t e e n k u n d e 1 9 3 9 : 8 2 ( 1 9 3 9 ) , invalid (Art 3 6 1

1442] Echinocactus

ehenaeanthus

Echinocactus

hankeanus

var. minor

Labouret. M o n o g r . Cact., 2 5 4 ( 1 8 5 3 ) , invalid (Art. 4 3 . 1 ) . [4431

R u m p l e r in Forster's H a n d b . Cact. ed. 2 . 4 7 1 ( 1 8 8 5 ) . T L : N e i t h e r extant

illustrated. O b s . : Published in s y n o n y m y of E. humilis. hankeana

( R u m p l e r ) D o l z ex Backeberg, Feddes Repert. 5 1 : 6 0 ( 1 9 4 2 ) . [ 4 4 5 ]

Neoporteria

hankeana

( R u m p l e r ) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 6 ( 1 9 6 6 ) . [ 4 4 6 |

Echinmactus

fobeanus

mm

[444]

Neochilenia

Mieckley, M o n a t s s c h r . K a k t . - K u n d e 17: 1 8 7 ( 1 9 0 7 ) . T L : U n k n o w n ; not extunt « r

illustrated. 1447] Chilenia

fobeana

( M i e c k l e y ) Backeberg, K a k t e e n k u n d e 1 9 3 9 : 8 2 ( 1 9 3 9 ) . [ 4 4 8 ]

Neochilenia

fobeana

(Mieckley) Backeberg. Feddes Repert. 5 1 : 6 0 ( 1 9 4 2 ) . [4491

Neoporteria

jussieui

var. fobeana

( M i e c k l e y ) D o n a l d & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 7 1 1 9 6 6

«5Й

S E M Р1.ЛТЕ X V I Rows 1 - 3 , Eriosyce senilis: 4 , E. villosa,

1. var, senilis,

F K - 4 2 2 ; 2 , var. elquiensis,

F K - 4 6 2 ; 3 , var. coimasensis,

F K - 2 6 6 . Row

FK-71.

163

Kattermann:

Eriosyce

2 2 . 3 a . E r i o s y c e t a l t a l e n s i s subsp. and var. e c h i n u s I Putter) K a t t e r m a n n , supra, Pyrrhocactus

echinus

119. N 5 1 ]

Ritter, T a x o n 12: 3 3 ( 1 9 6 3 1 . T L : N Chile. A n t o f a g a s t a , Cerro Colozo, T : Ritter 5 3 7 ( U ) .

1452] Horridocactus Neoporteria

echinus echinus

Pyrrhocactus Pyrrhocactus

(Ritteri Backeberg, Descr. Cact. N o v . 3 : 7 ( 1 9 6 3 ) . (4531

(Ritteri F e r r y m a n in P r e s t o n - M a f h a m , Cact. III. Diet., 1 4 2 ( 1 9 9 1 1 . [4541

echinus

var. minor

glaucescens

Ritter, T a x o n 12: 3 3 ( 1 9 6 3 ) . T : Ritter

5 3 7 a ( U ) . 14551

Ritter, Taxon 12: 3 3 ( 1 9 6 3 ) . T L : N C h i l e . El Cobre, T : Ritter 5 3 8 ( U ) . [4561

Neochilenia

glaucescens

(Ritter) Backeberg, Descr. Cact. Nov. 3: 9 ( 1 9 6 3 ) . [ 4 5 7 )

Neoporteria

eriocephala

var. glaucescens

(Ritter) D o n a l d & Rowley. Cact. Succ. J. G r . Brit. 2 8 : 5 6 ( 1 9 6 6 ) ,

invalid (Art. 4 3 . 1 ) . 1458] 2 2 . 3 b . E r i o s y c e t a l t a l e n s i s var. f l o c c o s a i Ritter) K a t t e r m a n n , supra. Pyrrhocactus

floccosus

1 1 9 . 14591

Ritter, T a x o n 12: 3 2 ( 1 9 6 3 ) . T L : N Chile, prov. A n t o f a g a s t a . coastal region. T : Ritter

5 4 5 ( U ) . [4601 Neochilenia

floccosa

Neochilenia

eriocephala

(Ritter) Backeberg, Descr. Cact. N o v . 3: 9 ( 1 9 6 3 ) . [4611 Backeberg, Die Cact. 3: 1 8 3 0 ( 1 9 5 9 ) , invalid ( A r t s 9 . 5 . 3 7 . 1 ) . T L : B a s e d on living

material from Chile, without loc. [ 4 6 2 Г Neoporteria

eriocephala

(Backeberg) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 7 ( 1 9 6 6 ) , invalid (Arts

9 . 5 , 3 7 . 1 ) . [4631 2 2 . 4 . E r i o s y c e t a l t a l e n s i s subsp. p i l i s p i n a (Ritter) K a t t e r m a n n , supra, Pyrrhocactus

pilispinus

119. [ 4 6 4 |

Ritter. Succulenta 1 9 6 2 ( 4 ) : 4 2 ( 1 9 6 2 ) . T L : N Chile, prov. A t a c a m a , Barquito, 1 9 5 4 .

T : Ritter 2 1 7 ( Z S S ) . [ 4 6 5 ] Neochilenia

pilispina

(Ritter) Backeberg, Descr. Cact. N o v . 3 : 9 ( 1 9 6 3 ) . | 4 6 6 ]

Neoporteria

pilispina

(Ritter) D o n a l d & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 7 ( 1 9 6 6 ) . | 4 6 7 1

2 3 . E r i o s y c e o c c u l t a K a t t e r m a n n , supra, Pyrrhocactus

occultus

on Echinocactus

1 1 9 . H o l o t y p e : Kattermann

sensu S c h u m a n n ( 1 9 0 3 ) поп Philippi. [4691

2 4 . E r i o s y c e e s m e r a l d a n a (Ritter) K a t t e r m a n n , supra, Chileorebutia

3 9 1 ( D B G ) . 14681

( S c h u m a n n ) Ritter, Succulenta 1 9 5 9 : 131 ( 1 9 5 9 ) , invalid (Art. 3 6 . 1 , 3 7 . 1 ) . O b s . : B a s e d occultus

esmeraldana

119. 14701

Ritter, T a x o n 12: 1 2 3 ( 1 9 6 3 ) . T L : N Chile, E s m e r a l d a , T : Ritter 5 1 8 ( U ) . [ 4 7 1 ]

Neochilenia

esmeraldana

(Ritter) Backeberg, Descr. Cact. N o v . 3: 9 ( 1 9 6 3 ) . | 4 7 2 ]

Neoporteria

esmeraldana

(Ritter) D o n a l d & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 6 ( 1 9 6 6 ) . 1473]

Thelocephala

esmeraldana

(Ritter) Ritter, K a k t . S u d a m e r . 3: 1 0 2 1 ( 1 9 8 0 ) . [ 4 7 4 ]

2 5 . E r i o s y c e a e r o c a r p a (Ritter) K a t t e r m a n n , supra, Chileorebutia

aerocarpa

1 1 9 . 1475]

Ritter, C a c t u s (Paris) 15(66): 8 < I 9 6 0 ) , incorrect n a m e (Art. 11.3). T L : Chile, F r e i n n * .

