Sea Slugs of the Red Sea (Pensoft Series Faunistica)

Sea Slugs of the

Red Sea

Nathalie Yonow

SEA SLUGS OF THE RED SEA Nathalie Yonow

SEA SLUGS OF THE RED SEA Nathalie Yonow

Sofia–Moscow 2008

SEA SLUGS OF THE RED SEA Nathalie Yonow Photographs: Binyamin & Shulamit Koretz, Johann Hinterkircher, Jürgen Kuchinke, Oren Lederman, Pam Kemp, Arik Diamant, Alex Double, Amir Stern, Angel Valdés, Bernard Picton, Chris Todd , Doreen Sharabati , Ewald Lieske, Gunnar Bemert, Götz Reinicke, Gordon Smith, Helmut Göthel, Hagen Schmid, Helmut Schuhmacher, Hans Sjöholm, Jacob Dafni, Johnny Walker, Kevin Metcalfe, Mohamed Habib, Moshira Hassan, Michel Jozon, Michael Schrödl, Nechama Ben-Eliahu, Nicole Kenward, Philibert Bidgrain, Razi Cohen, Schmuel Schrenkel, Tuula Metcalfe, Thomas Paulus, Thilo Wanner, Viviana Farstey, Volker Neumann Disclaimer: Where known the provenance and copyright have been assigned to each of the illustrations. All reasonable effort has been made to contact copyright holders. In the event copyright is claimed by individuals, please write to the publishers who will endeavour to acknowledge and correct such information in the next edition. Pensoft Series Faunistica No. 74 ISSN 1312-0174 ISBN 978-954-642-327-6 (HB) ISBN 978-954-642-423-5 (e-book) © PENSOFT Publishers All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the copyright owner. Pensoft Publishers Geo Milev Str. 13a, Sofia 1111, Bulgaria Fax: +359-2-870-42-82 [email protected] www.pensoft.net Printed in Bulgaria, 2008

TABLE OF CONTENTS

Introduction The Red Sea 7 The Opisthobranchs 18 Nineteenth and early twentieth century engravings Savigny 1817 34 Ehrenberg 1828 36 Rüppell and Leuckart 1828 & 1830 38 Vayssière 1912 46 Pruvot-Fol 1933 50 Checklist of species recorded from the Red Sea Key to species of Red Sea opisthobranchs 66 Species accounts Order Cephalaspidea 74 Order Anaspidea 96 Order Pleurobranchida 109 Order Sacoglossa 118 Order Nudibranchia Suborder Doridina 128 Suborder Dendronotina 225 Suborder Arminina 240 Suborder Aeolidina 244 “Incertae sedis” 271 Glossary 277 Acknowledgements 282 Bibliography 284 Index 294

52

1. The first chart of the Red Sea, measured and drawn by Carsten Niebuhr, a member of the first expedition sent to explore Arabia in 1761 [courtesy of www.ancestryimages.com].

INTRODUCTION THE RED SEA Physical Characteristics The Red Sea, lying between 30°N and 12°40’N, is located between Saudi Arabia and the African mainland. It is a long (1932 km) and narrow (average 280 km) body of water, generally shallow (500 m) with maximum depths of 2850 meters within the central rift. At its northern end the Red Sea divides around the Sinai Peninsula into two gulfs, the shallower Gulf of Suez to the west and the deeper (1800 m) Gulf of Eilat (Gulf of Aqaba) to the east. The opening of the Suez Canal in 1869 linked the Red Sea to the eastern Mediterranean. At the southern end, the narrow straits of Bab al Mandab, or “Gate of Tears” in Arabic, mark the boundary between the Red Sea and the Gulf of Aden, and beyond to the Arabian Sea and Indian Ocean (fig. 1). The shallow coastal shelf is extensive in the southern Red Sea, but is limited to only a few kilometres offshore in the central and northern regions. Most of this shelf is shallow, 0–5 m in depth, and covered with actively growing coral reefs (fig. 2). Beyond this zone, the sea bed drops in a series of stepped cliffs with a shelf at 50 m to the main trough at 500 m depth and gradually to 1000 m, followed by steep slopes to the central rift of 1500 m with localized deeps. The Gulf of Suez is shallow

Introduction 7

2. An aerial view of the shallow coastal shelf fringing the Arabian coastline (photo G. Bemert).

and flat-bottomed, while the Gulf of Eilat is steep-sided and deep. The latter is cut off from the Red Sea by a sill 250–300 m deep, similar to that at the Straits of Bab al Mandab, which is 100–130 m deep; both effectively limit sea water exchange between the Gulf of Eilat and the Red Sea and the Red Sea and the Gulf of Aden respectively. The Red Sea is the axis of the African rift system which is still active today. The Arabian Peninsula and African continent are slowly drifting apart; this rift extends from the Jordan Valley to the East African Rift Valley in a north-south direction and east-west from Aden to the Indian Ocean. As the land masses move apart, new crust is formed by hot magma in the deeps. The Red Sea is relatively young, originating in the Mesozoic, perhaps 180 million years ago. During its formation, the sea often covered the region. Much later, in the Miocene (25–5 million years ago), the entire basin underwent the ‘Great Evaporation’, connected to the ancient Mediterranean by a shallow strait but cut off from the Indian Ocean in the south. Five million years ago, land uplifts closed the connection to the Mediterranean in the north and the Indian Ocean broke through in the south, carrying with it Indo-Pacific faunas. A number of glaciations then occurred in the region, the last one ending 20,000–15,000 years ago. It is only since then that the tropical coral communities have re-established themselves. Fluctuations in depth, salinity, and temperature affected the fauna and flora over long periods, suggesting that present day environmental and physical conditions are in delicate balance. At the moment, this balance is maintained by the inflow of surface waters through the Bab al Mandab and loss through evaporation. The sediments are largely biogenic classic calcium carbonate, derived from the bordering reefs and shells of pelagic organisms (fig. 3). 3. Deposits of the pelagic pteropod opisthobranch Creseis acicula from the depths of the Gulf of Eilat, dating 14,000 years (photo V. Farstey).

8 Introduction

High surface temperatures and large salinity changes characterize the waters of the Red Sea. Temperatures range from 18°C in February to 32°C from June to September. Within the 50 m depth contour, temperatures of 30–38°C occur with a maximum of 45°C in lagoons behind the fringing reefs. Salinity ranges from 36.5 ‰ in the south to 40.5 ‰ in the north. No rivers discharge into the Red Sea: rain is sparse, only 10–55 mm per year, and evaporation is high. The water deficit is made up by input from the Bab al Mandab, as the weak flow through the Suez Canal is negligible. Flow rates are affected by the different physical and chemical characteristics of the Red Sea and Gulf of Aden waters, the underwater topography, and the winds. Currents are affected by the Indian Ocean monsoons, causing drifting northwards in winter and southwards in the summer, but these are relatively weak. Tides in the Red Sea are not affected by those of the Indian Ocean. Instead, there is a local oscillatory tide of small amplitude which causes high water at one end and low water at the other. The range is greatest at each end, approximately 60 cm at the mouth of the Gulf of Suez and 90 cm in the south. In the Gulfs of Eilat and Suez, the spring tidal ranges are 1.2 m and 1.5 m respectively. Vertical variations of 20–30 cm occur along the coastlines, locally affected by meteorological conditions. The tides oscillate around the central part of the Red Sea at 20–21° N which has almost no daily tidal changes, near Suakin (Sudan) and Jeddah (Saudi Arabia). The tides are of semi-diurnal period, with a difference of approximately 12 hours between successive high and low water at any given location. Because of the oscillatory nature of the tides, there is a six-hour difference between times of high water (or low water) at the two ends of the Red Sea. Marine Ecosystems and Habitats The Gulf of Suez boasts the most northerly coral reefs of the Indian Ocean, which consist of small coral patches at depths of between 1–5 meters lying on calcareous sandy and silty substrata. Fringing reefs can be found on the African coast approximately 50 km south of Suez, marking the beginning of a long stretch of reef extending southwards. The Gulf of Eilat supports only a narrow fringing reef extending into the Red Sea along both shores. The fringing reefs extend for short distances offshore, up to 70–80 meters, before either sloping off gently or producing steep drop-offs. Further south, offshore reefs occur on the Arabian side, and islands and atolls appear along this coastline. Waters are calm, well lit, warm, and clear, providing perfect conditions for coral reef growth (fig. 4). It is in this ecosystem that the most flamboyant opisthobranchs are to be found, crawling exposed on coral, rubble, rocks, and ledges. Various species of sea grasses colonize the shallow lagoons behind coastal and offshore reefs, existing from mid-tidal levels to

Introduction 9

4. A drop-off reef face to the sandy sea bed at approximately 40–50 meters depth, with Dendronephthya and whip corals (photo G. Bemert).

70 meters (fig. 5). Sea grass beds tend to have high productivity, and serve to stabilize the sediments. Recent studies show that they are one of the richest and most productive marine ecosystems. Many opisthobranchs are herbivorous, and although sea grass beds are not favoured for diving or snorkelling, careful searching will reveal beautiful and varied forms. Sea grasses form an interesting biotope, and many herbivorous opisthobranchs form massive breeding aggregations here. Not only are the aggregations of interest, but some of

10 Introduction

the larger species of opisthobranchs can be found here, some weighing several kilograms each! The flat reef tops are good hunting places for opisthobranchs, with depths of 0.5 meters extending sometimes for 10–20 meters offshore, sloping gently to 1.5–5 meters (fig. 6). This region is affected by shallow wave and tidal movement. There is some coral growth with micro atolls of brain corals, but much of the substrate is comprised of rock with small to huge rocks and boulders, occasionally covered by algal felt and

Introduction 11

5. Left above: Sea grass bed (photo G. Bemert). 6. Right above: Shallow reef flat (photo G. Bemert). 7. Below: A gully along the edge of a fringing reef (photo G. Bemert).

silt. Many rocks are movable, but the larger boulders can withstand water movement and are microhabitats in themselves, covered in young corals, algae, sponges, ascidians, and hydroids. Opisthobranchs can often be found on and around these boulders, feeding on the epifauna and flora. Variously sized gullies are also found at these depths, the currentswept walls supporting a rich epilithic flora and fauna (fig. 7). Biogeography and Endemism

8. Bar graph showing the endemism levels of known groups for the Red Sea.

The study of endemism of a region describes its uniqueness, denoted by the percentage of flora and fauna which only occurs in that region and nowhere else. The real extent of endemism in the Red Sea, and indeed anywhere, is difficult to estimate due to the extensive collections and comparisons of faunas necessary. However, some taxa have been well studied and in fish, an endemism level of 17% in the Red Sea appears to be accurate (Ormond & Edwards 1987). However, this is higher for butterfly fishes, 41% (Righton et al. 1996), cardinal fishes, 22% (Randall 2003) and gobies, 27% (Goren, pers. comm.). Levels of 8% are reported for scleractinian corals (De Ventier et al. 2000) but 56% for soft corals

12 Introduction

9. Pie diagram showing the percentages of Red Sea opisthobranchs occurring in the other zoogeographical regions.

belonging to the Xeniidae (Reinicke 1997). 14% for echinoderms and 9% for algae have been estimated (Head 1987, Campbell 1987); polychaetes are estimated to have a 7.5% endemic population (calculated from Wehe & Fiege 2002). Cowries show higher endemism levels of 35% (calculated from Foin & Ruebush 1969) as do echinoids (Nebelsick 1996). Endemism of the molluscan fauna is estimated at 16% (Dekker & Orlin 2000). Of the 275 species of opisthobranchs recorded from the Red Sea, 70 are found only in this locality, representing 24% endemism (fig. 8). The distribution of molluscs throughout the Red Sea is not homogeneous, with many more species being present in the southern region. This distribution pattern also occurs with echinoids: 25 species are recorded from the northern part of the Red Sea and 43 from the southern region (Nebelsick 1996). Nineteen species of cowries are present in the northern Red Sea, but 24 in the central Red Sea and 37 in the south (Foin & Ruebush 1969). A study of the Polyplacophora reveals 23 species for the Red Sea with only two endemic species (9%), five of which are limited to the southern Red Sea (Strack 1993). Pteropods are also more speciose in the southern Red Sea (Almogi-Labin, pers. comm.). Opisthobranchs also differ from the general molluscan rule of greatest affinity to the Gulfs of Aden and Arabia and the western Indian Ocean: only two opisthobranch species which were thought to be endemic have recently been found in the Gulf of Oman and five species also occur only in the Gulf of Aden. In contrast to other groups for which information is available, there are only 8% of the species in common with the western Indian Ocean compared to 31% affinity with the broader Indo-West Pacific region (fig. 9). For example, cluster analysis of coral species show greatest affinities with the Gulf of Aden and the Persian Gulf species, and this cluster shows close similarities with the western Indian Ocean fauna (De Ventier et al. 2000). Polychaetes show a very high affinity with the nearby regions of the Gulf of Aden,

Introduction 13

10. Portrait of Petrus Forskål [courtesy of www.goran.waldeck.se/ Ento3.html].

Arabian Sea, and Arabian Gulf, and Wehe & Fiege (2002) calculate 20% endemism for the “Arabian region” alone. Remarkably, chitons also show similar affinities, with all the non-endemic species also recorded from the western Indian Ocean (Strack 1993). While molluscs in general show a trend of higher diversity in the southern half of the Red Sea, opisthobranchs again follow an opposite trend with 29 species found only in the Gulf of Eilat and not in the southern Red Sea. It remains to be seen whether these observations are real (possibly due to different habitats, shipping, or other causes) or artificial (and will decrease with increased taxonomic effort). Numerous species are not yet formally described and others recently described may yet be found in the western Indian Ocean or further afield. However, some species are different from their Indian Ocean counterparts (some are in need of new names) demonstrating a real difference with the Indian Ocean. History of Opisthobranch Research in the Red Sea

11. The title page of Rüppell’s copy of Forskål’s great work!

12. Portrait of Carsten Niebuhr in full dress given him by the Imam of Sana’a, Yemen (from Maempel 1992).

The first scientific expedition specifically to the Red Sea was the ill-fated Danish “Arabia Felix” expedition (1761–1767) commissioned by King Frederick V to explore the geological and biological Arabia of the bible. The expedition spent October 1762 to August 1763 in the Red Sea, destination Yemen. Petrus Forskål (1732–1763), a child prodigy enrolled at the University of Uppsala at the age of ten, was a Swedish botanist and student of Linnaeus, and one of the five of six scientists (plus one artist and a ship’s surgeon) who died, of malaria or the plague in Yemen, at the age of 31 (fig. 10). He was a diligent observer of marine plants and animals and his collections and detailed field notes were extensive. Forskål’s third major work, including the opisthobranchs, was edited posthumously by the sole survivor, Niebuhr, in 1775 (as Forskål 1775, fig. 11). C. Niebuhr (1733–1815) sailed from Mocha (Yemen) for India, and then on to Europe via the Persian Gulf and Tigris River, with albums, diaries, and illustrations which formed the basis of subsequent large publications. Carsten Niebuhr (fig. 12) was the first geographer to accurately measure and chart the Red Sea, and his copper engraving was used 100 years later by the British Royal navy (see fig. 1). A recent review details the history of the mollusc collections and the taxonomic status of the molluscan species described by Forskål (Yaron et al. 1986), including three opisthobranchs. A wonderful biography of the expedition is provided by Maempel (1992). The second important expedition was Napoleon I’s great “Expedition d’Egypte” from 1798 to 1801, with 167 scientists and naturalists, including the natural historian Jules César de Savigny (1777–1851): he was 21 years old (fig. 13). Savigny’s beautiful engraved plates of marine organisms, including fifteen opisthobranchs, were published in 1817, but he became blind before he was able to write the descriptions. The publication spanned 17 years, and was a major scientific achieve-