1 9 5 6 , T : Ritter 4 9 8 ( U ) . O b s . : First invalidly published in C a c t u e (Parie) 1 4 ( 6 4 ) : Suppl. ( 1 9 5 9 ) . 1 4 7 6 ] Neochilenia

aerocarpa

(Ritter) Backeberg. Die Cact. 3: 1 8 2 6 ( 1 9 5 9 ) , invalid ( A r t . 3 7 . 1 ) . O b s . : B a s e d on R m e r

(1959). 1477] Thelocephala

aerxx-arpa

(Ritter) Ritter, K a k t . S u d a m e r . 3: 1 0 1 0 ( 1 9 8 0 ) , invalid. O b s . : B a s i o n y m cited f r e e *

first (invalid) publication. 14781 Neoporteria

reichei

Thelocephala Ritter

164

var. aerocarpa

(Ritter) F e r r y m a n in P r e s t o n - M a f h a m . Cact. III. Diet., 1 4 7 ( 1 9 9 I i .

nuda Ritter, K a k t . S u d a m e r . 3 : 1 0 0 4 ( 1 9 8 0 ) . T L : C h i l e , dept. Freirina, coastal region. 1 9 6 L T:

1 4 2 5 ( U ) . [4801

S E M PLATE XVII Pollen and epidermal features. Row I, left, Eriosyce F K - 5 7 2 , 0 . 0 5 6 7 m m . Row 2 , left, E. isiayensis

perforations, grain 0 . 0 4 5 m m ; right, E. tenebrica F K - 2 7 4 , 0 . 0 6 1 m m ; right. E. subgibbosa centre, s t o m a t a of E. subgibbosa

aurata

F K - 1 7 ; grain is 0 . 0 6 0 m m ; right, E.

{Islaya molendensis),

strausmna

showing the least n u m b e r of spinulae and

F K - 4 0 2 , 0 . 0 4 7 m m . R o w 3 , left, E. curvispina

var.

F K - 2 5 5 . Row 4 , left, vertical section of epidermis o f £ . senilis

F K - 2 5 5 ; right, s t o m a t a of E. aurata

curvispina FK-418;

FK-17.

165

Kattermann: 2 6 . E r i o s y c e k r a u s i i (Ritter) K a t t e r m a n n , supra, Chileorebutia

krausii

5 0 2
( 1 9 5 9 ) (Art. 3 7 . 1 ) ; nom. prov. for Echinocactus krausii

Thelocephala

malleolata malleolata

Neoporteria

reichei

Thelocephala

S c h u m a n n non R u m p l e r . [ 4 8 2 |

Ritter, T a x o n 12: 1 2 3 ( 1 9 6 3 ) . T L : Chile, N of C h a n a r a l . T : Ritter

5 1 7 ( U ) . (4851

(Ritter) Backeberg, Descr. Cact. Nov. 3 : 9 ( 1 9 6 3 ) . | 4 8 6 |

var. malleolata

malleolata odieri

Chileorebutia

var. mebbesii

(Ritter) Ritter, K a k t . S u d a m e r . 3: 1 0 1 7 ( 1 9 8 0 ) . 1484]

Neochilenia

Neoporteria

odieri

(Ritter) Backeberg. Die Cact. 3: 1 8 3 2 ( 1 9 5 9 ) , invalid (Art. 3 7 . 1 ) . | 4 8 3 ]

krausii

Chileorebutia

1 1 9 . [4811

Ritter, C a c t u s (Paris) 15(66): 6 - 7 with fig. 1 1 9 6 0 ) , incorrect n a m e (Art. 11.3). T L : Chile,

Copiapo, 1 9 5 6 . T : Ritter Neochilenia

Eriosyce

(Ritter) D o n a l d & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 8 ( 1 9 6 6 ) . 14871

(Ritter) Ritter, K a k t . S u d a m e r . 3: 1 0 2 0 1 1 9 8 0 ) . 14881

var. malleolata

malleolata

(Ritter) F e r r y m a n in P r e s t o n - M a f h a m , Cact. III. Diet., 1 4 6 ( 1 9 9 1 ) . [4891

var. sofitaria

Ritter, T a x o n 12: 1 2 3 1 1 9 6 3 ) . T L : Chile, S of C h a n a r a l , T : Ritter 5 1 7 a

(U). [490] Neochilenia

malleolata

Neoporteria

reichei

var. solitaria

f. solitaria

Thelocephala

malleolata

Thelocephala

longirapa

(Ritter) Backeberg, Descr. Cact. Nov. 3 : 9 ( 1 9 6 3 ) . | 4 9 1 |

(Ritter) D o n a l d & Rowley, Cact. Succ. ,1. G r . Brit. 2 8 : 5 8 ( 1 9 6 6 ) . [4921

var. solitaria

(Ritteri Ritter, K a k t . S u d a m e r . 3: 1 0 2 0 ( 1 9 8 0 ) . [493f

Ritter, K a k t . S u d a m e r . 3: 1 0 1 9 ( 1 9 8 0 ) . T L : Chile, Copiapo, 1 9 6 1 , T : Ritter

1321 (Ul.

1494]

2 7 . E r i o s y c e t e n e b r i c a t Ritteri K a t t e r m a n n , supra, Thelocephala

tenebrica

1 1 9 . [4951

Ritter, K a k t . S u d a m e r . 3 : 1 0 0 1 11980). T L : Chile, prov. A t a c a m a , 1 9 6 1 . T : Ritter 1 0 9 2

(U). [496] Thelocephala

fankhauseri

Ritter, K a k t . S u d a m e r . 3: 1 0 0 2 ( 1 9 8 0 ) . T L : C h i l e , H u a s c o , D o m e y k o , T : Ritter 1 4 5 1

(U). 1497] 2 8 . E r i o s y c e l a u i J. L u t h y , supra,

1 2 1 . [4981

2 9 a . E r i o s y c e c h i l e n s i s ( H i l d m a n n ex S c h u m a n n ! K a t t e r m a n n , supra, Echinix-actus

chilensis

Neoporteria Chilenia

chilensis

chilensis

Neix-hilenia

119. |var. c h i l e n s i s ] 1 4 9 9 1

H i l d m a n n ex S c h u m a n n . G e s a m t b . K a k t . . 4 2 3 ( 1 8 9 8 ) . T L : Chile, A n d e s , not extant

or illustrated. N e o t y p e (designated here): Kattermann

FK-3 (DBG). |500]

( H i l d m a n n ex S c h u m a n n ) Britton & Rose. Cact. 3 : 9 9 1 1 9 2 2 ) . [5011

( H i l d m a n n ex S c h u m a n n ) Backeberg. K a k t e e n k u n d e 1 9 3 9 : 8 2 ( 1 9 3 9 ) . [5021

chilensis

I H i l d m a n n ex S c h u m a n n ) Backeberg, Cact. Succ. J. ( U S ) 2 3 : 8 8 ( 1 9 5 1 ) . [5031

Pyrrhocactus

chilensis

( H i l d m a n n ex S c h u m a n n ) Ritter, Succulenta 1 9 5 9 : 131 ( 1 9 5 9 ) . [ 5 0 4 |

Echinocactus

chilensis

var. confinis

H i l d m a n n ex S c h u m a n n . G e s a m t b . K a k t . , 4 2 4 ( 1 8 9 8 ) . T L : N o t extant or

illustrated. 15051 Chilenia

chilensis

var. confinis

( H i l d m a n n ex S c h u m a n n ) Backeberg in Backeberg & K n u t h , K a k t u s - A B C .

3 0 0 ( 1 9 3 5 ) , invalid (Art. 4 3 . 1 ) . [ 5 0 8 ] Neoporteria

chilensis

var. confinis

i H i l d m a n n ex S c h u m a n n ) Ito, Expl. Diagr. Austroechinocacl., 2 1 4 ( 1 9 5 7 c

D o n a l d & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 5 1 1 9 6 6 ) . [5071 Chilenia

chilensis

var. cytindracea

Backeberg in Backeberg & K n u t h . K a k t u s - A B C , 3 0 0 ( 1 9 3 5 ) , invalid l A r t .

3 6 . 1 , 4 3 . 1 ) . [5081 Neoporteria

chilensis

var. cylindracea

chilensis

f. cylindracea

(Backeberg) Ito, Expl. Diagr. Austroechinocact., 2 1 4 ( 1 9 5 7 ) , invalid.

[5091 Neoporteria Chilenia

pseudoehiiensis

Pyrrhocactus

krausii

(Backeberg) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 5 ( 1 9 6 6 ' . [ 5 1 8 f

Backeberg. Blatt. Kakteenf. 1 9 3 8 ( 3 ) : A n h a n g 13 ( 1 9 3 8 ) , invalid (Art. 4 3 . 1 ) .

Ritter. K a k t . S u d a m e r . 3 : 9 5 3 11980). T L : Loc. u n k n o w n , prob. coastal Chile, T :

1 4 5 0 ( U ) . O b s . : N o t Chileorebutia

krausii

chilensis

var. albidiflorus

T : Ritter 5 5 9 ( U l . 15141

166

Rtmr

Ritter. 15121

2 9 b . E r i o s y c e c h i l e n s i s var. a l b i d i f l o r a (Ritteri K a t t e r m a n n , supra, Pyrrkixactus

(5113

1 1 9 . [5131

Ritter, K a k t . S u d a m e r . 3 : 9 2 7 119801 T L : Chile, lllapel. P i c h i d a n e « .

Kattermann:

Eriosyce

3 0 . 1 a . E r i o s y c e s u b g i b b o s a i H a w o r t h I K a t t e r m a n n , supra, Echinocactus

subgibbosus

119. [subsp. and var. s u b g i b b o s a

Neoporteria Chilenia

subgibbosa

subgibbosa

1515)

H a w o r t h , Phil. M a g . 10: 4 1 9 (18311. T L : Chile, V a l p a r a i s o , T : T. Bridges

extant or illustrated. N e o t y p e (designated here): Kattermann

s.n. not

1 ( D B G ) . [5161

I H a w o r t h I Britton & Rose, Cact. 3: 9 7 ( 1 9 2 2 ) . | 5 1 7 )

( H a w o r t h ) Backeberg in Backeberg & K n u t h , K n k t u s - A B C , 3 0 1 ( 1 9 3 5 ) , invalid (Art.

43.1). [518] Echinocactus

exsculptus

subgibbosus Echinocactus

O t t o ex Pfeiffer, E n u m . C a c t . . 6 5 ( 1 8 3 7 ) , illegit. (Art. 6 3 . 1 ) . O b s . : T y p e a s for E.

H a w o r t h . 15191 acutissimus

O t t o & Dietrich. Allg. G a r t e n z . 3: 3 5 3 11835). T L : Chile or B u e n o s A i r e s (?), not

e x t a n t or illustrated. [ 5 2 0 | Chilenia

acutissima

(Otto & Dietrich) Backeberg in Backeberg & K n u t h . K a k t u s - A B C , 3 0 0 ( 1 9 3 5 ) . invalid

(Art. 4 3 . 1 ) . [521J Echinocactus

caxtaneoides

C e l s ex S a l m - D y c k . Cact. Hort. Dyck. 1 8 4 9 : 1 6 5 ( 1 8 5 0 ) . T L : N o t extant

or

illustrated. [ 5 2 2 ) Chilenia

castaneoides

( C e l s ex S a l m - D y c k ) Backeberg in Backeberg & K n u t h . K a k t u s - A B C , 3 0 0 ( 1 9 3 5 ) ,

invalid (Art. 4 3 . 1 ) . [5231 Neoporteria Echinocactus Chilenia

castaneoides

rastrata

Echinocactus

( C e l s ex S a l m - D y c k ) W e r d e r m a n n , K a k t e e n k u n d e 1 9 3 8 : 6 4 ( 1 9 3 8 ) . I524J

rostratus

Jacobi, Allg. G a r t e n z . 2 4 : 1 0 8 ( 1 8 5 6 ) . | 5 2 5 ]

(Jacobi) Backeberg. K a k t e e n k u n d e 1 9 3 9 : 8 2 ( 1 9 3 9 ) . [ 5 2 6 ]

cttpulatus

Forster, H a m b u r g . G a r t . B l u m e n z e i t . 17 161 ( 1 8 6 1 ) . T L : Based on specimen in Hort.

Hauge; p r e s u m a b l y not preserved. 1527] Chilenia

heteracantha

Backeberg in Backeberg & K n u t h . K a k t u s - A B C , 4 1 7 1 1 9 3 5 ) . invalid (Art. 4 3 . 1 ) . T L :

B a s e d on material from Chile. [ 5 2 8 ] Neoporteria

heteracantha

W . T . M a r s h a l l . Cact., 1 4 3 < 19411. O b s . : also publ. by Backeberg in Cact. Succ. J.