14 Introduction

13. A medallion of Jules César Savigny at 24 years old (from Pallary 1931).

14. The frontispiece of the publication of the Description de l’Egypte , spanning 17 years (from Drouin 2001).

ment (fig. 14). Seven opisthobranchs are reproduced in this book (p. 35). His pupil Jean Victor Audouin produced a brief interpretation of the molluscs (1826) which was completed by the Italian malacologist Arturo Issel in 1869 and much later by Paul Pallary (1926), a Frenchman living in North Africa and a shell collector and dealer. Bouchet & Danrigal (1982) published copies of the plates, with a history of both the plates and the collections. The German zoologists Christian G. Ehrenberg (1795–1876) (fig. 15) and Wilhelm F. Hemprich (1796–1825) made extensive invertebrate collections during an archaeological expedition to Egypt between 1820 and 1826. They separated from the main party and spent nine months on the Sinai Peninsula, where Ehrenberg studied the marine life of the Red Sea. Hemprich died of fever in Ethiopia and was buried there. When Ehrenberg returned to Berlin, he published their discourses as ten coloured plates in 1828 and the accompanying text in 1831; the opisthobranch plate is reproduced here for the first time (p. 37). Eduard Rüppell (1794–1884) (fig. 16) also travelled in the Red Sea during this period, 1821–1827, and subsequently published beautiful colour plates and monographs, of which the opisthobranchs appeared in 1828 and 1830 with F. S. Leuckart; they are reproduced here for the first time since their publication (pp. 39–45). An important review of all records of Red Sea molluscs published upon since the time of Linnaeus was provided in a book by Issel in 1869. Seventy years after Rüppell’s expedition to the Red Sea, the Austrian research vessel “Pola” (fig. 17) cruised the northern Red Sea during 1895–1896, and the southern half in 1897–1898. Dr. Rudolf Sturany published the gastropod material,

Introduction 15

15. Christian Ehrenberg (from Klausewitz 2002).

16. Portrait of Eduard

Rüppell (from Klausewitz 2002).

17. The Austrian research vessel S.M.S. ‘Pola’ in the northeastern Red Sea, anchored off the island of Jazirat an Noman (photo K. Natterer, courtesy of G. Schefbeck).

18. Portrait photograph of Sir Charles Norton Edgecumbe Eliot taken in 1920 by Bassano [courtesy of www.npg.org.uk/live/ search/person.asp?].

19. Albert Vayssière, director of the Laboratoire Marion 1921–1924 [courtesy of www.com.univ-mrs.fr/ histoire].

including a few deep water species of shelled opisthobranchs, in 1903. In 1904–1905 Cyril Crossland, a British zoologist with tropical experience, was sent to investigate the marine biota of the Sudanese Red Sea by Professor W. A. Herdman, and his collections of opisthobranchs were described in two publications by C. N. E. Eliot in 1908 and 1911 (fig. 18). Albert Vayssière (fig. 19) was the director of the Laboratoire Marion in Marseille, and specialized in opisthobranchs. He published descriptions and plates of opisthobranchs collected by Dr. Felix Jousseaume in 1894 in Aden and the southern Red Sea, and by Charles Gravier in an expedition in 1904 to the Bay of Tadjourah (Vayssière 1906, 1911a, 1911b, 1912). His colour plate from the 1912 paper is reproduced here (p. 47). The next major work on Red Sea opisthobranchs resulted from the Cambridge Expedition to the Suez Canal in 1924, which set out to assess the effects of the canal on migrations of animals since its opening. This substantial work describes four cephalaspideans and fifteen nudibranchs from the Red Sea, in addition to listing all the species recorded in the Red Sea (a total of 82) and a list of all species recorded in the Mediterranean (O’Donoghue 1929). The French “Mission Robert Ph. Dollfus en Egypte” (December 1927–March 1929, s/s ‘Al Sayad’) collected opisthobranchs in the northern Red Sea and in Lake Timsah in the Suez Canal. Alice Pruvot-Fol (Museum National d’Histoire Naturelle, Paris) published her findings on these opisthobranchs in 1933. The colour plate is reproduced here for the first time since its publication (p. 51). The John Murray Expedition (1933–1934) with the Egyptian vessel ‘Mabahiss’ (fig. 20) was the third expedition in the first half of this century to collect a significant number of opisthobranchs in the Red Sea. These were published by Nellie Eales (British Museum of Natural History, London) in 1938. Heller & Thompson (1983) published on part of the opisthobranch collections of the British Suakin Expedition, describing a number of new species. The remainder of the collection

16 Introduction

20. The University of Alexandria research vessel “Mabahiss” which made several expeditions throughout the Red Sea [courtesy of www.oceanography. egyguide.net/about_us.htm].

remains unpublished; C. Todd provided many original slides, and a few species have been located on www.seaslugforum.net. All are included as records in the checklist provided in this book. Barash and Danin worked from the Gulf of Eilat and documented the opisthobranch fauna, especially in comparison with migrations to and from the Red Sea (1972, 1977, 1980). Gohar and his colleagues, based at the Marine Station of Al Ghardaqa (also known as Hurghada) on the Egyptian coast, detailed the reproduction and developmental biology of numerous species (1957–1991). During the summer of 1983, the ‘Ibn Batuta’ (fig. 21) circumnavigated the Red Sea, with the express purpose of collecting bivalves, prosobranchs and opisthobranchs. This expedition culminated in a book (Sharabati 1984) and a series of taxonomic papers (Yonow 1986–2000). 21. m/v “Ibn Batuta”,

named after the great Arab explorer of the 14 th century (photo N. Yonow).

Introduction 17

THE OPISTHOBRANCHS Taxonomy and Biology The Opisthobranchia are highly evolved relatives of the gastropod sea snails: their evolution has led them to lose the shell completely in the most evolved orders, but representatives of each stage still survive today (fig. 22). They are all macroscopic, ranging from 2 mm to 500 mm in length, and all species are marine except a few species of tiny sand-dwell-

22. The five stages in the evolution of opisthobranchs, from the heavily shelled species to the naked ones. The area of each group approximates the number of species it contains (adapted from Willan & Morton 1984, drawn by Anna Ratcliffe).

18 Introduction

1 Animal still retains a spiral shell and possibly an operculum. 2 Animal has a reduced shell, displays enlargement of body. 3 Animal has a slug-like body, internal shell, single gill remaining. 4 Animal has bilaterally symmetrical body, no shell, secondary gills developing. 5 Digestive gland extending into cerata

ing Acochlida (fig. 23). There are more than 6000 species of opisthobranchs (Behrens & Hermosillo 2005); they may not number as many as the prosobranch snails, but they show more variety in their flamboyant appearance, behaviour, diet, and defensive adaptations. They inhabit coral reefs, boldly exposed, hidden under or between boulders, burrowing in soft sediments, or crawling on the surface. The earliest opisthobranchs were burrowing, infaunal animals; the head was flat and broad and the shell more streamlined in shape than in other gastropods, as in the present day acteonids (cephalaspideans, figs. 24). Most are soft-bodied and often brilliantly coloured, but have efficient defences against predators. Lack of a protective shell in the more advanced species has led to active methods of defence, an amazing array of body forms, and flamboyant colours and patterns to aid camouflage, mimic other toxic animals, or deter would-be predators with warning signals. No longer does the animal withdraw into its shell and wait for danger to pass, it actively acquires or produces these mechanical and chemical defences: calcareous sponge spicules, cnidarian nematocysts, toxic chemicals, acids, and luminescent products. Sea hares (fig. 26) can squirt a jet of purple ink when threatened. Pleurobranchs (fig. 30) secrete strong acids as concentrated as pH 1 from specialized epidermal cells of the mantle. Aeolid nudibranchs (fig. 38) store and use stinging cells (nematocysts) from the tentacles of their cnidarian prey, while dorid nudibranchs (figs. 33, 34) simply concentrate toxic substances from the sponges or other organisms upon which they feed.

23. A newly described species of acochlidian from the Gulf of Eilat, Hedylopsis ballantinei (from Sommerfeldt & Schrödl 2005).

25. Members of chelidonurids have a head shield, lateral parapodia, and a posterior shield containing a small shell.

24. A shelled cephalaspidean which has a reduced operculum.

26. A typical sea hare with two pairs of rolled anterior tentacles and large parapodia.

Introduction 19

Systematics

27. A thecosome with a conical shell, Clio convexa (photo V. Farstey).

The subclass Opisthobranchia is currently divided into ten orders (the higher taxonomy is in a state of flux), of which the five benthic orders are covered in this book. The following orders will not be dealt with further, although they do occur in the Red Sea. The Thecosomata and Gymnosomata are collectively known as pteropods, the name derived from the modification of the foot (“pod”) into paired swimming wings (“ptero”). The thecosomes (meaning “encased body”) or the sea butterflies are pelagic, bearing a conical (fig. 27), elongate, or spiral (fig. 28) shell or an internal ‘false’ shell, and wing-like extensions of the foot which enable movement in the plankton upon which they feed with a mucous net. Sporadically they suffer mass mortalities, and beaches can be awash with dead and dying specimens (fig. 29). The gymnosomes (“naked body”) have no adult shell and also live in the plankton, feeding mainly on the thecosomes! Finally, the Acochlida are tiny opisthobranchs, shell-less and smaller than 10 mm, living between sand grains and gravel (fig. 23).

28. A thecosome with a spiral shell, Limacia trochiformis (photo V. Farstey).

29. Mass mortality of

Creseis acicula in the Gulf of Eilat during January 2005 (photo V. Farstey).

Etymology The Latin names of most taxa, from classes to orders, families, genera, and species are derived from Greek and Latin because, until the nineteenth century, the language of science was Latin. Some are named after gods and goddesses, others for distinctive characters. The sub-class name Opisthobranchia is derived from the Latin, meaning rear (“opistho”) gills (“branchia”). This refers to a state of detorsion of the typical gastropod snail form, the result of which the gills became located at the back of the body; with this condition, the advanced opisthobranchs show both

20 Introduction

external and internal bilateral symmetry. The names of the orders Cephalaspidea and Anaspidea both originate from the common Greek root “aspidea,” meaning shield. The prefixes mean head and none, respectively, therefore “head shield” and “no shield”. The other three orders covered in this book, the Pleurobranchida, the Sacoglossa, and the Nudibranchia, differ in their roots. Pleurobranchida is derived from the Greek “plevro” for side (lateral); hence “pleurobranchus” means side gill. Sacoglossans (“glossa” in Greek means tongue, referring to the radular ribbon) store their used teeth in a sac, and the Nudibranchia are aptly named “naked gills” from the Latin because the vast majority bears the gills exposed and uncovered, and therefore unprotected, on the dorsum. Order Cephalaspidea Cephalaspidea are amongst the most primitive opisthobranchs, often retaining a shell and an operculum (fig. 24). However, during evolution, the shell has become less important, and some cephalaspideans have a reduced internal shell or none at all. The head is often flattened for burrowing in soft substrata, and is termed the cephalic shield. The shelled forms have a large foot, and those with internal shells have a posterior shield and lateral parapodia (fig. 25). They feed on foraminiferans, polychaetes, bivalves, and prosobranchs, as well as other opisthobranchs; only a few are herbivorous. The range in morphology of cephalaspideans is reflected in the large number of families included in the order. Order Anaspidea Anaspidea are opisthobranchs in which the shell may be external, internal, or absent, demonstrating once again that all the stages in their evolution have been successful. The body is generally elongate and high, with large parapodia which enable swimming in some species (fig. 26). The head is often obvious with two pairs of tentacles: both the rhinophores and oral tentacles are rolled and large, giving rise to the common name sea hares. Most species of anaspideans are herbivorous. Order Pleurobranchida Commonly known as the side-gilled opisthobranchs, the Pleurobranchida have a large plume-like gill located between the mantle and the foot on the right side of the body (fig. 30). The shell is internal, usually located centrally on a generally ovoid dorsum; one family has lost the shell completely. They generally have an oral veil with rolled oral tentacles and a pair of rolled rhinophores. Most pleurobranchs can secrete sulphuric

Introduction 21

acid as a defensive mechanism. The majority of species are found in the shallow sublittoral, but a few species occur deeper offshore. These slugs are grazing carnivores, feeding on sponges, tunicates, and other sessile invertebrates, and have strong jaws and a broad radula. Order Sacoglossa Sacoglossans, like the nudibranchs, are highly specialized opisthobranchs but are herbivorous, with only rare exceptions. They are relatively small and delicate creatures which may have an internal or external shell or none at all. The shell may be large and rounded, or small and cap-like. The most unusual sacoglossans have a bivalved shell as does Julia exquisita (fig. 31). Body form in the shell-less species varies almost as much as it does in the Nudibranchia: the animals can look like the elongated dorids, or like aeolids with large flattened cerata. However, these cerata do not bear cnidosacs, although they do support ramifications of the digestive gland. The majority bear parapodia, which may be upstanding or held close to the body (fig. 32). The most obvious external character-

30. A side-gilled opisthobranch with its gill showing between the mantle and the foot.

31. Julia, a bivalved sacoglossan.

32. A typical sacoglossan

Elysia with extended parapodia and a single pair of rolled tentacles anteriorly.

22 Introduction

istic distinguishing sacoglossans from nudibranchs are the paired enrolled rhinophores (when present); all nudibranchs bear tapered, lamellated, or branching rhinophores but never rolled ones. Oral tentacles are very small or absent, and the gills are lacking or not obvious. The radula is very distinctive, bearing a single tooth per row which is used for piercing single algal cells. The fluid contents of the algal cells are then sucked out with a muscular pharynx which acts as a powerful pump. The teeth are sharp and blade-like, used one at a time and produced in a continuous series. The worn teeth are stored in a sac when they detach from the ribbon. Many of these herbivorous opisthobranchs have a symbiotic relationship with the chloroplasts extracted from the algae. The chloroplasts continue to photosynthesize, providing extra nourishment, and can sometimes be seen in faint rows or patches inside the parapodia. Those species which do not feed on algae feed on the eggs of other opisthobranchs. Order Nudibranchia The nudibranchs are amongst the most beautiful and flamboyant of the gastropods, as well as the most highly evolved. They lack a shell as an adult although many planktotrophic species hatch with a larval shell, which is lost at metamorphosis when the animal assumes its adult body form (see fig. 40). Body form varies enormously and is related to the diet and habitat of each animal. Nudibranchs may be oval and flattened, long and thin, and with or without dorsal projections of many shapes, sizes, and functions. Oval flattened species tend to feed on encrusting sponges, barnacles, and ascidians. They are usually cryptic, and well camouflaged through shape, colour, texture, and behaviour. Some less active species restrict their diet to a single food organism only, most often a sponge, on which they become so closely dependent that they can be found partially imbedded in it. Nudibranchs which feed on erect arborescent organisms (hydroids, bryozoans, gorgonians, anemones) tend to be elongated and more active and aggressive. In most nudibranch species, however, diet is not so rigid, and they take a variety of prey organisms, although they often limit themselves to one particular group of animals, such as ascidians, anemones, barnacles, or hydroids. Food preference in a single species can change with age, geographical distribution, and bathymetric depth. In some suborders and many families, the body is dorsally papillate or covered with finger-like projections, the cerata; in two sub-orders, these are used to store nematocysts derived from their cnidarian prey which the opisthobranch then uses for its own defence. In other groups, the cerata contain extensions of the digestive gland only, defensive secretory glands, or calcareous spicules. The evolutionary trend in opisthobranchs has been a reduction in the importance of the shell, and increased emphasis on biological and

Introduction 23

chemical defences. This has become most noticeable in the nudibranchs, none of which have a shell as an adult. Many species extract mechanical and chemical defence mechanisms from their prey. Along with this has evolved the use of colour and pattern: many nudibranchs have evolved a precise resemblance to their prey organisms, to such an extent that some match their body colour to that of their food; some go so far as to become imbedded in their sponge or ascidian prey. Others use bright colours and patterns as warning signs, to advertise their noxious taste to would-be-predators. One can also find mimicry among nudibranchs, with non-poisonous species imitating the colour patterns of poisonous species: Aldisa sp. 2 (p. 156) resembles the noxious phyllidiids, for example. They are virtually immune to attack by fish or other predators, and with their soft agile bodies and chemosensory rhinophores, they are perfectly equipped to exploit every possible ecological niche. The order is subdivided into four sub-orders. The Doridina are the largest group of nudibranchs, generally with paired lamellated rhinophores located anteriorly and a clump of gills surrounding the anus posteriorly. Morphology varies and there are two loose groupings, the phanerobranchs (where the gills are not contractile into a pocket, fig. 33) and the cryptobranchs (gills contract completely into a pocket, fig. 34). The phanerobranchs often have extra appendages on the head and near the gills; these do not bear digestive gland ramifications and are termed papillae. They are thought to serve as protection for the vital organs, the rhinophores and gills. In addition to differences in morphology, the diet is different: phanerobranch dorids feed on a variety of invertebrates while the crypto-

33. A phanerobranch dorid with a reduced mantle and projections alongside the rhinophores and gills to detract predators from these vital organs.

35. Dendronotaceans have a paired series of branching gills along the body, as well as complicated rhinophores and a digitate oral veil.

24 Introduction

34. A typical cryptobranch dorid with an oval mantle, a gill circlet retracting into a single pocket, and often rhinophore sheaths also.

37. The vertically lamellated rhinophore of an arminid.

36. An arminid nudibranch with an oral veil and closely set rhinophores.

branch dorids consume sponges of all kinds. Doris was a Greek goddess, the daughter of Oceanus and Tethys in Greek mythology. The Dendronotina are characterized by a reduced mantle with paired branching gills along the back (fig. 35). The name is derived from the Greek “dendro” meaning tree, referring to the tree-like gills on the notum. The head has an extended oral veil, usually with digitiform papillae. The rhinophores are complicated, issuing from cupped, flared, or branching sheaths, and are themselves papillate or branched. Members feed on alcyonarian soft corals and cnidarians. The Arminina are a small, heterogeneous group of nudibranchs, very much in need of systematic revision (fig. 36). This sub-order contains nudibranchs which do not fit into any other category and hence morphology is extremely varied: species can resemble dorids and aeolids. If dorid-like, the rhinophores have vertical lamellae (fig. 37) and there is usually an expanded oral veil. Others have smooth tapering rhinophores and an undulating mantle edge, or, if aeolid-like, then show crinkly spiral lamellae on the rhinophores and the cerata are present all around the front of the animal (e.g. Madrella, p. 243), a situation never found in the aeolids. The Aeolidina are elongated, and the dorsum is covered in cerata of various shapes and sizes (fig. 38). These contain extensions of the digestive gland and have a specialized structure at the tip, the cnidosac, in which the nematocysts are stored (fig. 39). They have no gills as such and respiration occurs by diffusion, probably through the cerata. As well as sensory rhinophores, the long oral tentacles are also sensory in function, and propodial tentacles are generally present. Some species can move rapidly and others are aggressive. The name Aeolidina is derived from Greek mythology: Aeolus was custodian of the four winds.

Introduction 25

38. A typical aeolid nudibranch with long rhinophores and oral tentacles, and its dorsum covered in finger-like cerata.

39. A single ceras showing the digestive gland leading to the cnidosac at the tip to store unexploded nematocysts.

Defence With the loss of a protective shell and the necessary development of active methods of defence, body shape, colour, and pattern have assumed a huge variety of forms. More simple mechanical defence is exhibited by some of the sponge-eating dorids (e.g. Platydoris, Jorunna, Discodoris) which incorporate the calcareous spicules into their dorsal epidermis. These species can be hard to the touch, rough, or spiky. Other dorids have incorporated the sponge toxins into their epidermis and are thereby more colourful, such as the chromodorids and phyllidiids. Aeolids feeding on anemones, hydroids, and jellyfish have special cerata on their dorsum to store nematocysts (e.g. Pteraeolidia, Baeolidia). When in danger, the cerata bristle, pointing to the source of danger, and the nematocysts are released. Plocamopherus emits flashes of light from the knobs on its mantle margin when swimming away from danger. The acids and chemicals found in opisthobranchs have attracted much recent research, and it appears that each species has its own complement of complex toxins. These natural products may be synthesized by the opisthobranch or acquired, concentrated, and/or toxified by the animal from its prey species. The toxins serve different functions such as, amongst others, feeding deterrence and communication (e.g. alarm or reproductive state). The products are often located in special extensions of the body which are dispensable, such as processes near the rhinophores and gills; these extensions are usually brightly coloured, detracting predators from vital organs, and they are often regenerated when nipped off (e.g. Ceratosoma). Others have glands around the edge of the mantle in which the products are stored (e.g. Chromodoris, Hypselodoris) while still others have biologically active substances generally located in the dorsum (e.g. Phyllidia, Phyllidiella), the region exposed to potential predators. These secretions have been found to occur in the five orders covered in this book. In the non-nudibranchs, the secretions are products of the animal’s own metabolism whilst in the nudibranchs, the chemicals and nematocysts are derived from the diet organisms. Respiration Respiration mechanisms differ greatly in the opisthobranchs. Some primitive species still possess a gill in the mantle cavity, such as the shelled Cephalaspidea. In the Anaspidea, the gill is barely protected by the mantle and can be seen in some species, while in the Pleurobranchida it is clearly visible on the right side between the foot and the mantle. However, in most of the Nudibranchia, newly developed dorsal structures function as gills (“branchia”). In the Aeolidina, respiration is effected by osmosis through the cerata: there are no specially developed respiratory structures. In the Phyllidiidae and some Arminina, secondary gill lamellae have developed and are located ventrally, lying between the foot and the mantle.

26 Introduction

Reproduction and development All opisthobranchs are hermaphrodite: thus, the reproductive organs of male and female are both present in adult specimens, and are usually positioned closely together and jointly referred to as the gonad or ovotestis. The gonad is located on the right-hand side of the body, and the gonopore, where exchange of sperm is effected, is also on the right near the head. It is often visible in dorids, for example, where the colour pattern may be deflected around it, or it may be present as a prominent swelling as in some aeolids. Individuals are capable of fertilizing each other; after fertilization, the animals separate and crawl away to lay their spawn on suitable substrata. Many non-nudibranchs mate in chains, the posterior animal acting as male to the one in front, which in turn acts as male to the one in front of it. Spawn is characteristic for each species, in shape and form, colour, size, and arrangement of eggs within. The eggs hatch either into planktonic veligers, a swimming and feeding larval phase which then settles and begins benthic life, or into a minute crawling juvenile (fig. 40). Larval life and mode of development are related to the type of egg; eggs and larval shells have been classified, and are relat-

Introduction 27

40. The development of a pelagic lecithotrophic species, Paradoris erythraeensis [upper branch] which loses its shell at seven days and metamophoses into a juvenile nudibranch and Glossodoris cinta [lower branch] which has direct development and loses its cap-like shell in its egg case at 22 days old before metamorphosing into a juvenile (adapted from Gohar & Aboul-Ela 1959 and Gohar & Soliman 1967d, drawn by Anna Ratcliffe).

ed to life strategies. The lifespan of opisthobranchs is generally less than one year but more primitive species, larger species, species which live in temperate and cold water, and species living at depths may live longer, but probably not more than a few years. The longest-lived tropical opisthobranch is Dolabella auricularia (p. 101), living for five years in an aquarium in Hawaii (Hadfield & Switzer-Dunlap 1984), a similar lifespan to the Antarctic species Philine gibba (Seager 1982). Diet and feeding Feeding has become a specialized function among opisthobranchs. Although most species are active predators, several opisthobranch families feed on the eggs of cephalopods or teleost fish, as well as eggs of gastropods and even other opisthobranchs. Oophagous opisthobranchs, including nudibranchs and sacoglossans, are usually slim and pale so as to pass unnoticed among cream and white coloured eggs. Some have even lost their excretory system, as they selectively feed on young eggs and there is no waste. Generally, the cephalaspids, pleurobranchs, and nudibranchs are carnivorous, while the anaspids and most sacoglossans feed on algae.

41. The jaws of

Marioniopsis rubra (from Yonow 2000).

Most opisthobranchs have a pair of jaws (fig. 41) and a radula (fig. 42) for this purpose; both are made of chitin. The radula is a ribbon bearing few or numerous rows of teeth, the number and shape of which varies between orders, families, genera, and even species. In both grazing herbivores and carnivores, the radula is broad (many teeth per row) and long (many rows). The radula is long and narrow in species such as Flabellina and Nembrotha feeding on erect hydroids and bryozoans. The sacoglossans have a single sabre-like tooth in each row, used for piercing single algal cells. Species without a radula include Philinopsis, which engulfs its opistho-

28 Introduction

branch prey whole; Melibe, which captures small crustaceans with an enlarged tentacular hood around its mouth; and the porostome genera, such as Phyllidia, Fryeria, and Dendrodoris, which suck sponge tissue after digesting it externally. The many feeding strategies are a direct result of shell loss and concomitant exploitation of marine habitats. Locomotion Opisthobranchs, like most shelled gastropods, creep along the substratum on the sole of the foot. This movement is brought about by the control of smooth muscle fibres lying longitudinally and dorso-ventrally in the foot. Control is so precise that it allows movement of localized areas of the foot: for example, Aplysia sp. (p. 100) and Chelidonura flavolobata (p. 76) can lift the head and remain attached by the tail. Most opisthobranchs move very slowly, remaining where food is abundant if they are undisturbed. However, many species can crawl at speed (for a sea slug!), such as Chelidonura sandrana (p. 80) and many aeolids. In addition, when alarmed or when finding themselves in an unsuitable region, many species can swim, a behaviour only possible when the shell is buoyant, internal or absent. Swimming opisthobranchs are a spectacular sight, the movement often accompanied by flashes of colour. Motion is usually effected by dorso-ventral or lateral flexions of the body. Species in the Red Sea which can swim include Hexabranchus sanguineus (fig. 43 and p. 144), Nembrotha megalocera (p. 137), Plocamopherus indicus (p. 140), Sebadoris nubilosa (p. 153), Melibe rangii (p. 237), and Notarchus indicus (fig. 44 and p. 105). It is likely that swimming has evolved as an escape reaction in some species and a defensive reaction in others. Once perfected, this swimming behaviour benefited a useful secondary function, such as for

Introduction 29

42. Sections of the median and lateral teeth of the radula of Marioniopsis viridescens (from Yonow 2000).

43. Swimming in

Hexabranchus marginatus, showing the undulating wave passing along the mantle margin (from Edmunds 1968).

44. Jet propulsion in

Notarchus indicus, studied in the Gulf of Eilat (from Schuhmacher 1973b).

feeding or forming breeding aggregations, or simply a method of rapid displacement to a more suitable substrate. Notarchus indicus, common in the sea grass beds of the northern Red Sea, has developed a novel method of locomotion, jet propulsion (fig. 44). Its parapodia have fused along most of their lengths to form a sac with an anterior opening, through which water is drawn in and then propelled out rapidly, causing the animal to summersault into the water column (Schuhmacher 1973b).

30 Introduction

Mimics Polyclad flatworms are often misidentified as opisthobranchs, and a selection of mimics is included in this book. They belong to the phylum Platyhelminthes (from the Greek for flat and worm) and the class Turbellaria, the free-living flatworms. They are extremely primitive, and have a mouth and central gut branching to each side, but no anus. Generally they are very flattened, but many marine tropical species, especially those mimicking opisthobranchs, are able to produce the body into a pair of sensory “tentacles” anteriorly, and some have pustules. The tropical ones, belonging to the Polycladida, and indeed some temperate

Maritigrella cf. fuscopunctata JK

Pseudoceros scintillatus JK

Pseudoceros bimarginatus JK

species, are extremely colourful. Almost all species are marine, usually benthic, and common in shallow water habitats and coral reefs. Their bright colours have evolved for defence against fish, their main predators which have excellent eye sight. As in many opisthobranchs, colour has evolved with the development of toxic or distasteful chemicals, and there may be a benefit in advertising these compounds. Just as there are groups of nudibranchs with similar warning colourations, there are

Introduction 31

Pseudoceros scriptus JK

Pseudoceros cf. dimidiatus JK

Pseudoceros sp.

JK

Pseudoceros sp.

JK

flatworms with patterns similar to the toxic nudibranchs. So, not only are the flatworms mimicking toxic animals for protection, but they also produce their own virulent cocktails (Schupp et al. 1999). A selection of species occurring in the Red Sea which resemble and can be mistaken for opisthobranchs are presented in this book. They are included with the species they are mimicking. Collection of Opisthobranchs In the Red Sea, opisthobranchs are most easily located in the intertidal and shallow sublittoral waters on ‘hard’ shores, on the reef tops and along the coral reefs. The sandy beaches, unless colonized by sea grasses, are too exposed, and the rocky shores are too steep, being almost vertical. In the intertidal and on the reef flat, the best places to look are underneath, on, and around large rocks and boulders, remembering to replace EVERYTHING back to its original position : animals and plants

32 Introduction

living on the upper surfaces of rocks are different from those beneath, and if the rocks are left turned over, both communities will die, as well as those animals sheltering in the particular under-boulder habitat. On the coral reefs, where most rocks and corals are cemented together, opisthobranchs can still be spotted with a keen eye. Ledges and cave floors littered with coral rubble are also good places to look. The only way to preserve the colours and patterns of opisthobranchs is to photograph them and prepare detailed drawings. In preservative, most species eventually lose shape and colour, resulting in amorphous blobs of interest only to the scientist. The animals are best studied when alive, with notes on size, colours, habitat, and behaviour, and then returned to their habitat. Snorkellers and divers seriously interested in the rare or undescribed species in this book can be of great value to the scientist: specimens are necessary to establish the names of species, with their deposition in museums as types. These types form the international reference collections with which all subsequent specimens are compared in order to establish their identity. After detailed notes and drawings have been made, opisthobranchs should be relaxed in 7% magnesium chloride in sea water. Alternatively, they can be refrigerated and then frozen in sea water, but this is not so efficient. When the animal does not react to prodding, it is best preserved in 4–5% formaldehyde, adding it drop by drop to a minimum amount of sea water containing the opisthobranch. Remember that pure formalin is 40% solution in water of the gas formaldehyde, so you will be preserving your animals in a 10% solution of formalin. Beware – it is an unpleasant and toxic solution. If formalin is unavailable, 70% alcohol is adequate. Drawings, field notes, photographs and specimens can be sent to the author, who will be happy to assist you in identification and techniques (Department of Biological Sciences, University of Swansea, Singleton Park, Swansea SA2 8PP, Wales, United Kingdom; [email protected]).