( U S ) 23(4): 1 1 8 ( 1 9 5 1 ) . [ 5 2 9 ) Neoporteria

subgibbosa

t heteracantha

i W . T . M a r s h a l l ) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 8

( 1 9 6 6 ) . [5301 Chilenia

subcylindrica

Backeberg in Backeberg & K n u t h . K a k t u s - A B C , 4 1 7 ( 1 9 3 5 ) , invalid (Art. 4 3 . 1 ) . T L :

B a s e d on material from Chile, foot of A n d e s . | 5 3 1 | Neoporteria

subcylindrica

Neoporteria

castanea

(Backeberg) Backeberg, Cact. Succ. J. ( U S ) 2 3 : 1 1 9 ( 1 9 5 1 ) . invalid. 15321

var. tunensis

Ritter, T a x o n 12: 3 4 ( 1 9 6 3 ) . T L : Chile. T u n a , near S a n Fernando, T : Ritter

2 3 6 a ( U ) . 1533] 3 0 . 1 b . E r i o s y c e s u b g i b b o s a var. l i t o r a l i s Neoporteria

litoralis

Neoporteria

subgibbosa

Kin. i

K a t t e r m a n n . supra,

f. litoralis

tRitter)

Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 8 ( 1 9 6 6 ) . [ 5 3 6 |

3 0 . 1 c . E r i o s y c e s u b g i b b o s a var. c a s t a n e a (Ritter) K a t t e r m a n n , supra, Neoporteria

119. | 5 3 4 |

Ritter. Succulenta 1 9 5 9 : 2 8 ( 1 9 5 9 ) . T L : C o q u i m b o . , T : Hitter 2 1 9 ( Z S S ) . [5351

castanea

1 1 9 . 1537]

Ritter, T a x o n 12: 3 4 ( 1 9 6 3 ) . T L : Chile, prov. Talca. Villa Prat. T : Ritter 2 3 6 ( U ) . | 5 3 8 ]

3 0 . 2 a . E r i o s y c e s u b g i b b o s a subsp. and var. c l a v a t a (Sohrens ex S c h u m a n n ) K a t t e r m a n n , supra, EchiniK-actus

clavatus

S o h r e n s ex S c h u m a n n , M o n a t s s c h r . K a k t . - K u n d e 10: 2 7 ( 1 9 0 0 ) . T : Sohrens

1 1 9 . [5391 s.n., not

extant. L e c t o t y p e (designated here): S c h u m a n n , I.e. 2 5 (illustration). [ 5 4 0 | Neoporteria

clavata

(Sohrens ex S c h u m a n n ) W e r d e r m a n n , K a k t e e n k u n d e 1 9 3 9 : 7 9 ( 1 9 3 9 ) . | 5 4 1 ]

Neoporteria

clavata

var. procera

Ritter, T a x o n 12: 3 4 ( 1 9 6 3 ) . T L : N Chile, E of L a Serena, T : Ritter 4 8 3 ( U ) .

1542] 3 0 . 2 b . E r i o s y c e s u b g i b b o s a var. n i g r i h o r r i d a (Backeberg) K a t t e r m a n n , supra, Neoporteria

nigrihorrida

119. [5431

Backeberg, K a k t e e n k u n d e 1 9 3 9 : 8 1 ( 1 9 3 9 ) . T L : B a s e d on material from Chile,

Coquimbo. L e c t o t y p e 'designated here): Illustration in Backeberg & K n u t h . K a k t u s - A B C . 3 0 1 1 1 9 3 5 ) . [5441 Chilenia

nigrihorrida

Backeberg in Backeberg & K n u t h , K a k t u s - A B C , 4 1 7 ( 1 9 3 5 ) , invalid (Art. 4 3 . 1 ) . (5451

Chilenia

nigrihorrida

var. major

Backeberg in Backeberg & K n u t h . K a k t u s - A B C , 4 1 7 ( 1 9 3 5 ) , invalid (Art.

4 3 . 1 ) . [5461

167

Kattermann: Chilenia

nigrihorrida

var. minor

Eriosyce

Backeberg in Backeberg & K n u t h , K a k t u s - A B C , 4 1 7 * 1 9 3 5 ) , invalid (Art.

4 3 . 1 ) . [547J Neoporteria

microsperma

Ritter, Succulenta 1 9 6 3 : 6 ( 1 9 6 3 ) . T L : 1 5 k m E of La S e r e n a , T : Ritter

535 (U).

[548] 3 0 . 2 c . E r i o s y c e s u b g i b b o s a var w a g e n k n e c h t i i (Ritter) K a t t e r m a n n , supra, Neoporteria Ritter

wagenknechtii

119. 15491

Ritter. Succulenta 1963( 1): 5 (19631. T L : Chile, N of IM Serena, Juan Soldado, T :

7 1 5 ( U ) . 1550]

Neoporteria

wagenknechtii

var. парта

Ritter, Succulenta

1963(1): 5 ( 1 9 6 3 ) . T L : Chile. N of La Serena.

Choros Bajos. T : /titter 7 1 4 ( U ) . [5511 Neoporteria

rapifera

Ito, Cact., 4 2 1 (19811. Based on N. wagenknechtii

var. napina.

3 0 . 2 d . E r i o s y c e s u b g i b b o s a var. v a l l e n a r e n s i s (Ritteri K a t t e r m a n n . supra, Neoporteria

vallenarensis

|552]

119. [ 5 5 3 ]

Ritter, Kakt. S u d a m e r . 3: 1 0 3 2 ( 1 9 8 0 ) . T L : Chile, between V a l l e n a r and Freirina.

1 9 6 0 , T : Witter 1 0 8 8 ( U ) . [ 5 5 4 ] 3 1 . E r i o s y c e s o c i a b i l i s (Ritter) K a t t e r m a n n , supra, Neoporteria

sociabilis

Neoporteria

siKiabilis var. napina

T : Ritter

119. [ 5 5 5 ]

Ritter, Succulenta 1 9 6 3 : 3 ( 1 9 6 3 ) . T L : N C h i l e , Totoral Bajo, T : Ritter 6 5 5 ( U ) . [ 5 5 6 ] Ritter, Succulenta 1963(1): 3 ( 1 9 6 3 ) . T L : N Chile. Carrizalillo, Freirina,

655a (U). [557]

3 2 . 1 . E r i o s y c e s e n i l i s (Backeberg) K a t t e r m a n n . supra, Neoporteria

senilis

Echinocactus

senilis

senilis

T: Philippi

119. [subsp. s e n i l i s ] [ 5 5 8 ]

Backeberg in Backeberg & K n u t h , K a k t u s - A B C . 2 6 2 ( 1 9 3 5 ) . O b s . : Treated as nom. nov.

for Echinocactus

Philippi. [ 5 5 9 ]

Philippi, Gartenflora 3 5 : 4 8 5 , t. 1 2 3 0 a ( 1 8 8 6 ) , nom. illeg. (Art. 6 4 . 1 ) . T L : Chile, O v a l l e ,

s.n. ( S G O 4 1 2 8 3 ; see M u n o z Pizarro 1 9 6 0 : 9 2 ) . O b s . : N o t E. senilis Beaton In Loudon, G a r d .