Introduction 33

NINETEENTH & EARLY TWENTIETH CENTURY ENGRAVINGS OF OPISTHOBRANCHS SAVIGNY PLATES Savigny, J. C. 1817. Description de l’Egypte, ou receuil des observations et des recherches qui ont été faites en Egypte pendant l’Expédition de l’armée française, publiée par ordre du gouvernement. Histoire Naturelle. Imprimerie Royale, Paris. Planches, vol. II. Audouin, J. V. 1826. Explications sommaire des planches de Mollusques de l’Egypte et de la Syrie publiées par J. C. Savigny... Histoire Naturelle, Animaux Invertébrés 1 (4): 7–56. Savigny’s engraved plates were prepared from the original coloured “albums” or velums in great detail by the best artists. The mollusc plates were destined to be described by Audouin who was an ornithologist, and he provided only very short explanations. The shelled species are not covered here but are on the checklist. The engravings reproduced here only illustrate the new species; they stand as the types as no material remains in the Museum National d’Histoire Naturelle in Paris. AUDOUIN’S NAMES

CURRENT NOMENCLATURE

Bursatella savignana Bursatella leachii savignana Audouin: Bursatella leachii savignana is circumglobal, and has entered the Mediterranean via the Red Sea. Despite this, it has not been found in the Red Sea recently. It reaches 100 mm in length and is patchy in colour, brown green with reticulations and clear green areas with a central blue spot; the body may be covered in long papillae. It is also figured on Rüppell & Leuckart’s engraving (p. 41) as a new species, Notarchus lacinatus. Pleurobranchus oblongus

Pleurobranchus albiguttatus (Bergh), p. 110

Doris concentrica ? Doriopsis granulosa Pease: There is a strong possibility that Audouin’s species is Doriopsis granulosa. The species is characterized by a flap over the gill cavity with the gills arranged horizontally. This is clearly intentional in the engraving as the gills of other species of Doris are 3-dimensional and arranged in a circlet. Doriopsis granulosa is covered in evenly sized tubercles, which are not necessarily as ‘concentric’ as depicted in the engraving. Colour is extremely variable, from shades of yellow through to orange, green, and brown. Doriopsis granulosa is recorded from the Indo-West Pacific. Doris marmorata

Hypselodoris infucata (Rüppel & Leuckart), p. 193

Doris immaculata Dendrodoris immaculata (Audouin): I can find nothing in the scientific literature referring to Doris immaculata; the engraving appears to be of a species of Dendrodoris.

34 Nineteenth & early twentieth century engravings of opisthobranchs

Savigny plates [from Bouchet & Danrigal 1982]

Nineteenth & early twentieth century engravings of opisthobranchs 35

Doris tigrina Dendrodoris tigrina (Audouin): It has been decided to retain this species, although it was not present in these collections. Rüppell & Leuckart recognized it, but for some strange reason named their specimen D. punctata, which is here regarded as a synonym. Abraham (1877) also recognized them as similar. The body is elongate oval, pale or white in colour and covered in large and small black spots, and cream spots. The margin is produced in a ruffled skirt. [Clearly there are no elongated tentacles in Dendrodoris as in the engraving.] Tritonia elegans

Tritoniopsis elegans (Audouin), p. 232 EHRENBERG PLATE Ehrenberg, C. G. 1828, 1831. Decas 1. Mollusca. Symbolae physicae, seu icones et descriptiones animalium e vertebraterum sepositis insectis quae ex itinere per Africam Borealum et Asiam Occidentalem – nov ae ant illustratae redierunt (pages unnumbered) (plates 1828, text 1831). Ehrenberg documented, amongst many plants and animals, the molluscs of North Africa and the Red Sea in this volume. The work was produced in two parts, the plates in 1828 and the text in 1831. The colour plate of opisthobranchs, reproduced here for the first time, bears the names of three new species. Cryptophthalmus olivaceus is valid under the new rules of the ICZN (1999) but Hexabranchus praetextus is not. The sacoglossan Stiliger ornatus is also considered valid but has not been recorded from the Red Sea since.

EHRENBERG’S NAMES

CURRENT NOMENCLATURE

Fig. I

Hexabranchus praetextus

Hexabranchus sanguineus (Rüppell & Leuckart), p. 144

Fig. II

Cryptophthalmus olivaceus

Phanerophthalmus olivaceus (Ehrenberg), p. 94

Fig. III

Stiliger ornatus Ehrenberg: This species has not been collected again from the Red Sea, but has been recorded from Mozambique, Japan, and Heron Island (Great Barrier Reef), although Japanese descriptions differ from the type description. The animal is rather beautiful with yellowish ground colour and black-banded cerata. The head is also marked with striking black pigmentation.

36 Nineteenth & early twentieth century engravings of opisthobranchs

Ehrenberg plate [courtesy of The Natural History Museum, London]

Nineteenth & early twentieth century engravings of opisthobranchs 37

RÜPPELL AND LEUCKART PLATES Rüppell, W. P. E. S. & Leuckart, F. S. 1828, 1830. Mollusca. In: Atlas zu der Reise im nordlichen Afrika von Eduard R üppell. Zool. Neue wirbellose Thiere des Rothen Meeres, pp. 15–22 and Tab. 4, 5 (1828), pp. 23–47 and Tab. 7–11 (1830). Difficulties surround the publication dates of the many new species described by Rüppell & Leuckart, which was published in parts, despite Winckworth’s (1942) detailed paper. Pages 15–22 were published in 1828, while pages 23–47 were published in 1830. This is important as there are clashes with Ehrenberg’s publication on opisthobranchs, the plates with names dated 1828 and the text in 1831. Both publications name the same two new species; because Ehrenberg placed names on his colour plate and they are clearly recognizable, they should have priority, as does Phanerophthalmus olivaceus. However, Hexabranchus sanguineus Rüppell & Leuckart, 1830 takes precedence over Hexabranchus praetextus Ehrenberg, 1828 because the name has been used so frequently in the literature (ICZN 1999). Despite brief descriptions, nearly all the species illustrated by Rüppell & Leuckart in these plates have been identified and are illustrated in this book. RÜPPELL & LEUCKART’S NAMES

CURRENT NOMENCLATURE

fig. 1 fig. 2 3

Tritonia rubra Tritonia glama Tritonia cyanobranchiata

Marioniopsis rubra (R & L), p. 229 Marionia glama (R & L), p. 225 Marioniopsis cyanobranchiata (R & L), p. 226

Tab. 5

fig. 1 fig. 2 fig. 3

Pleurobranchus citrinus Pleurobranchus forskalii Plocamopherus ocellatus

Berthellina citrina (R & L), p. 109 Pleurobranchus forskalii (R & L), p. 112 Plocamopherus ocellatus (R & L), p. 142

Tab. 7

fig. 1 fig. 2

Tab. 8

fig. 1 fig. 2 fig. 3

Tab. 4 fig.

Aplysia argus Aplysia dactylomela Rang, p. 96 Notarchus laciniatus Bursatella leachii savignana Audouin: See Savigny’s plate for explanations on this species (p. 34). Doris sanguineus Hexabranchus sanguineus (R & L), p. 144 Doris fumata Dendrodoris fumata (R & L), p. 211 Doris albo-limbata Dendrodoris nigra (Stimpson), p. 212 This species appears to be a species of Dendrodoris. It is most similar to Dendrodoris nigra, as well as similar to Vayssière’s (1912) Doriopsis rosea, which has been synonymized with Dendrodoris fumata; however, Doris albo-limbata was not mentioned in the revision by Brodie et al. (1997). Dendrodoris nigra is more frequently encountered in Red Sea collections than Dendrodoris fumata, and Rüppell & Leuckart most probably recorded it as D. albo-limbata. The size of the gill circlet and the colour is also very suggestive, and it is regarded as a synonym of D. n igra.

38 Nineteenth & early twentieth century engravings of opisthobranchs

Tab. 4 Rüppell & Leuckart [plates 4 and 5 courtesy of Leone Lemmer, The Australian Museum; plates 7–11 courtesy of Daniel Geiger, Santa Barbara Museum of Natural History, California]

Nineteenth & early twentieth century engravings of opisthobranchs 39

Tab. 5 Rüppell & Leuckart

40 Nineteenth & early twentieth century engravings of opisthobranchs

Tab. 7 Rüppell & Leuckart

Nineteenth & early twentieth century engravings of opisthobranchs 41

Tab. 8 Rüppell & Leuckart

42 Nineteenth & early twentieth century engravings of opisthobranchs

Tab. 9 Rüppell & Leuckart

Nineteenth & early twentieth century engravings of opisthobranchs 43

Tab. 10 Rüppell & Leuckart

44 Nineteenth & early twentieth century engravings of opisthobranchs

Tab. 11 Rüppell & Leuckart

Nineteenth & early twentieth century engravings of opisthobranchs 45

Tab. 9

Doris punctata Dendrodoris tigrina (Audouin): The body form is identical to that of several species of Red Sea dendrodorids. The colour is grey with black and white spots, which is similar to D. tigrina (Audouin) and D. n igropunctata (Vayssière). According to Pruvot-Fol (1933), Rüppell & Leuckart recognised Audouin’s Doris tigrina as the identity of their species but strangely, gave it a new name. Abraham (1877) also considered these two species the same.

fig. 1

fig. fig. fig.

2 3 4

fig.

5

Doris quadricolor Doris obsoleta Doris tinctoria Doris pulchella

Chromodoris quadricolor (R & L), p. 176 Chromodoris obsoleta (R & L), p. 184 Chromodoris tinctoria (R & L), p. 184 Risbecia pulchella (R & L), p. 201

Tab. 10 fig. fig. fig.

fig. 1 2 3 4

Doris pallida Doris impudica Doris infucata Eolida bella

Glossodoris pallida (R & L), p. 191 Gymnodoris impudica (R & L), p. 135 Hypselodoris infucata (R & L), p. 193 ? Phyllodesmium magnum Rudman, p. 263

Tab. 11 fig.

fig. 1 2

Phyllidia pustulosa Bulla smaragdina

Fryeria rueppeli (Bergh), p. 215 Phanerophthalmus olivaceus (Ehrenberg), p. 94

VAYSSIÈRE PLATE AND LINE DRAWINGS Vayssière, A. 1912. Recherches zoologiques et anatomiques sur les Opisthobranches de la Mer Rouge et du Golfe d’Aden, partie 2. Annales de la Faculté des Sciences, Marseille 20: 5–157. Vayssière worked on collections from the Red Sea and the Bay of Tadjoura (Gulf of Aden) made by two Frenchmen. He described many opisthobranchs, but only illustrated one plate in colour. The other plates are of anatomical characters, with a scattering of whole animals. His descriptions are much fuller than Rüppell & Leuckart’s or Ehrenberg’s, and many species are recognized and recorded in this book. VAYSSIÈRE’S NAMES fig. 1

CURRENT NOMENCLATURE

Casella atromarginata Glossodoris atromarginata (Cuvier): Vayssière described G. atromarginata from the Gulf of Aden; subsequently both White (1951) and Marcus & Marcus (1960) recorded this species from the Red Sea. White described both G. cincta and G. atromarginata in her collections, while Marcus & Marcus discuss at length the differences between the two species. Despite this, I do not think G. atromarginata occurs in the Red Sea; my own extensive collections and my photographers’ continuing searches have revealed utterly no records to date. Both White and the Marcuses collected in the Indian Ocean, and it is possible that some error occurred in the labeling or sorting of specimens.

46 Nineteenth & early twentieth century engravings of opisthobranchs

Vayssière plate [courtesy of The Natural History Museum, London]

Nineteenth & early twentieth century engravings of opisthobranchs 47

fig. 2 fig. 3 fig. 4, 5 fig. 6 fig. 7 fig. 8 fig. 9 fig. 10

Doriopsis rosea Dendrodoris fumata (Rüppell & Leuckart), p. 211 Doriopsis nigra Dendrodoris nigra (Stimpson), p. 212 Trevelyana striata Analogium striatum (Eliot), p. 129 Orodoris miamirana Orodoris miamirana Bergh, p. 207 Halgerda willeyi Halgerda willeyi Eliot, p. 163 Chromodoris runcinata Hypselodoris infucata (Rüppell & Leuckart), p. 193 Plakobranchus ocellatus Plakobranchus ocellatus van Hasselt, p. 124 Plocamopherus indicus Plocamopherus indicus Bergh, p. 142 This appears to be the only subsequent record of Bergh’s species in the literature. Bergh did not illustrate his new species, but Vayssière did, comparing his specimen with Bergh’s description and Alder & Hancock’s description and excellent illustrations of P. ceylonicus Kelaart. There are certainly differences between Vayssière’s specimen and P. ceylonicus as described by Thompson (1975), but there are also many similarities. Both have an elongated, tuberculate body with dendritic appendages; in both colour is pale orange or pale brown, and both have either a yellow foot or yellow markings. Differences include three tripinnate gills in P. indicus but four in P. ceylonicus, one pair of knobs and two or more respectively, rhinophores with many fine lamellae in the first species and only 24 in the second, with a raised irregular sheath in P. indicus and slightly raised smooth edge in P. ceylonicus. If the variability of P. ceylonicus includes these characters, the two can be synonymized. Meanwhile, they are retained as separate species until further material can be examined.

fig. 11,12 fig. 13 fig. 14,15

Hallaxa indecora Chromodoris tenui-linearis Phyllidia dautzenbergi

Hallaxa indecora (Bergh), p. 150 Hypselodoris nigrostriata (Eliot), p. 197 Phyllidiopsis dautzenbergi (Vayssière), p. 222

fig. 78

Tritoniopsis gravieri Vayssière: The single tiny specimen, 11 mm preserved, was collected from the Gulf of Aden: the description closely resembles that of M. cyanobranchiata, yellow or pale orange with some black patches in the middle of the pustulate dorsum, and pale blue gills along the sides. The gills alternate in size, but this may be due to the small size. The radular teeth are not dissimilar, taking into account the size of this specimen. White (1951) subsequently recorded this species also from the Gulf of Aden.

fig. 105

Doriopsis jousseaumei Dendrodoris jousseaumei (Vayssière): Doriopis has long been regarded a synonym of Dendrodoris. This species was described from twenty preserved specimens (10–18 mm) collected from the Gulf of Aden; one would surely expect it to be recorded again. Vayssière was very meticulous in his descriptions of the dendrodorids, and yet I can find nothing similar in the literature on opisthobranchs from the Indian Ocean or Red Sea.

fig. 117

Bornella digitata

Bornella stellifer (Adams & Reeve), p. 233

48 Nineteenth & early twentieth century engravings of opisthobranchs

fig. 144

Gravieria rugosa Sclerodoris rugosa (Vayssière): There are no pits which would identify this species with S. tuberculata; nor do the tubercles have long extensions like S. apiculata. It is described as hyaline white, with opaque yellowish white lines along the crests. The yellow white lines are composed of a multitude of small yellow granulations. The crests are very low, with a longitudinal central one, and the mantle ample and elliptical. The rhinophore sheath is finely denticulate, the six bipinnate gills could be tripinnate. This appears so very similar to a photograph of an uncollected species on p. 274.

Vayssière line drawings

Nineteenth & early twentieth century engravings of opisthobranchs 49

PRUVOT-FOL PLATE Pruvot-Fol, A. 1933. Mission Robert Ph. Dollfus en Egypte. Opisthobranchiata. Memoires de l’Institut d’Eg ypte 21: 89–159. Although this is the most recent publication on the ‘old’ expeditions to the Red Sea and their respective collections of opisthobranchs, it is also the most difficult. Many of the five new species and other species are still unknown or unrecognizable, and there are many typographical and other errors in her work. PRUVOT-FOL’S NAMES

CURRENT NOMENCLATURE

fig. I

Asteronotus Hemprichi

Asteronotus cespitosus (van Hasselt), p. 162

fig. II, III

Ceratosoma cornigerum

Ceratosoma tenue Abraham, p. 203

fig. IV–VI

Armina semperi (Bergh) var. erythraea: There is no recent work on this rather difficult group of nudibranchs. This is most probably Armina semperi and the lack of the beautiful blue on the foot and oral veil is due to preservation. The radula appears to be distinctive for each species that has been well studied, and Pruvot-Fol’s match Bergh’s and Eales’ descriptions well enough. The largest of four specimens measured 70 mm (in preservative) and all were collected by dredging. (Eales 1938 had 21 specimens from the Gulf of Aden.)

fig. VII, VIII

Glossodoris dollfusi

fig. IX

Glossodoris ? albomaculata (Pease) var. erythraea ? Chromodoris obsoleta (R & L), p. 184 Described from ten preserved specimens, possibly darker than pale yellow or white, with bright white spots, a bright yellow margin, and black sub-marginal band. The dorsum and foot bear unevenly sized, rather large, rounded white spots which are opaque and slightly raised. There are 11–13 gills of unequal sizes; the radular formula is undetermined. Pease’s species has not been rediscovered, nor has Pruvot-Fol’s. However, in 1951, she suggested that these might be synonyms of C. obsoleta. Chromodoris obsoleta is the most similar species in the Red Sea, with a deep purple submarginal and yellow orange marginal banding; the fawn reticulations can produce dense white patches (she illustrates spots). The ten specimens from Suez indicate a fairly common species, as is C. obsoleta in the Red Sea today.

fig. X

Glossodoris quadricolor

fig. XI

Philine Vaillanti Issel: Currently the only species of Philine to be recorded in the Red Sea; older records are from bottom dredge samples. Pruvot-Fol says it is very like Philine aperta (Linnaeus) from the North-East Atlantic, but pale salmon in colour and “much larger”. Shells can be found washed ashore after storms in the Gulf of Eilat (J. Dafni, pers. comm.).

Hypselodoris dollfusi (Pruvot-Fol), p. 192

Chromodoris quadricolor (Rüppell & Leuckart), p. 76

50 Nineteenth & early twentieth century engravings of opisthobranchs

Pruvot-Fol plate [courtesy of The Natural History Museum, London]

Nineteenth & early twentieth century engravings of opisthobranchs 51

CHECKLIST OF SPECIES RECORDED FROM THE RED SEA

This list is (hopefully) complete for all species recorded in the scientific literature from the Red Sea. They include expedition reports, biological and ecological observations, and chemical studies on single species, as well as a number of species photographed more than once but for which no material has been collected; some of these have been confidently assigned to a species or genus, but photographs of the others not listed here can be found in the chapter entitled “Incertae Sedis” (p. 271). The species printed in bold type face are illustrated in this monograph, those with asterisks * * * represent new records for the Red Sea, and the underlined species are endemic for the Red Sea. Those species which are found in the Red Sea and in the Gulf of Aden (but not in the western Indian Ocean) are considered endemic, while those which were only recorded from the Gulf of Aden but not the Red Sea are not considered endemic. The species which have a plus sign + after the recorder need confirmation or further study. ORDER CEPHALASPIDEA

ACTEONIDAE Acteon sp. Pupa affinis (A. Adams, 1855) Pupa tessellata (Reeve, 1842) Pupa cf. solidula (Linnaeus, 1758) Pupa sp. HYDATINIDAE Hydatina physis (Linnaeus, 1758) Hydatina zonata (Lightfoot, 1786)

RINGICULIDAE Ringicula acuta Philippi, 1849

Lamy 1938 as A. flammeus + Lamy 1938 Sharabati 1984 as P. sulcata, Taylor & Reid 1984 Sturany 1903, Mastaller 1979, Sharabati 1984 all as P. solidula (uncommonly washed up as shells, Eilat, J. Dafni) Sturany 1903 and Mastaller 1979 both as P. sulcata +

Sturany 1903, Vayssière 1906, Mastaller 1978, 1979, Sharabati 1984, Yonow 2000 Vayssière 1906 and Eales 1938 (Gulf of Aden) both as H. velum, Barash & Danin 1972 and Mastaller 1979 both as Hydatina amplustre (listings only)

Issel 1869 (Gulf of Aden), Mastaller 1979, Audouin 1826 as R. savignyi, Yonow 1992 as Ringicula sp. (commonly washed up as shells, Eilat, J. Dafni)

DIAPHANIDAE Colpodaspis thompsoni Brown, 1979

Heller & Thompson 1983

SCAPHANDRIDAE Acteocina fusiformis (A. Adams, 1850)

Audouin 1826 and Issel 1869 both as Tornatina

52 Checklist of species recorded from the Red Sea

Acteocina inconspicua (H. Adams, 1852) Acteocina involuta (Nevill, 1871) Acteocina simplex (A. Adams, 1850) Utriculastra mucronata (Philippi, 1849) CYLICHNIDAE Cylichna mongii (Audouin, 1826) Cylichna villersii (Audouin, 1826) Roxania lithensis (Sturany, 1903) RETUSIDAE Pyrunculus fourierrii (Audouin, 1826) Retusa desgenettii (Audouin, 1826) Retusa tarutana Smythe, 1979 Ventomnestia girardi (Audouin, 1826) PHILINIDAE Philine vaillanti Issel, 1869

olivaeformis, Tomlin 1927 as A. olivaeformis, Lamy 1938, Mastaller 1979 (commonly washed up as shells, Eilat, J. Dafni) Lamy 1938 as Tornatina Dekker & Orlin 2000 Lamy 1938 and Mastaller 1979 both as Tornatina Dekker & Orlin 2000 Audouin 1826 as Bulla, Issel 1869, Lamy 1938, Barash & Danin 1972, Mastaller 1979 Audouin 1826 as Bulla, Issel 1869 Sturany 1903 as Atys (deep water) Audouin 1826 as Bulla, Issel 1869 as Cylichna mica Audouin 1826 as Bulla, Issel 1869 as Cylichna pulvisculus Dekker & Orlin 2000 Audouin 1826 as Bulla, Lamy 1938, Mastaller 1979 [= V. bizona A. Adams, = V. villosa (Gould)]

Issel 1869, Sturany 1903, O’Donoghue 1929, Pruvot-Fol 1933 (commonly washed up as shells, Eilat, J. Dafni)

AGLAJIDAE Chelidonura flavolobata Heller & Thompson, 1983 Heller & Thompson 1983, Yonow 1990, 2000 Chelidonura fulvipunctata (Baba, 1938) * * * Chelidonura livida Yonow, 1994 Engel & van Eeken 1962 (part) as Aglaja cyanea (nigra?) , Yonow 1990 as C. africana, Yonow 1994a, 2000 Chelidonura punctata Eliot, 1903 *** Chelidonura sandrana Rudman, 1973 Yonow 1994a Philinopsis cyanea (Martens, 1879) Engel & van Eeken 1962 (part) as Aglaja cyanea (nigra?) , Mastaller 1979, Yonow 1990, 1992, 2000 Philinopsis reticulata (Eliot, 1904) Yonow 1990 as Aglaja phaeoreticulata (syn. nov.) Odontaglaja guamensis Rudman, 1978 *** GASTROPTERIDAE Sagaminopteron cf. ornatum Tokioka & Baba, 1964 Yonow 2000 Sagaminopteron cf. psychedelicum Carlson & Hoff, 1974 Yonow 2000 Siphopteron sp. *** HAMINOEIDAE Atys cylindricus (Helbling, 1779)

Sturany 1903, Vayssière 1906, Eales 1938, Lamy 1938, Checklist of species recorded from the Red Sea 53

Mastaller 1978, 1979, Sharabati 1984, Taylor & Reid 1984 (commonly washed up as shells, Eilat, J. Dafni) Atys ehrenbergi (Issel, 1869) Issel 1869 as Alicula (fossil shell), Sharabati 1984 as A. cf. naucum (uncommonly washed up as shells, Eilat, J. Dafni) Atys naucum (Linnaeus, 1758) Sturany 1903, Vayssière 1906, Eales 1938, Mastaller 1979 Atys semistriatus Pease, 1860 Heller & Thompson 1983 Diniatys dentifer (A. Adams in Sowerby, 1850) Tomlin 1927, Barash & Danin 1972 (both Gulf of Suez) Diniatys dubius (Schepman, 1913) *** Haminoea cyanocaudata Heller & Thompson, 1983 Heller & Thompson 1983 Haminoea cyanomarginata Heller & Thompson, 1983 Heller & Thompson 1983 Haminoea pemphis (Philippi, 1847) Vayssière 1906, Lamy 1938 as H. tenera (commonly washed up as shells, Eilat, J. Dafni) Liloa curta (A. Adams in Sowerby, 1850) Lamy 1938 and Barash & Zeniper 1993 both as Cylichna semisulcata SMARAGDINELLIDAE Phanerophthalmus collaris Eales, 1938 Eales 1938 (Gulf of Aden) Phanerophthalmus albocollaris Heller & Thompson, 1983 Heller & Thompson 1983 Phanerophthalmus olivaceus (Ehrenberg, 1828) Ehrenberg 1828 as Cryptophthalmus, Rüppell & Leuckart 1830 as Bulla smaragdina, Vayssière 1912 as Cryptophthalmus (Gulf of Aden), Sturany 1903, Heller & Thompson 1983 as P. smaragdinus, Yonow 2000 Smaragdinella calyculata (Broderip & Sowerby, 1829) Lamy 1938, Barash & Zeniper 1993 (rare as shells, Eilat, J. Dafni) Smaragdinella sieboldi A. Adams, 1864 Dekker & Orlin 2000 BULLIDAE Bulla ampulla Linnaeus, 1758

Issel 1869, Sturany 1903, Vayssière 1906 and as B. striata (Gulf of Aden), Eales 1938, Lamy 1938, Hughes 1977, Mastaller 1979, Sharabati 1984, Taylor & Reid 1984, Barash & Zeniper 1993, Yonow 2000

ORDER ANASPIDEA

AKERIDAE Akera soluta (Gmelin, 1791)

* * * (shells rarely washed up after winter storms, Eilat, J. Dafni)

APLYSIIDAE Aplysia cornigera Sowerby, 1869 Aplysia dactylomela Rang, 1828

Eales 1979, Mastaller 1979, Barash & Danin 1980 Rüppell & Leuckart 1828 as A. argus, Sturany 1903 as Tethys argus, Hughes 1977, Eales 1979, Mastaller 1979, Barash & Danin 1980

54 Checklist of species recorded from the Red Sea

Aplysia fasciata Poiret, 1789

Sturany 1903 as T. leporina, Eales 1960 as A. gracilis, Eales 1979, Barash & Danin 1980 (Mediterranean species) Aplysia oculifera Adams & Reeve, 1850 Engel & van Eeken 1962, Eales 1960, 1979, Fishelsen 1971, Mastaller 1979, Barash & Danin 1980, Barash & Zeniper 1994, Plaut et al. 1995, 1998 Aplysia parvula Guilding in Mörch, 1863 Eales 1960 (Gulf of Aden), Eales 1979, Mastaller 1979, Barash & Danin 1980 Aplysia sp. Sharabati 1984 as A. ?fasciata, Yonow 2000 Dolabella auricularia (Lightfoot, 1786) Ehrenberg 1831 and Tomlin 1927 both as D. hemprichii, Vayssière 1906 as D. rumphii, Engel & van Eeken 1962, Hughes 1977, Eales 1979, Barash & Danin 1980, Sharabati 1984; both Sturany 1903 and Mastaller 1979 as D. gigas (listings only) Dolabrifera dolabrifera (Cuvier, 1817) Sturany 1903 as D. cuvieri, Hughes 1977, Eales 1979, Mastaller 1979, Barash & Danin 1980 Petalifera petalifera Rang, 1828 Vayssière 1906 as Aplysiella gravieri and as A. g. var. alba, Engel & van Eeken 1962, Eales 1979, Mastaller 1979, Barash & Danin 1980 (Mediterranean species) Petalifera ramosa Baba, 1959 *** Phyllaplysia sp. Engel & van Eeken 1962 Syphonota geographica (Adams & Reeve, 1850) Engel & van Eeken 1962, Eales 1979, Barash & Danin 1980 NOTARCHIDAE Bursatella leachii savignana Audouin, 1826 laciniatus

Audouin 1826, Rüppell & Leuckart 1828 as Notarchus , Sturany 1903, Hägg 1904 as N. brevipes, Eales & Engel 1935, Barash & Danin 1972, Eales 1979, Barash & Danin 1980 Notarchus indicus Schweigger, 1820 Sturany 1903 as B. savignana, Engel & van Eeken 1962, Schuhmacher 1973b, Eales 1979, Mastaller 1979, Barash & Danin 1980, Yonow 2000 Stylocheilus longicauda (Quoy & Gaimard, 1824) *** Stylocheilus striatus (Quoy & Gaimard, 1832) Vayssière 1906 as Notarchus indicus (part), Bergh 1908 as Aclesia erythraea, Eales 1938, Sharabati 1984 ORDER PLEUROBRANCHIDA

PLEUROBRANCHIDAE Berthella stellata (Risso, 1826)

Heller & Thompson 1983 as B. stellata albocrossata [Mediterranean species, = B. pellucida (Pease)] Berthellina citrina (Rüppell & Leuckart, 1828) Rüppell & Leuckart 1828 as Pleurobranchus, Vayssière 1906 (and Gulf of Aden); Vayssière 1898 and Pruvot-Fol 1933 both as Berthella granulata, O’Donoghue 1929, Gohar & Aboul-Ela 1957a, Engel Checklist of species recorded from the Red Sea 55

& van Eeken 1962, Eales 1970, Fishelsen 1971, Marbach & Tsurnamal 1973, Hughes 1977, Mastaller 1979, Heller & Thompson 1983, Yonow 2000 Berthellina saidensis (O’Donoghue, 1929) O’Donoghue 1929 + Pleurobranchus albiguttatus (Bergh, 1905) Audouin 1826, O’Donoghue 1929, Barash & Danin 1972 (listing only) all as P. oblongus (syn. nov.) Pleurobranchus forskalii Rüppell & Leuckart, 1828 Forskål 1776 as Lepus marinus, Rüppell & Leuckart 1828, Issel 1869 as P. ruppelli, Vayssière 1898 as P. forskalii, P. perrieri, P. semperi; Vayssière 1906 (Gulf of Aden), Engel & van Eeken 1962; Hughes 1977, Mastaller 1979 (both only listings) Pleurobranchus grandis Pease, 1868 Yonow 2000 Pleurobranchus cf. grandis Pease, 1868 Bemert & Ormond 1971, Heller & Thompson 1983, Yonow 2000 all as P. forskalii Pleurobranchus mamillatus Quoy & Gaimard, 1832 *** Euselenops luniceps (Cuvier, 1817) Pruvot-Fol 1933 as Euselenops ?amboinei ORDER SACOGLOSSA

OXYNOIDAE Oxynoe viridis (Pease, 1861) JULIIDAE Julia exquisita Gould, 1862

*** Jousseaume 1888 and Lamy 1928 both as J. bourbonica, Mienis 1973 as Berthelinia sp.

PLAKOBRANCHIDAE (= ELYSIIDAE) Elysia tomentosa Jensen, 1997 Sharabati 1984 as Bursatella sp. Elysia trilobata Heller & Thompson, 1983 Heller & Thompson 1983 Elysia sp. 1 Eliot 1908 and Marcus 1980 both as Elysia grandifolia, Heller & Thompson 1983 as Elysia sp. Elysia sp. 2 Sharabati 1984 as Elysia sp. Elysiella pusilla Bergh, 1872 *** Thuridilla decorata (Heller & Thompson, 1983) Heller & Thompson 1983, Sharabati 1984, Yonow 1990 all as Elysia Thuridilla ? indopacifica Gosliner, 1995 * * * Plakobranchus ocellatus van Hasselt, 1824 Vayssière 1912 (Gulf of Aden), Hughes 1977, Heller & Thompson 1983, Yonow 1990, Barash & Zeniper 1994 as P. cholorphaeus POLYBRANCHIIDAE Cyerce elegans Bergh, 1870 Polybranchia orientalis (Kelaart, 1858)

***

STILIGERIDAE Stiliger ornatus Ehrenberg, 1828

Ehrenberg 1828 (see p. 37)

Sharabati 1984 as Cyerce sp.