M a g . 15: 5 5 0 ( 1 8 3 9 ) . [ 5 6 0 ] Neoporteria

gerocephala

Echinocactus Neoporteria

senilis

multicolor

Ito, Expl. Diagr., 2 1 1 ( 1 9 5 7 ) , nom. illeg. (Art. 6 3 . 1 ) . O b s . : later nom. nov. for Philippi. 1561] Ritter. T a x o n 12: 3 3 11963). T L : C h i l e , C o q u i m b o , T : Ritter 2 4 3 ( U ) . [ 5 6 2 ]

3 2 . 2 . E r i o s y c e s e n i l i s subsp. c o i m a s e n s i s (Ritter) K a t t e r m a n n , supra, Neoporteria

coimasensis

Neoporteria

robusta

119. [ 5 6 3 ]

Ritter, T a x o n 12: 3 4 ( 1 9 6 3 ) . T L : Chile, prov. A c o n c a g u a . T : Ritter 4 7 3 ( U ) . [564J

Ritter, T a x o n 12: 3 4 ( 1 9 6 3 ) . T L : Chile, prov. S a n t i a g o , E from M o n t e n e g r o , T : Ritter 4 7 3 c


villosus

462 (DBG). [566]

Monville, Hort. Univ. 1: 2 2 3 ( 1 8 3 9 ) . T L : N o t extant or illustrated. N e o t y p e i designated here):

Kattermann Neoporteria

1 1 9 . H o l o t y p e : Kattermann

119. [ 5 6 7 |

71 ( D B G ) . [ 5 6 8 ]

villosa

(Monville) Berger. K a k t e e n , 3 4 3 ( 1 9 2 9 ) . | 5 6 9 !

villosa (Monville) Backeberg, Blatt. K a k t . 1 9 3 5 ( 1 2 ) : I unpaged I ( 1 9 3 5 ) . incorrect n a m e (Art. 1 1 . 3 ) .

Bridgesia 1570] Chileniopsis

villosa

Echinocactus

(Monville) Backeberg in Backeberg & K n u t h . K a k t u s - A B C , 2 8 4 ( 1 9 3 5 ) . | 5 7 0 a ]

polyraphis

Pfeiffer ex S a l m - D y c k , Cact. Hort. Dyck. 1 8 4 9 : 1 6 6 ( 1 8 5 0 ) . T L : Not e x t a n t or

illustrated. [ 5 7 1 ] Bridgesia

polyraphis

Chileniopsis

(Pfeiffer ex S a l m - D y c k ) Backeberg, Blatt. K a k t . 1 9 3 5 ( 1 2 ) : [unpaged! ( 1 9 3 5 ) . [ 5 7 2 ]

polyraphis

(Pfeiffer ex S a l m - D y c k ) Backeberg in Backeberg & K n u t h , K a k t u s - A B C . 2 8 4 ( 1 9 3 5 ) .

polyraphis

(Pfeiffer ex S a l m - D y c k ) Backeberg, Cact. Succ. J. ( U S ) 2 3 : 1 1 8 ( 1 9 5 1 ) . [ 5 7 4 ]

1573] Neoporteria Chilenia

atrispinosa

Backeberg, Blatt. K a k t . 1938(3): [unpaged] ( 1 9 3 8 ) , invalid (Art. 4 3 . 1 ) . T L : Based on

material from Chile, coastal region, Huasco. [5751 Neoporteria

1K8

atrispinosa

Backeberg, K a k t e e n k u n d e 1939(5): 81 ( 1 9 3 9 ) . [5761

Kattermann: Chilenia

cephalaphora

Eriosyce

Backeberg, Blatt. Kakt. 1938(3): limpagedl 11938). invalid (Art. 4 3 . 1 ) . T L : Based on

material from Chile, coastal region, H u a s c o . [ 5 7 7 ) Neoporteria

cephalaphora

Neoporteria

laniceps

Backeberg, K a k t e e n k u n d e 1 9 3 9 : 8 1 ( 1 9 3 9 ) . [5781

Ritter, T a x o n 12: 3 4 < 1 9 6 3 ) (as 'planiceps',

orth. err.). T L : N Chile, Totoral Bajo, T :

Ritter 4 8 3 ( U ) . [5791

N A M E S T O BE E X C L U D E D FROM Echinocactus

ambiguus

cannot be an Eriosyce Chilenia

ambigua

Echinopsis

ERIOSYCE

H i l d m a n n ex S c h u m a n n , G e s a m t b . K a k t . , 3 1 1 ( 1 8 9 8 ) . O b s . : From the description this (FK). |580)

( H i l d m a n n ex S c h u m a n n ) Backeberg, K a k t e e n k u n d e 1939: 8 2 ( 1 9 3 9 ) . [5811

catamarcensis

W e b e r in Bois, Diet, hort., 4 7 1 ( 1 8 9 8 ) . T L : A r g e n t i n a , C a t a m a r c a , T :

Schickendantz

s.n. (fide S c h u m a n n , G e s a m t b . K a k t . , 2 4 4 . 1 8 9 8 ) . O b s . : 1 m tall, therefore not part of Eriosyce.

Britton

& Rose, T h e Cact. 3: 1 4 6 ( 1 9 2 0 ) list it a s a species possibly referable to Trichocereus,

include

Echinocactus

catamarcensis

but

Spegazzini as a s y n o n y m . Later, and without c o m m e n t , they used the latter

n a m e as the basionym for Malacinarpus

catamarcensis

(Spegazzini) Britton & Rose (I.e. 3: 197. 1 9 2 2 ; see

no. 4 a ) . [5821 Cereus

catamarcensis

Pyrrhocactus Neoporteria Echinocactus Neoporteria

W e b e r in Bois, Diet, hort., 4 7 1 ( 1 8 9 8 ) , pro. s y n . 15831

catamarcensis catamarcensis cupreatus

( W e b e r ) Backeberg in Backeberg & K n u t h . K a k t u s - A B C , 2 6 2 ( 1 9 3 5 ) . [5841 ( W e b e r ) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 5 ( 1 9 6 6 ) . | 5 8 5 |

Poselger ex R u m p l e r in Forster's H a n d b . Cact. ed. 2 , 6 0 2 ( 1 8 8 6 ) . [5861

tuberisulcata

var. cupreata

(Poselger) Donald & Rowley, Cact. Succ. J. Gr. Brit. 2 8 : 5 8 11966).