56 Checklist of species recorded from the Red Sea

ORDER NUDIBRANCHIA DORIDINA

GONIODORIDIDAE Goniodoridella savignyi (Pruvot-Fol, 1933) Goniodoris joubini (Risbec, 1928) Goniodoris castanea (Alder & Hancock, 1845) Trapania toddi Rudman, 1987 GYMNODORIDIDAE Analogium striatum (Eliot, 1908) Analogium sp. Gymnodoris alba (Bergh, 1877) Gymnodoris ceylonica (Kelaart, 1858) Gymnodoris citrina (Bergh, 1875)

Pruvot-Fol 1933 as Goniodoris (photographic records from Eilat & Egypt)

***

Eliot 1908 (northern European species) Rudman 1987b (single 3 mm specimen) Eliot 1908 as Trevelyana, Vayssière 1912 (Gulf of Aden), O’Donoghue 1929, Sharabati 1981, Yonow 1990 all as Gymnodoris, Yonow 2000

*** ***

Yonow 1990 Aboul-Ela 1959 as Trevelyana bicolor, Yonow 1990 as Polycera sp. indet. Gymnodoris impudica (Rüppell & Leuckart, 1828) Rüppell & Leuckart 1828 as Doris, Haas 1920, White 1951 [= G. rubropapulosa (Bergh) (syn. nov.)] Gymnodoris inornata (Bergh, 1880) ?Abraham 1876 as Trevelyana concinna Gymnodoris cf. nigricolor Baba, 1960 Rudman & Darvell 1990 Gymnodoris sp. *** POLYCERIDAE Nembrotha megalocera Yonow, 1990 Tambja affinis (Eliot, 1904) Tambja limaciformis (Eliot, 1908) Thecacera pacifica (Bergh, 1883)

Yonow 1990, 2000, Cervera et al. 1996 Bemert & Ormond 1971, Sharabati 1984, Yonow 1990 Eliot 1908 as Nembrotha Eliot 1908 as Ohola

TRIOPHIDAE Plocamopherus indicus Bergh, 1890 Vayssière 1912 (Gulf of Aden) Plocamopherus ocellatus Rüppell & Leuckart, 1828 Rüppell & Leuckart 1828, Eliot 1908, Haas 1920, O’Donoghue 1929, White 1951 AEGIRIDAE Aegires citrinus Pruvot-Fol, 1930 Aegires villosus Farran, 1905

*** ***

HEXABRANCHIDAE Hexabranchus sanguineus (Rüppell & Leuckart, 1830) Rüppell & Leuckart 1830 as Doris, Ehrenberg 1828 and Vayssière 1912 both as H. praetextus, Abraham 1876 and Sturany 1903 both as H. suezensis, Hägg 1903 as H. plicatus, Eliot 1908, Eales 1938, Aboul-Ela 1959, Gohar & Soliman 1963, Schuhmacher 1973a, Hughes 1977, Mastaller 1979, Gat & Fainzilber 1984, Yonow 1990 Checklist of species recorded from the Red Sea 57

Hexabranchus sp.

***

ACTINOCYCLIDAE Actinocyclus sordida (Rüppell & Leuckart, 1830) Rüppell & Leuckart 1830 as Doris (assigned to Actinocyclus by O’Donohue 1929; not considered by Valdés 2002) + Actinocyclus verrucosus Ehrenberg, 1831 Ehrenberg 1831 Hallaxa indecora (Bergh, 1905) Vayssière 1912 (Gulf of Aden), O’Donoghue 1929 as H. decorata DISCODORIDIDAE Discodoris fragilis (Alder & Hancock, 1864) Vayssière 1912 (Gulf of Aden), Engel & van Eeken 1962, Gohar & Soliman 1967f all as D. concinna Discodoris granulata (Ehrenberg, 1831) Ehrenberg 1931 as Doris, O’Donoghue 1929 (not considered by Dayrat 2006) + Discodoris schmeltziana Bergh, 1880 *** Discodoris sp. Eliot 1908 as D. amboinensis (?) (not considered by Dayrat 2006) + Paradoris erythraeensis (Vayssière, 1912) Vayssière 1912 (Gulf of Aden), Aboul-Ela 1959, Gohar & Aboul-Ela 1959 all as Discodoris Sebadoris nubilosa (Pease, 1871) Crossland 1937 and White 1951 both as Thordisa cr osslandi, Soliman 1980 as S. crosslandi DORIDIDAE Aldisa sp. 1 Aldisa sp. 2 Archidoris vayssierea O’Donoghue, 1929 Archidoris sp. Atagema ornata (Ehrenberg, 1831) Atagema spongiosa (Kelaart, 1858) Doriopsis granulosa Pease, 1860 Doris sp. 1 Doris sp. 2

*** ***

Vayssière 1912 as A. staminea (Gulf of Aden), O’Donoghue 1929 + Pruvot-Fol, 1933 as A. o’donoghuei + Ehrenberg 1831, Gohar & Soliman 1967g as Trippa spongiosa Gohar & Soliman 1967g as Trippa areolata ?Audouin 1826 as Doris concentrica (see p. 35) Ehrenberg 1828 as Actinocyclus fragilis (removed by Valdés 2002) + Ehrenberg 1828 as Actinocyclus velutinus (removed by Valdés 2002) + Eliot 1908 as D. exanthemata +

Doris sp. 3 Hoplodoris grandiflora (Pease, 1860) *** Hoplodoris pustulata Abraham, 1877 Yonow et al. 2002 [= H. bifurcata (Baba) (syn. nov.)] Sclerodoris apiculata (Alder & Hancock, 1864) Vayssière 1912 (Gulf of Aden) and Eales 1938 both as Halgerda Sclerodoris tuberculata Eliot, 1904 *** Sclerodoris rugosa (Vayssière, 1912) Vayssière 1912 as Gravieria (Gulf of Aden) (see p. 49) Spongiodoris sp. Pruvot-Fol 1933 as S. rigida +

58 Checklist of species recorded from the Red Sea

HALGERDIDAE Artachaea clavata Eliot, 1908 Artachaea intermedia Eales, 1938 Artachaea verrucosa Eliot, 1908 Asteronotus cespitosus (van Hasselt, 1824)

Asteronotus denticulata (Eliot, 1908) Halgerda willeyi Eliot, 1904

KENTRODORIDIDAE Jorunna funebris (Kelaart, 1858)

Eliot 1908 Eales 1938 (Gulf of Aden) Eliot 1908 Ehrenberg 1831, Eliot 1908, Pruvot-Fol 1933, Crossland 1937, Eales 1938 all as A. hemprichii, Gohar & Soliman 1967e, Hughes 1977, Yonow 1990, 2000 (and Gulf of Aden) Eliot 1908 as Peronodoris Vayssière 1912 (Gulf of Aden), Eales 1938, Hughes 1977, Rudman 1978, Sharabati 1981, Yonow 1990, 2000 (and Gulf of Aden)

Eliot 1908 and O’Donoghue 1929 both as Kentrodoris labialis, Engel & van Eeken 1962 and Fishelsen 1971 both as Kentrodoris annnuligera, Mastaller 1979, Sharabati 1984

Jorunna pantherina Angas, 1864 Jorunna sp.

*** ***

PLATYDORIDIDAE Platydoris scabra (Cuvier, 1804) Yonow 1990 Platydoris striata (Kelaart. 1858) Platydoris sp.

Marcus & Marcus 1960, Hughes 1977, Soliman 1978,

***

Eliot 1908 as P. incertae +

CHROMODORIDIDAE Ardeadoris egretta Rudman, 1984 Cadlinella ornatissima (Risbec, 1928) Cadlinella sp. Chromodoris africana Eliot, 1904

*** *** ***

Eliot 1908, Gohar & Aboul-Ela 1959, Marcus & Marcus 1960 all as C. quadricolor, Sharabati 1981, Yonow 1989 Chromodoris annulata Eliot, 1904 Crossland 1937, Gohar & Aboul-Ela 1957c, AboulEla 1959, Sharabati 1984, Yonow 1989, Lütmann et al. 2006 Chromodoris aspersa (Gould, 1852) Eliot 1908, O’Donoghue 1929, Gohar & Soliman 1967b all as C. inornata, Rudman 1983, Yonow 1989 Chromodoris charlottae (Schrödl, 1999) Eliot 1911 as C. ?inopinata, Yonow 1989 and 2000 as Chromodoris sp., Schrödl 1999 as Glossodoris Chromodoris fidelis (Kelaart, 1858) Sharabati 1984, Yonow 1989 Chromodoris geminus Rudman, 1987 Bemert & Ormond 1971, Yonow 1989 Chromodoris obsoleta (Rüppell & Leuckart, 1830) Rüppell & Leuckart 1830 as Doris, Eliot 1911 as C. reticulata var., ?Pruvot-Fol 1933 as Glossodoris albomaculata erythraea, Mastaller 1979, Yonow 1989 Checklist of species recorded from the Red Sea 59

Chromodoris quadricolor (Rüppell & Leuckart, 1830) Rüppell & Leuckart 1830 as Doris, Sturany 1903, Eliot 1908, Pruvot-Fol 1933, White 1951, Aboul-Ela 1959, Marcus & Marcus 1960, Engel & van Eeken 1962, Fishelsen 1971, Hughes 1977, Mastaller 1979, Sharabati 1981, Bemert & Ormond 1971, Yonow 1989, 2000 Chromodoris strigata Rudman, 1987 *** Chromodoris tennentana (Kelaart, 1859) Mienis 1984 as C. vicina (single photograph) Chromodoris tinctoria (Rüppell & Leuckart, 1830) Rüppell & Leuckart 1830 as Doris, Eliot 1911, Haas 1920, Gohar & Soliman 1967c Chromodoris verrieri (Crosse, 1875) Yonow 1989 Chromodoris sp. Eliot 1908 as H. maculosa, Yonow 1989 and Perrone & Doneddu 2001 both as C. cf. decora Durvilledoris lemniscata (Quoy & Gaimard, 1832) Rudman 1984 Durvilledoris pusilla (Bergh, 1874) *** Glossodoris atromarginata (Cuvier, 1804) Vayssière 1912 (Gulf of Aden), White 1951, Marcus & Marcus 1960; Haas 1920 and Hughes 1977 both as listings only +++ Although Marcus & Marcus distinguish between G. cincta and this species, I have never seen specimens or photos of G. atromarginata from the Red Sea. Glossodoris cincta (Bergh, 1888) O’Donoghue 1929, White 1951, Engel & van Eeken 1962 all as Casella foxi, Gohar & Soliman 1967d as Casella obsoleta, Yonow 1989, 2000 Glossodoris electra Rudman, 1990 *** Glossodoris hikuerensis (Pruvot-Fol, 1954) Yonow 1989, 2000 Glossodoris pallida (Rüppell & Leuckart, 1830) Rüppell & Leuckart 1830 as Doris, Ehrenberg 1831 as D. xantholeuca, Eliot 1908, O’Donoghue 1929, Rudman 1984, Yonow 1989, 2000 Glossodoris plumbea (Pagenstecher, 1877) Pagenstecher 1877 as Doris, Sturany 1903, Gohar & Aboul-Ela 1959 as Casella atromarginata Glossodoris symmetricus Rudman, 1990 *** Glossodoris sp. Engel & van Eeken 1962 as G. rufomarginata + Hypselodoris dollfusi (Pruvot-Fol, 1933) Pruvot-Fol 1933 as Glossodoris, ?White 1951 as G. luteorosea, Gosliner & Behrens 2000 (Arabian Gulf) Hypselodoris infucata (Rüppell & Leuckart, 1830) Audouin 1826 as Doris marmorata (syn. nov.), Rüppell & Leuckart 1830 as Doris; Ehrenberg 1831, Issel 1869, Sturany 1903 all as Brachyclanis pantherina; ?Ehrenberg 1831 as Pterodoris picturata, Eliot 1908 as C. diardii; Vayssière 1912 (Gulf of Aden), O’Donoghue 1929, White 1951, Engel & van Eeken 1962 all as Chromodoris runcinata, Mastaller 1979, Yonow 1989, Johnson & Valdés 2001 Hypselodoris maculosa (Pease, 1871) Rudman 1986a, Yonow 1989 Hypselodoris maridadilus Rudman, 1977 Sharabati 1984, Yonow 1989 Hypselodoris nigrostriata (Eliot, 1904) Vayssière 1912 as C. tenui-linearis (Gulf of Aden) 60 Checklist of species recorded from the Red Sea