15871 Echinopsis

korethroides

W e r d e r m a n n ex Backeberg, N e u e K a k t e e n , 8 4 (19311. T L : N A r g e n t i n a , A n d e s , not

extant or illustrated. O b s . : Referred to Echinopsis Eriosyce

korethroides

Echinocactus

lindleyi

formosa

by Rausch < 1 9 7 7 ) . [ 5 8 8 |

( W e r d e r m a n n ) Backeberg in Backeberg & K n u t h . K a k t u s - A B C . 2 7 3 ( 1 9 3 5 ) . 15891 Forster, H a m b u r g . Gart. B l u m e n z e i t . 17: 1 6 2 ( 1 8 6 D . T L : Probably from Peru; not

extant or illustrated. O b s . : A nebulous n a m e ; see Britton & Rose. T h e Cact. 3: 9 0 ( 1 9 2 2 ) . It cannot be Eriosyce Neopctrtena

(Islaya)

from the description given ( F K ) . [5901

W / e y i (Forster) Donald & Rowley, Cact. Succ. J. G r . Brit. 2 8 : 5 7 ( 1 9 6 6 ) . 15911

169

Kattermann:

Eriosyce

Alphabetical List of Synonyms All names are followed, first, by the number of the taxon to which they are referred in the systematic account and documented in Appendix IV, and second by a serial number in 1 I by which they can be more quickly traced in Appendix TV. Valid names are printed in roman type; invalid and illegitimate names, and names of doubtful application in italic type. For the scope of the list, see page 131. Acanthocalycium

Chileniopsis

andreaeanum

6|62|

polyraphis villosa

[Echinocaclusl 3 3 1573]

fuscus

33 [570a|

geissei

Bridgesia

22.2?

var.

polyraphis

3 3 [5721

Chileocactus

villosa

3 3 15681

froehlichianus

1 0 [1321

aerocarpa

25[4761

duripulpa

2 0 . 2 b [3581

albicans

Л f [2271

hankeanus 10? [ ! 9 9 |

22.2?

horridus

Chileorebutia

horrid im

lOe 1171]

villosus

33 [569]

Cereus catamarcensis

esmeraldana glahrescens

21.2 [373]

molendensis

napina

2 6 [482)

malleolata var. solitaria

atrispinosa castaneoides cephalaphora chilensis var.

confinis

var.

cylindracea

[581]

odieri

2 1 . 1 a 1366]

reichei

21.1?

[5231

saxifraga

33 1577] 29a [502]

Delaetia

29a | 5 0 6 |

woutersiana

22.2

[435]

2.9a [5081 22.2?

[442]

Echinocactus

22.2*

|448]

acutissimus

fusca

22.2?

(4391

heteracantha

30.1a

[5281

aspillagae

jussieui

19.3?

[334]

auratus

kunzei

1 1 a [2121

napina

20.1|344]

10? 1193] 20.11349] 8[72|

napinus

2 0 . 1 [3421

n id и*

11? 1228]

nigricans

12?[240[

iK-cultus

19.3? [337 \

odieri var.

2 1 . 1 a 1363] mebbesii

2i./.?|379]

polyraphis

3 3 [5711

reichei

21.1?

rostratus

3 0 . 1 a 1525]

sandillon

l a [3]

sanjuanensis

ebenacantha

nigricans

[389]

9.2 [1271

fobeana

nidus

2 6 1485] 2 6 1490] 20.1 [347]

33 [ 5 7 5 ] 30.1a

[590] millarioides

mitis

Ж la 15211

ambigua

lindleyi mam

ambiguus

[5801 16 12771

bulbocalyx castaneoides

11? [ 2 3 0 ]

catamarcensis

12? [2371

centeterius

|383]

4a | 4 1 |

senilis 3 0 . 1 a 1520]

[3321

11a [ 2 0 9 ]

21.3 [377]

krausii

Chilenia arutissima

19.3?

kunzei

fulva 1583]

8 |69|

jussieui

24 [471]

[444]

lOe 11721

islayensis Cactus curvispinus

[437]

11? [2251

32.1 15601

soehrensii

10? 1197]

strausianus

4 a [351

sultgihhosus

3 0 . 1 a [5161

l a [2]

supertextus

10? 1 1 9 6 ]

3[26|

tuberisulcatus

lOe [1671

3 0 . 1 a [5221

umadeave

51541

4a [38] 10? [ 1 8 8 ]

Echinopsis

nigrihorrida

30.2b

[545]

ceratistes

70?[1871

catamarcensis

[582]

var.

major

30.2b

15461

chilensis

2 9 a [5001

korethroides

[5881

var.

minor

30.2b

1547]

29a 1505]

nigricans

var. w n f l n i s

occulta

19.3? [ 3 3 9 ]

clavatus

3 0 . 2 a 15401

odieri

2 1 . 1 a [3651

cupreatus

[5861

pseudoch Hen sis reichei mslr.ilii

29a

15111

cupulatus

2U?№7\

curvispinus

3 0 . 1 a [5261

ehenaeanthus

subcylindrica

30.1a

15311

subgibbosa

30.1a

{5181

10[1331 22.2?

1441]

algarrobensis ausseliana ceratistes

l a 161 la [ 5 ] la [9]

22.2?

[4431

var. combarbalensis

exsculptus

30.1a

[5191

var. coquimbensis

l a f 11J

fobeanus

22.2?

14471

var. jorgensis

l a [121

var. mollensis

l a 113]

var.

minor

froehlichianus

170

3 0 . 1 a [5271

12?1236]

Eriosyce

10? 1198]

la [10|

Kattermann: [Eriosyce ceratistes)

Eriosyce

I Horridocactus

1 Neochilenial

var. tranquillaensis

la | 161

var. vallenarensis

l a 1141

22.2/

[4491

var. zorillaensis

l a 1151

Islaya

fusca

22.2!

1440]

l a 171 1589)

Imnlnr

8 [78]

glaucescens

2 2 . 3 a 14571

brevicylindrica

Я 1801

gracilis

2 2 . 1 b [4111

l a (8)

copiapoides

8 1821

hankeana

ihotzkyanae korethroides lapampaenais

vallenarensis

11a | 2 1 9 |

megacarpa

2 1221

var. chalaensis

rodentiophila var. lanata

2 \20]

var. pseudomollendensis

sandillon Spinibarbis

la | 4 | l b 1181

Friesia

8 |86|

Gymnocalycium

var. orientalis andicola var. mollensis var. robustus armatus

krausii

26 [ 4 8 3 ]

S 1891

kunzei

11a [2131

var.

tenuispina

8 [911

lembekei

8 [92]

malleolata

var. neglecta

8 1961

mitis

8 [70]

monte-amargensis

var. copiapoides

8 1831

napina

var. minor

8[76]

napina var. mitis

22.1a

[3971

neoreichei

20.2a

[356]

10a 11401 12 [2341

8 [75]

minuscula

8 11071

molendensis

8 [731

omasensis

8[109] 8 [981 8 1971

odoriflora

8 [991

paucicostata

var. curvispina

8[108]

unguispina

reichei

10a [1451 19.1 [3011 us

marksianus nigricans paucicostatus var. viridis pygmaeus robustus trapichensis tuberisulcatus

curvispinus

18 12981

11? [2261

var. gracilis

catamarcensis mammiUarioides

15 1271]

lissocarpus

Malacocarpus

18 | 2 9 5 ]