Mexichromis katalexis Yonow, 2001 Rudman 1983 as M. mariei (single specimen) Noumea alboannulata Rudman, 1986 Rudman 1986a (single specimen) Noumea flava (Eliot, 1904) *** Noumea norba Marcus & Marcus, 1970 Yonow 1989 as N. purpurea (single specimen) Noumea simplex (Pease, 1871) *** Noumea sudanica Rudman, 1985 Rudman 1985 (single specimen) Risbecia ghardaqana (Gohar & Aboul-Ela, 1957) Crossland 1937 as Chromodoris sp., Gohar & Aboul-Ela 1957c, Aboul-Ela 1959, Rudman 1987a Risbecia pulchella (Rüppell & Leuckart, 1830) Rüppell & Leuckart 1830 as Doris; Gohar & Aboul-Ela 1957c, Aboul-Ela 1959, Mastaller 1979 all as Chromodoris, Sharabati 1981, Yonow 1990 (1989), 2000 Thorunna africana Rudman, 1984 Eliot 1908 as T. furtiva, Rudman 1984 Ceratosoma tenue Abraham, 1876 Rochebrune 1894 as C. jousseaumei and C. rhopalicum, Haas 1920 as C. trilobatum, Pruvot-Fol 1933 and Crossland 1937 both as C. cornigerum, White 1951 as C. jousseaumei, Rudman 1988, Yonow 2000 (Gulf of Aden) Ceratosoma trilobatum (Gray, 1827) *** Miamira magnifica Eliot, 1910 *** Orodoris miamirana Bergh, 1875 Vayssière 1912 (Gulf of Aden) DENDRODORIDIDAE Dendrodoris coronata Kay & Young, 1969 * * * Dendrodoris cuprea (Ehrenberg, 1831) Ehrenberg 1831 as Doris, Engel & van Eeken 1962, Mastaller 1979 (listing only) Dendrodoris elongata Baba 1936 ?Eliot 1908 as Doridopsis sp. Dendrodoris fumata (Rüppell & Leuckart, 1830) Rüppell & Leuckart 1830 as Doris; Eliot 1908, White 1951, Hughes 1977, Mastaller 1979 all as D. rubra, Vayssière 1912 as Doriopsis rosea (Gulf of Aden), Gohar & Soliman 1967a, Haas 1920 Dendrodoris immaculata (Audouin, 1826) Audouin 1826 as Doris (see p. 35) Dendrodoris jousseaumei (Vayssière, 1912) Vayssière 1912 as Doriopsis (Gulf of Aden) (see p. 49), O’Donoghue 1929 Dendrodoris leptopus Ehrenberg, 1831 Ehrenberg 1831 Dendrodoris lugubris Ehrenberg, 1831 Ehrenberg 1831 (type species) Dendrodoris nigra (Stimpson, 1855) Rüppell & Leuckart 1830 as Doris albo-limbata (syn. nov.) (see p. 38); Eliot 1908, Vayssière 1912 (Gulf of Aden) both as Doriopsis, O’Donoghue 1929, Engel & van Eeken 1962, Hughes 1977, Mastaller 1979, Yonow 1990 Dendrodoris nigropunctata (Vayssière, 1912) Vayssière 1912 as Doriopsis, Haas 1920, O’Donoghue 1929 Dendrodoris tigrina (Audouin, 1826) Audouin 1826 as Doris (see p. 35), Rüppell & Leuckart 1830 as Doris punctata (syn. nov.) (see p. 43), Vayssière 1912 (Gulf of Aden) and Issel 1869 both as D. punctata Checklist of species recorded from the Red Sea 61

Dendrodoris tuberculosa (Quoy & Gaimard, 1832) Engel & van Eeken 1962 as D. pustulosa Dendrodoris sp. *** Doriopsilla sp. White 1951 + PHYLLIDIIDAE Fryeria rueppelii (Bergh, 1869)

Rüppell & Leuckart 1828 and Eales 1938 both as P. pustulosa, Vayssière 1912 (Gulf of Aden), O’Donoghue 1929, White 1951 as P. varicosa, Yonow 1986, 1988, 1990 (and Gulf of Aden) Phyllidia multifaria Yonow, 1986 Heller & Thompson 1983 as P. varicosa, Yonow 1986, 1988, 1990 (and Gulf of Aden) Phyllidia schupporum Fahrner & Schrödl, 2000 Fahrner & Schrödl 2000a Phyllidia undula Yonow, 1986 Sharabati 1981, Yonow 1986, 1988 Phyllidia varicosa Lamarck, 1801 Ehrenberg 1831 as P. arabica, Vayssière 1912 (Gulf of Aden), Sharabati 1981, Soliman 1986; Sharabati 1984, Yonow 1988, 2000 (also Gulf of Aden) all as P. arabica, Fahrner & Schrödl 2000b Phyllidia sp. Eliot 1908 as P. varicosa Phyllidiella pustulosa (Cuvier, 1804) Heller & Thompson 1983; Yonow 1986, 1988, 2000 (Gulf of Aden) also as P. melanocera (= juveniles) Phyllidiopsis cardinalis Bergh, 1873 *** Phyllidiopsis dautzenbergi (Vayssière, 1912) Vayssière 1912 as Phyllidia (Gulf of Aden), O’Donoghue 1929, Heller & Thompson 1983, Sharabati 1984, Yonow 1986, 1988 Phyllidiopsis monacha (Yonow, 1986) Yonow 1986 as Phyllidia (single specimen) Phyllidiopsis sinaiensis (Yonow, 1988) Heller & Thompson 1983 only fig. 8D as P. varicosa, Yonow 1986, 1988, 1990 also as P. quadrilineata, Fahrner & Schrödl 2000c Phyllidiopsis sudanensis (Heller & Thompson, 1983) Heller & Thompson 1983 as Phyllidia (single specimen) DENDRONOTINA

TRITONIIDAE Marionia glama (Rüppell & Leuckart, 1828) Rüppell & Leuckart 1828 as Tritonia, Yonow 2000 Marioniopsis cyanobranchiata (Rüppell & Leuckart, 1828) Rüppell & Leuckart 1828 as Tritonia, Sturany 1903, Eliot 1908, Sharabati 1984, Yonow 2000 Marioniopsis levis (Eliot, 1904) Avila et al. 1999 as M. fulvicola (syn. nov.), Yonow 2000 Marioniopsis rubra (Rüppell & Leuckart, 1828) Rüppell & Leuckart 1828 as Tritonia, Sharabati 1981, Yonow 2000 Marioniopsis viridescens (Eliot, 1904) Yonow 2000 Tritoniopsis elegans (Audouin, 1826) Audouin 1826 as Tritonia, Pruvot-Fol 1933, White 1951 as Tritoniopsis sp., Engel & van Eeken 1962 Tritoniopsis gravieri Vayssière, 1912 Vayssière 1912 (see p. 47), White 1951 (both Gulf of Aden)

62 Checklist of species recorded from the Red Sea

BORNELLIDAE Bornella stellifer (Adams & Reeve in Adams, 1848) Collingwood 1881 as B. marmorata, Vayssière 1912 as B. digitata (both Gulf of Aden), Yonow 2000 Bornellopsis kabretiana O’Donoghue, 1929 O’Donoghue 1929 DOTOIDAE Doto orcha Yonow, 2000

Yonow 2000 (single specimen)

LOMANOTIDAE Lomanotus vermiformis Eliot, 1908

Eliot 1908, Pruvot-Fol 1933

SCYLLAEIDAE Scyllaea pelagica Linnaeus, 1758 Crosslandia viridis Eliot, 1902

Forskål 1775 as S. ghomfodensis, Yonow 2000 Eliot 1908 as C. fusca

TETHYIDAE Melibe bucephala Bergh, 1902 Melibe rangii Bergh, 1875

O’Donoghue 1929, single photo H. Debelius Bergh 1875, Haas 1920; Eliot 1908, Engel & van Eeken 1962, Barash & Danin 1972, Schuhmacher 1973b, Mastaller 1979, Yonow 2000 all as M. bucephala

ARMININA

ARMINIDAE Armina semperi (Bergh, 1860)

Pruvot-Fol 1933 as A. semperi erythraea, Eales 1938 (Gulf of Aden, deep sea)

Dermatobranchus albus (Eliot, 1904) *** Dermatobranchus glabrus (Eliot, 1908) Eliot 1908 Dermatobranchus gonatophorus van Hasselt, 1824 *** Dermatobranchus ornatus (Bergh, 1874) * * * Dermatobranchus striatus van Hasselt, 1824 Eales 1938, Gohar 1948, Aboul-Ela 1959 Dermatobranchus sp. *** MADRELLIDAE Madrella ferruginosa Alder & Hancock, 1864

***

AEOLIDINA

FLABELLINIDAE Flabellina bicolor (Kelaart, 1858) Sharabati 1984, Yonow 2000 “Flabellina” sp. *** Flabellina bilas Gosliner & Willan, 1991 * * * Flabellina rubrolineata (O’Donoghue, 1929) O’Donoghue 1929 as Coryphellina, Mienis & Gat 1986, Yonow 2000

Checklist of species recorded from the Red Sea 63

FACELINIDAE Facelina rhodopos Yonow, 2000 “Facelina” sp.

Yonow 2000

***

AEOLIDIIDAE Aeolidia habessinica Ehrenberg, 1828 Ehrenberg 1828 + Aeolidiella alba Risbec, 1928 *** Aeolidiella indica Bergh, 1888 Eliot 1908 as A. orientalis + Baeolidia fusiformis Baba, 1949 Yonow 2000 Baeolidia moebii Bergh, 1888 O’Donoghue 1929, Engel & van Eeken 1962 Cerberilla annulata (Quoy & Gaimard, 1832) * * * (single photo B. & S. Koretz) Limenandra nodosa Haefelfinger & Stamm, 1959) * * * (single photo B. & S. Koretz) (Mediterranean species) Limenandra sp. Yonow 2000 as Limenandra nodosa Spurilla australis Rudman, 1982 Yonow 2000 (photographs only) Spurilla major (Eliot, 1903) Yonow 2000 (single specimen) EUBRANCHIDAE Dunga ocellata (Alder & Hancock, 1864) Eubranchus rubropunctatus Edmunds, 1969

*** ***

GLAUCIDAE Caloria indica (Bergh, 1896) *** Favorinus japonicus Baba, 1949 *** Favorinus tsuruganus Baba & Abe, 1964 * * * Glaucus atlanticus Forster, 1777 Thompson & McFarlane 1967 (Gulf of Aden) Glaucilla marginata Bergh, 1868 *** Phyllodesmium colemani Rudman, 1991 * * * Phyllodesmium crypticum Rudman, 1981 * * * Phyllodesmium fastuosum Ehrenberg, 1831 Ehrenberg 1831 Phyllodesmium hyalinum Ehrenberg, 1831 Ehrenberg 1831, White 1951, Gohar & Aboul-Ela 1957b as P. xeniae, Engel & van Eeken 1962 as Favorinus horridus brevitentaculus, Yonow 2000 Phyllodesmium magnum Rudman, 1991 ?Rüppell & Leuckart 1828 as Eolida bella (see p. 46), Yonow 2000 Pteraeolidia ianthina (Angas, 1864) Marcus & Marcus 1960 as P. semperi, Hughes 1977, Sharabati 1981, Yonow 2000 EMBLETONIIDAE Embletonia gracile Risbec, 1928 TERGIPEDIDAE Catriona susa Marcus & Marcus, 1960 Cuthona sibogae (Bergh, 1905) Phestilla lugubris (Bergh, 1870) Phestilla melanobrachia Bergh, 1874

Yonow 2000

Marcus & Marcus 1960

***

Yonow 2000 Sharabati 1984 as Cuthona sp., Yonow 2000

64 Checklist of species recorded from the Red Sea

Incertae sedis – Actinodoris sponsa Ehrenberg 1831: Rudman (1984) stated it was impossible to know the identity of Actinodoris sponsa but that it belonged to the genus Chromodoris, similar to C. quadricolor and C. africana. – Pterodoris brachyphylla Ehrenberg 1831: although the species was not discussed in revisions (Rudman 1984, Johnson & Valdés 2001), the genus Pterodoris was suppressed in preference for Hyspelodoris. – Deridobranchus argus Ehrenberg 1831: I have not yet found any mention of this species in the literature. – Doris erythraea Ehrenberg 1831: O’Donoghue (1929 p. 724) lists this species as Glossodoris, citing Chromodoris erythraea Abraham (1877) and Chromodoris ? erythraea Bergh (1877), which are non-existent entities. Pruvot-Fol (1933) simply includes Ehrenberg’s species in her list. Notes species recorded by J. Dafni can be found on www.dafni.com/mollusca/Families3.htm species recorded by B. & S. Koretz can be found on www.koretz.net/Eilat/Invertebrates/ Molluscs/Opistobranchs.htm (sic)

Checklist of species recorded from the Red Sea 65

KEY TO SPECIES OF RED SEA OPISTHOBRANCHS

The following is a key to the opisthobranch molluscs illustrated in this book: it will not key out previously recorded or other species not illustrated in the “Species Accounts”. The majority key out as species, although some genera key out if the representative species are few or similar in form but easily distinguished by colour pattern. Some species key out because they are not typical of their families or genera, and their conspecifics may key out elsewhere. A few species may key out twice depending upon the condition of the animal and interpretation of the characters. The terminology used has been simplified as much as possible, and the terms utilized have been defined or illustrated in the introduction and glossary. 1 –

Animal with an obvious shell, even if internal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Animal without an obvious shell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

2.

Shell white, oval or globular, but animal within is pigmented with patches of drab browns and greens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Globular shell, animal small (< 15 mm) and brightly coloured . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Larger bubble shell (20–60 mm) with coloured patches or lines . . . . . . . . . . . . . . . . . . . . . . . . . 5

– – 3. – – –

Animal translucent white with opaque white spots, faint ochre band between rather small eyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Liloa curta Animal with opaque white spots, black speckles, and green patches; dense brown band between prominent large eyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diniatys dentifer Body with glistening white pigment, brown patches in shell region . . . . . . . . . . . Atys semistriatus Body patchy brown and pale green, white patch between faint eyes . . . . . . . . . . . Diniatys dubius

4. – – – – –

Shell and animal white but with purple margins . . . . . . . . . . . . . . . . . . . . Haminoea cyanomarginata Animal green with long tail and small shell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oxynoe viridis Animal pink with orange margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Sagaminopteron cf. ornatum Animal yellow and white with brown margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . Siphopteron sp. Animal black with white patches containing yellow spots . . . . . . . . . . . . . . . Colpodaspis thompsoni Animal cream or grey with orange or brown patches surrounded by black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sagaminopteron cf. psychedelicum

5. – –

Shell with patches in varying shades of brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bulla ampulla Shell with three white bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydatina zonata Shell with spiral brown lines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

6. –

Animal shades of pink with bright blue margins, many brown lines . . . . . . . . . . .Hydatina physis Animal translucent white, few brown lines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Atys naucum

7. – –

Animal with parapodial lobes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Animal without parapodial lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Parapodial lobes difficult to detect, colourful tuberculate body . . . . . . . . . . . . . . Petalifera ramosa 66 Key to species of Red Sea opisthobranchs

8. –

Parapodia held close to body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Parapodia larger, held open above the body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

9. –

Obvious rolled rhinophores anteriorly, single metapodium . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 No rhinophores, body in four parts, sometimes two tails . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

10. Pair of rhinophores only (no obvious oral tentacles) (Plakobranchidae) . . . . . . . . . . . . . . . . . 11 – Rhinophores and oral tentacles clearly visible, both rolled (Aplysiidae) . . . . . . . . . . . . . . . . . . 12 11. Very long and thin, colourful, usually small < 15 mm . . . . . . . . . . . . . . . . . . . . . . . Thuridilla spp. – Flattened with variously coloured eye spots,< 35 mm . . . . . . . . . . . . . . . . . Plakobranchus ocellatus – Grass green with mottled beige rhinophores, approx. 10 mm . . . . . . . . . . . . . . . . Elysiella pusilla 12. Very large, up to 300 mm, truncated posterior . . . . . . . . . . . . . . . . . . . . . . . . . Dolabella auricularia – Small, < 50 mm, rounded posterior and flattened body . . . . . . . . . . . . . . . . Dolabrifera dolabrifera – Very small, 10 mm, elongated dark green body . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllaplysia sp. 13. Black with bright colour patterns, tail in two parts, the left usually longer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chelidonura spp. + Philinopsis spp. – Pale body, small and cryptic, eyes visible. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14. – – – –

White or cream with red, brown, green pattern . . . . . . . . . . . . . . . . . . . . . . . Odontoglaja guamensis Pale green or olive green with notch on head shield . . . . . . . . . . . . . . . Phanerophthalmus olivaceus Pale lilac with white marking on head shield . . . . . . . . . . . . . . . . . . . .Phanerophthalmus albocollaris Dense green, on algae (with rhinophores) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elysiella pusilla Dark green, rolled rhinophores and tentacles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllaplysia sp.