10? | 2 0 2 ]

heinrichianus

2 1 . 1 a 13671

paucispinosa

f'roehluh

grandiflorus

9 . 2 [1281 19.3/

odieri

10c1159]

11a | 2 1 4 |

geissei

nuda

4a [391 10 [134] 104 11941 21.1/

[3841

Neochilenia aerocarpa

2.5 14771

andreaeana aricensis

fi [591 9.2 [1211

aire

21.If

|392|

10? 1204]

calderana

2 2 . 1 b 1408]

17b12881

earneoflora

27.213751

17c 1293]

chilensis

29a[503]

17a 12821

chorosensis

19.1 13051

12? 12411

confinis

2 2 . 2 [4241

deherdtiana

2 2 . 2 [4281

dimorpha

13 12451 1.9.1 [3071 19.2a [3201

1404]

duripulpa

2 0 . 2 b 13591

/«#11831

eriocephala

2 2 . 4 6 14621

22.1b

14.1 12511

occulta

10a 1142]

eriosyzoides ianus

nigriscoparia

paucispina

10 11361

garaventae

[370]

20.1 [ 3 4 5 ] 20.1 13501

neofusca

2 2 . 3 a [4531 krausii

21.1b

8[1061

14.112491

engleri

2 6 14911

811011

lOd [1641

echinus

26 |486| 2 0 . 1 [3511

minor

14.2c [ 2 6 3 ]

kesselringian

8 1941

maritima

carrizalensis

curvispinus

2 0 . 2 o [3531

var. solitaria

var. hrevispina

krainziana

14.2a 12541

crispus

[3351

8 [87]

spinosior

10b11501

atroviridis choapensis

9.2 11181 19.3/

10b [1541 10a 11381

var. descendens

I355J

var.

islayensis

Horridocactus

2 2 . 1 b 1416]

jussieui

10? [2011

aconcaguensis

intermedia iquiquensis

81105]

grandis

Hildmannia

[4451

14.2b 12591 20.2a

811031

10? | 1 8 9 )

froehtichiana

22.21

huascensis imitans

flavida 5 [55)

centeterium

2 2 . 3 b 14611

divarieatiflora graodifloreos

umadearv

var.

8 |85l

floccosa fobeana

var.

niehhcsii

pilispina

21.1?

[3401 [3801

lOf [ 1 7 8 ] 2 2 . 2 [4261 2 2 . 4 [4661

pseudareichei

21.1/

pygmaea

2 2 . 1 b 14051

recondita reichei

13911

9 . 1 1113]

residua

I3S8I 2l.lt 9.2 124!

rupicola

2 2 . 1 a 1399]

scoparia

2 2 . 1 b 1413]

setosifbrn

1 9 . 2 b [3261

var. intermedia simulans

19.2a [3161 19.3 | 3 3 0 |

taltalensis

2 2 . 1 a [3951

totoralenais

I 4 . 2 d 12681

transitensis trapichensis wagenknechtii

l i b [2231 19.1 [3131 1 9 . 2 a 13181

Neoporteria andreaeana

6161J

aricensis

9 . 2 1122

armata

10c [1601

aspillagae

16 [2781

19.1 13121

eriosyzoides

11a [2161

atrispinosa

3 3 1576]

lOe 11691

esmeraldana

24[472]

atroviridis

14.2a [ 2 5 S |

171

Kattermann: I Neoporteria I bicolor

8(79]

bulbocalyx

a [28i

Eriosyce

I Neoporteria islayen sisj f. grandiflorens

[Neoporterial 8 188)

simulans

19.3 [3311

var. grandis

8193]

sociabilis

31 [556]

calderana

2 2 . 1 b 1409)

f. m i n o r

8 [771

carrizalensis

14.2c [2641

f. molendensis

8(741

strausiana

castanea

3 0 . 1 « 15381

f.

8 [901

subaiana

var. tunensis

3 0 . 1 a [5331

castaneoides

30.1a [524]

catamarcensis

15851

centeteria

10? [1921 3 3 1578]

cephalophora cliilensis

29a [ 5 0 1 ]

spinosior

jussieui var.

fobeana

kesselringiana

limariensis lindleyi

f. cylindracea

29a [510]

choapensis

lOd [1651 19.1|306| 3 0 . 2 a 15411

var. procera

3 0 . 2 a [5421

coimasensis

3 2 . 2 [5641

confinis

13 12461

crispa

14.112501 10 [ 1 3 7 ]

curvispina

var. aconcaguensis 10b [1511 kesselringiai ш 10? | 2 0 7 |

var.

var. lissocarpa f. orientalis var. vallenarensis deherdtiana dimorpha

engleri var.

var. glaucescens

18 [ 2 9 9 ]

fusca

32.1[561] 22.2?

heinrichiana heteracantha

[4381

15 [2731

geracephala hankeana

multicolor napina f.

1446)

19.1[3031 3 0 . 1 a [5291

horrida

lOe 11741

var. odoriflora

lOf[1791

huascensis

14.2b [260]

intermedia

22.1b [417]

iquiquensis

9.2 [119J

islayensis

8 [71]

|371|

32.1 [562] 20.1 [ 3 4 3 ]

mebbesii

var.

spinosior

nigrihorrida

21.1?

|382]

20.1

[3481

var. atroviridis

var.

nigricans

12? [ 2 3 9 ]

var. v e g a s a n a

10g[185]

villicumensis villosa

19.3?

wagenknechtii

21.1a [364]

var. p y g m a e a polyraphis

21.1?

[381]

8[110| 22.4 [467] 2 2 . 1 b [4061 33 |574] 22.1b [419] 3 0 . 2 c [5521

var, aerocarpa f. duripulpa var. malleolata f. eolitaria residua robusta rupicola sanjuanensis scoparia

9.1 [1141 21.1?

[386]

25[479] 20.2b [360]

4a 145) 30.2c[5501 3 0 . 2 c [5511

Notocactus reichei

21.1?

Г385]

2 6 [4871

Pyrrhocactus aconcaguensis var. orientalis andicola var. descendens var. mollensis var. robustus andreaeanus

10b 1149] 10b [ 1 5 3 | 10a [ 1 3 9 ] 10a 1141] 12 [2351 10a [1431 6160]

aricensis

9 . 2 1120]

armatus

10c [ 1 5 8 ] 16 [2791

26[4921

aspillagae

9.2 [ 1 2 5 ]

atrospinosus

4a 149] 14.2a 1253]

32.2 [565]

atroviridis

2 2 . 1 a [4001

bulbocalyx

3 [27]

4a[43]

calderanus

22.1b [407]

carrizalensis

14.2c1262]

8 1811 8 [95]

senilis

32.1 [ 5 5 9 |

8[84J

setiflora

in [ 4 8 ]

setosiflora

7(651 33 [570]

2 2 . 2 1427]

pulchella

reichei

var. napina

3 [31] 30.2d [554]

26[489]

rapifera recondita

5[57|

var. marayesensis

occulta

mebbesii

[5871 10? (2031

umadeave

odieri var.

10c [1611 14.2a [ 2 5 6 ]

cupreata

12? [ 2 4 2 ]

var. coiapoides

172

lOe [1701

var. a r m a t a

11? [ 2 2 9 ]

f. hrevispina

8 [1041

l i b [224]

tuberisulcata

vnlliana

omasensis

14.2d Г2691

transitensis

vallenarensis

[3381

22.1b [421]

2 2 . 1 b 14221

22.2 [ 4 3 Ц

f. brevicylindrica

var. divarieatiflora

transiens

30.2b [544 [

var. malleolata

2 2 . 1 a 1394]

var. carrizalensis 14.2c 12651

var. froehtichiana

pilispina

22.2?

totoralensis

21.1b

1458]

garaventae

3 0 . 1 b 15361

var. transiens

monte-amargensis

22.3a

2 4 [4731

f. litoralis taltalensis

var.

paucicostata

11a [ 2 1 7 ]

3 0 . 1 a [5301

3 0 2b 15481

[4631

esmeraldana

7 [671

[5321

f. heteracantha

micros perma

22.3h

eriosyzoides

17a[283] 17a [285J

melanacantha

nigricans

1 8 [2971 krausii

var. tunensis

nidus

3 ? [331

8 [1001 3 0 . 1 b [5351

marksiana

I l a [2201

2 2 . 3 a [4541

eriocephala

litoralis

10b [1551

19.2a [3211

echinus

8[102]

f. curvispina

neohankeana

dubia

30.1a [517]

159Ц

17b [ 2 8 9 )

19.1 [3081

30.1a

subgibbosa

12 1233]

29n [5071

clavata

subcylindrica

3 3 [5791

2 9 a 15091

chorosensis

[333| [4501

10? | 2 0 5 ] 11a[210|

var. cylindracea

4a [37] 15 | 2 7 5 |

19.3?

laniceps

var. confinis

31 [ 5 5 7 ]

22.2?

krainziana kunzei

var. napina

22.1b [414]

19.2b [ 3 2 7 ]

catama rcens is ( W e b e r ) Backeb. ( S p e g . ) Ritter

[584] 4a [ 4 0 ]

Kattermann: IPyrrhocactusI centeterius

10?[190]

I Pyrrhocactus 1 nigricans

Eriosyce | Thelocephala 1 12? [ 2 3 8 ]

esmeraldana

(Hxultus

23 [ 4 6 9 ]

fankhauseri

2 9 a 1504]

odoriflorus

Ю Г Ц77]

2 9 b [5141

pachacoensis

choapensis

lOd [1631

pamaensis

choroeenais

19.113041

chaniarensis

19.1 (3101

chilensis var. albidiflorus

coliguayensis

con fin is

10a 11461 13 12441

crispus

14.1 [2481

•:.!'- 1 - | - III.deherdtianus dimorphus

var. minor

floccosus ftoribunduH froehlichianus

longirapa

2 6 14941

22.4 [465]

malleolata

26 [488]

4 a 1501

napina nuda

9.1 [ 1 1 2 ]

odieri

21.1a [368]

9.2 [1231

reichei

21.1?

10g[182]

tenebriea

11a 1215]

var. v e g a s a n u s rupicola -.in jl . п и т i -i •

4a [ 4 2 ]

saxifrage

9.2 [ 1 2 6 ]

9.2 1130]

scoparius

22.1b [412]

10?[2001

setiflorus

10a 1144] 19.1 | 3 0 2 ] 10c 1173]

var.

twtosiflorus setosiflorus

4a [461

var. intermedius 19.2a [ 3 1 5 ] simulant) straueianuB subaianits

19.3 [3291 4a 136] 15 1274]

laltalt'iisis

22.1a [396]

tenuis

2 2 . 1 a [4011

Ю Г Ц761

i ..I i i r a Id i-l -

14.2d [267]

var. orientalis

10b 11561

transiens

22.1b [420]

var. robustus '•ll.l.-IVIh •

10g[186)

transitensis

l i b [222]

14.2b [2581

trapichensis

19.1 [ 3 1 1 ]

intermedius

22.1b [415]

truncatipetalus

17b|290|

tuberisulcatus

lOe [ 1 6 8 ]

19.3!

1336)

19.2a [ 3 2 3 ]

var. spinosior

19.2a [ 3 2 2 ]

ngianus

10? Г206]

umadeave var. marayesensis

11a1218]

vexatus

9.1 [115]

29a[5121

villicumensis

kunzei

11a 1211)

vollianus

12 12321

lissocarpus

17b [2871

var. gracilis m ammilla

rioides

marksianus var. tuncnsis

wagenknechtii

17c [2921

Reicheocactus

10? Ц 9 5 ]

ftnnbundus

17a [2841

megliolii

19.2a [317]

9.2 [1291

3 [291 7|66] 2 2 . 2 (4301

Rodentiophila atacamensis

2[24]

lanata

2 [211

megacarpa

2 [23]

var. denHispinus

2 2 . 2 (432J

var. elongatus

2 2 . 2 [4331

Thelocephala

var. flaviflorus

2 2 . 2 14341

aerocarpa

f.

22.2

duripul pa

woutersianus

7 [64] 4a [ 4 4 |

I7a[2811

melanacanthus neohankeanus

5 Г561 3 [SO]

vallenarensis

krausii limariensis

if)? 11911

4a [ 4 7 ]

/(VI1801

var. australis

Uebelmannia centeteria

1 9 . 2 b 13251

var. qratabOia

9.2 [117]

1390]

27 [496]

10g[184| 2 2 . 1 a [3981

22.3b | 4 6 0 |

var. aconcaguensis 10b [ 152)

kesselri

2 5 14801

robustus

22.1b [410]

jusaieui

2 6 [4931 20.1 [3461

residuus

gracilis

iquiquensis

[3541

3 ? [321 2 2 . 3 a [4551

reconditus

26 [484] 20.2a

2 2 . 3 a [4521

15|2721

minor

var. Bolitaria

2 2 . 1 b [4181

22.3a (456]

hcinrichianus

krausii lembckei

2 2 . 2 14291

2 2 . 1 b 1403]

garaventae

horrid us

10a [ 1 4 7 ] 2 2 . 2 [4251

pulchellus

glaucescene grandiflorus

platyacanthus

21.2 [374]

pygmaeus

18 [2961

(•ri(js> /unit's

pilispinus

21.3 [378]

10 [1351 19.2a [ 3 1 9 |

engleri

var. viridis

2 7 14971

fulva glabrescens

19.1 [ 3 0 9 |

dubius echinus

paucicostatua

4b [52]

2 4 14741

|436]

25 14781 20.2b [ 3 6 Ц

173