15. Elongated body, parapodia along complete length of body to tail, one pair of rolled tentacles (Plakobranchidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 – Body with distinct neck region, swollen posteriorly where parapodia are located, two obvious pairs of rolled tentacles (Aplysiidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 16. – – –

Pale green and ‘hairy’ in appearance, pink margins . . . . . . . . . . . . . . . . . . . . . . . . Elysia tomentosa Pale green with white patches and bright orange submargin . . . . . . . . . . . . . . . . . . . . Elysia sp. 1 Dark green with black patches, streaks, and margins . . . . . . . . . . . . . . . . . . . . . . . . . . Elysia sp. 2 Knobbly appearance, basically pale but colourful. . . . . . . . . . . . . . . . . . . . . . . . . . . Elysia trilobata

17. – – –

“Typical” sea hares with large upstanding parapodia . . . . . . . . . . . . . . . . . . . . . . . . . .Aplysia spp. Translucent globular body with short pointed papillae . . . . . . . . . . . . . . . . . . . . Notarchus indicus Bright yellow body with large pink spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stylocheilus longicauda Long and thin body with stripes and blue violet eye spots . . . . . . . . . . . . . . . . Stylocheilus striatus

18. Dorsum with elongate, branching, swollen or leaf-like projections (cerata) . . . . . . . . . . . . . . . 19 – Body elongated or oval. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 – Body not as above: long and thin, drab colours, with tiny transparent finger-like projections along mantle edge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lomanotus vermiformis

Key to species of Red Sea opisthobranchs 67

19. Single row of cerata not leaf-like or angular, rhinophore sheath may be present . . . . . . . . . . 20 – Multiple or single cerata leaf-like or angular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 – Multiple cerata in clusters, elongated body, medium to long oral tentacles, long rhinophores without sheaths (Aeolidina) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56 – Many cerata in fringe all around body, red in colour . . . . . . . . . . . . . . . . . . . . Madrella ferruginosa 20. Paired cerata inflated and tuberculate, animal translucent with no rhinophore sheath (Eubranchidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dunga ocellata + Eubranchus rubropunctatus – Paired cerata swollen and cup-shaped, animal white . . . . . . . . . . . . . . . . . . . . . . Embletonia gracile – Paired cerata swollen and tuberculate, animal brownish green . . . . . . . . . . . . . . . . . . . Doto orcha – Paired cerata branching vertically or horizontally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 21. Flattened cerata (easily autotomized) are distributed all over the body. . . . . . . . . . . . . . . . . . . 22 – Single cerata in a paired series along the sides of the body, sometimes branching; oral veil present anteriorly, complicated rhinophore sheaths (Dendronotina) . . . . . . . . . . . . . . . . . . . . 23 22. Cerata oval and swollen, translucent, iridescent blue spot at base, rolled rhinophores anteriorly (Polybranchiidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyerce elegans – Cerata irregular with white serrated margin, rolled rhinophores anteriorly (Polybranchiidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polybranchia orientalis – Cerata irregular with pustules, curled at tips, very long smooth rhinophores (Aeolidina) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllodesmium crypticum 23. Digitate oral veil and rhinophore sheaths, cream in colour with orange marbling and rings, leaf-like cerata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Bornella stellifer – One or two pairs of spatulate cerata, rhinophores enclosed in flared sheaths, greenish or brown body colour . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Scyllaea pelagica – Rhinophore sheaths with large flap, cylindrical cerata with papillae and tubercles, body translucent to opaque brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melibe rangii 24. Chunky body, single row of vertical branching gills on each side of dorsum, complex rhinophores and sheaths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Marionia, Marioniopsis, Tritoniopsis – Translucent elongated body, body and cerata with long papillae, rhinophore sheath with large flap and distal papillae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melibe rangii – Flattened body, length barely longer than breadth, four clusters of finger-like cerata on each side, blue; no rhinophore sheath . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glaucilla marginata – Flattened body with many branching cerata, white or orange, may have red marbling, branching rhinophore sheaths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Tritoniopsis elegans 25. Body without median or posterior dorsal gills . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 – Gills present as median or posterior arch, clump, or circlet on dorsum . . . . . . . . . . . . . . . . . . 32 26. Soft oval body, trapezoidal oral veil, short rolled rhinophores projecting forwards, single gill under mantle on right side (Pleurobranchida). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 – Body stiff, lamellate rhinophores, gills may be present between foot and mantle on both sides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

68 Key to species of Red Sea opisthobranchs

27. Large opisthobranchs, > 100 mm, red or red and white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 – Smaller, < 30 mm, orange, or brown and white, or pale with red/purple and white rings . . . 29 28. Dorsum with evenly sized, randomly distributed tubercles; often red or white with contrasting semi-lunar markings, wavy mantle margin, 100 mm. . . . . . . . . . . . . . . . . . . Pleurobranchus forskalii – Dorsum with red and white polygons containing tubercles surrounding a larger central one, posterior siphon present, 200 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Pleurobranchus grandis – Dorsum with regular polygons containing a central tubercle, pedal gland on sole of foot, no posterior siphon, 100 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pleurobranchus cf. grandis 29. Uniformly orange or yellow, fairly smooth dorsum . . . . . . . . . . . . . . . . . . . . . . . Berthellina citrina – Very pale with darker pigment and tubercles . . . . . . . . . . . . . . . . . . . . . Pleurobranchus albiguttatus – Pale with groups of tubercles, the central one the largest; white and violet rings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pleurobranchus mamillatus 30. Body broad, rounded anteriorly and tapered posteriorly; discontinuity in mantle where rhinophores emerge; rhinophores with longitudinal lamellae (Arminidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Dermatobranchus spp. – Body elongated or oval, usually tuberculate, black with orange/yellow pigment or black and pink; rhinophores with transverse lamellae, not emerging from anterior margin (Phyllidiidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 31. Body multi-coloured brown, yellow, red, green, etc; complex tubercles, bright yellow foot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllidiopsis cardinalis – Body black, white, and orange; orange rhinophores . . . . . . Phyllidia, Fryeria, Phyllidiopsis monacha – Body pink or white, and black; rhinophores pale with black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllidiella, Phyllidiopsis, Aldisa sp. 2 32. Gills present medially, mantle reduced or margin absent, body tapering posteriorly . . . . . . . . 33 – Gills present as posterior circlet, mantle skirt present but may be rolled up . . . . . . . . . . . . . . 40 33. Digitiform process present on each side of gills, rhinophore sheaths with flap-like extensions, bright orange with blue and black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Thecacera pacifica – Front edge branching, keel on tail, orange spots . . . . . . . . . . . . . . . . . . . . . . . . Plocamopherus spp. – Body stiff and rather rubbery . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 – Body soft and flexible. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 34. Body with spatulate head and mantle drawn out in horn behind gills . . . . . . . . . Ceratosoma spp. – Body covered in mushroom or club-shaped tubercles . . . . . . . . . . . . . . . . . . . . . . . . . Aegires spp. 35. Large (> 70 mm), black ground colour with large black rhinophores . . . . . . . . . . . . . . . . . . . 36 – Smaller, other colours . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 36. Body with orange stripes and some green markings . . . . . . . . . . . . . . . . . . . . . . . . . Tambja affinis – Body with yellow, purple, and red patches . . . . . . . . . . . . . . . . . . . . . . . . . . . Nembrotha megalocera

Key to species of Red Sea opisthobranchs 69

37. Translucent creamy body with orange markings on raised pustules or ridges (Gymnodorididae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 – Body otherwise coloured . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 38. Gills present as an arch, orange ridges on dorsum . . . . . . . . . . . . . . . . . . . . . . .Analogium striatum – Gills present as circlet, raised orange or yellow papillae or pustules . . . . . . . . . . .Gymnodoris spp. 39. – – –

Brick red colour with cream markings, some violet . . . . . . . . . . . . . . . . . . . . . .Tambja limaciformis Purple body with red ridges and white patches . . . . . . . . . . . . . . . . . . . . . . . . . . . . Analogium sp. Uniformly orange body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnodoris inornata Uniformly white body covered in round tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnodoris sp.

40. Body soft and fleshy; smooth, papillate, or tuberculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 – Body firm; smooth, rough, ridged or tuberculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 41. Gills located far posteriorly, rhinophores far anteriorly; soft and relatively smooth or with complex compound tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Dendrodoris spp. – Gills and rhinophores more centrally located, usually with brightly coloured patterns . . . . . . 42 42. Body with elongated papillae or tubercles. . . . . . . . . . . . . . Mexichromis katalexis + Cadlinella spp. – Body without papillae, relatively smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 43. Dorsum with rolled mantle skirt . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 – Dorsum without rolled edges, usually oval, may have permanent undulations (Chromodorididae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 44. Large, > 150 mm, usually red or pink, with six gills . . . . . . . . . . . . . . . . . . . . .Hexabranchus spp. – Tiny, < 10 mm, blue black with yellow spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . Goniodoris joubini 45. Dorsum simple, elongated or oval . . . . . . Chromodoris, Hypselodoris, Durvilledoris, Noumea, Risbecia – Mantle margin with permanent undulations along the sides . . . . Glossodoris, Thorunna, Ardeadoris 46. Body with large process over-hanging gills from behind . . . . . . . . . . . . . . . . . . . . Ceratosoma spp. – Body with very pronounced dorsal hump, with or without tubercles and papillae . . . . . . . . . 47 – Body with slight hump or flat; smooth, tuberculate, sculptured, papillate, etc. . . . . . . . . . . . . 48 47. Tiny, 15 mm, dark purple to brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hallaxa indecora – Red with sandy coloured spiky crest, 55 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atagema ornata – Drab colours with long filaments extending from tubercles around the margin, 40 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Sclerodoris apiculata – Mantle with one very large complex tubercle in front of the gills, large eye spots or pits around the dorsal hump, 140 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Atagema spongiosa 48. Body firm and rubbery, smooth but sculptured . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 – Dorsum with tubercles or papillae, may be rough to touch. . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 – Dorsum tuberculate with one or more central pit, red in colour . . . . . . . . . . . . . . . . . . . . . . . 55

70 Key to species of Red Sea opisthobranchs

49. Dorsum with peaks connected by ridges, white with black and yellow markings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Halgerda willeyi – Mantle with central ridge and four lobes; ventral surface with orange spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Miamira magnifica – Mantle with many lobes, ventral surface with green lines and blue eye spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Orodoris miamirana – Mantle with large tubercles, arranged in rings around the edge of the mantle, brown and white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asteronotus cespitosus – Rubbery, olive green to brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Actinocyclus verrucosus 50. – – –

Mantle colours bright . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mantle colours drab and cryptic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Body very flattened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Body with slight dorsal hump . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

51 52 53 54

51. Mantle white with blue purple tipped tubercles . . . . . . . . . . . . . . . . . . . . . . .Mexichromis katalexis – Mantle yellow or white with yellow lines, papillae long and pointed or club-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cadlinella spp. – Mantle with large tubercles, beige and browns with purple, yellow, etc. patches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Discodoris schmeltziana 52. – – –

Mantle with small evenly sized tubercles, shades of brown. . . . . . . . . . . . . . . . . . Hoplodoris spp. Mantle unevenly tuberculate, many tubercles with filament . . . . . . . . . . . . . . .Sclerodoris apiculata Mantle with soft papillae, especially long around margins . . . . . . . . . . . . . . . . . . Sebadoris nubilosa Mantle with low tubercles, pink and black resembling Phyllidiidae . . . . . . . . . . . . . . . Aldisa sp. 2

53. Body yellowish brown with orange margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Platydoris scabra – Body white covered in brown wavy lines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Platydoris striata – Mantle variously coloured in greys and browns but with darker symmetrical patches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Discodoris fragilis – Mantle very rounded, pale yellow with few black spots . . . . . . . . . . . . . . . . . Paradoris erythraeensis 54. Body white with black patches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Jorunna funebris – Body white or yellow with single black ‘spikes’ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Jorunna sp. – Body beige with pale brown patches and white spicules . . . . . . . . . . . . . . . . . . Jorunna pantherina 55. Mantle with ‘splashes’ of coral sand (ventral surfaces red); dorsum very unevenly tuberculate but always with a central pit, often with black ‘holes’ along the sides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sclerodoris tuberculata – Mantle evenly tuberculate with two pits on the dorsum and smaller raised pits on the mantle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aldisa sp. 1 56. – – –

Insipid white or pale colouration, sculptured rhinophores (usually) . . . . . . . . . . . . . . . . . . . . . Beige or brown but not white and not colourful . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cream to brown with lemon yellow rings along the body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bright, colourful, or patterned . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Key to species of Red Sea opisthobranchs 71

57 58 59 60

57. Tiny, 10 mm, flattened body, plump cerata, unusual swollen rhinophores with two constrictions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aeolidiella alba – Large, < 30 mm, with very long metapodium, finger-like cerata with recurved ends, rhinophores with two or three swellings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Favorinus japonicus – Very large, 50 mm, long knobbly cerata capped with white . . . . . . . . . . . . . . . . . Phestilla lugubris 58. – – –

Beige brown, elongated cerata pustulate on recurved ends only . . . . . . . . .Phyllodesmium hyalinum Beige, many crowded irregular flattened pustulate cerata . . . . . . . . . . . . . Phyllodesmium crypticum Brownish green, few long swollen cerata, opaque white patches . . . . . . . . Phyllodesmium colemani Dark brown speckling, flattened cerata translucent with white tips . . . . . . . . . . . . . Spurilla major

59. Dark brown speckling, short curled cerata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Limenandra sp. – Cream body, long cerata with one very long one in each group . . . . . . . . . . . . Baeolidia fusiformis 60. – – – – –

Uniformly coloured, not banded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61 Cerata with single ring or band of orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62 Cerata with two rings of different colours . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 Cerata with white or cream bands and red and white rings on cerata . . . . . . . . . . Flabellina bilas Cerata and tentacles banded in brown, green, violet, or blue; very long, 70 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pteraeolidia ianthina Cerata with coloured cap at tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64

61. Body orange, brown, or black; cerata with orange or white tips . . . . . . . . . Phestilla melanobrachia – Body blue and silver, flattened, with 3 or 4 horizontal groups of cerata . . . . . Glaucilla marginata 62. Rhinophores short and plump, lamellated and orange . . . . . . . . . . . . . . . . . . . . . . Flabellina bicolor – Rhinophores long and smooth, translucent and opaque white . . . . . . . . . . . . . . . “Flabellina” sp. – Rhinophores long and lamellated, translucent orange; swollen cerata with terminal nipple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Facelina rhodopos 63. Pink and lilac (or brighter orange and purple) bands around swollen cerata, body pink and appearing iridescent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . “Facelina” sp. – Blue and orange bands on swollen cerata, blue spot on head, body beige or pale orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spurilla australis 64. Cerata with at least one blue band and white tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65 – Cerata with yellow or orange tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 65. Orange cerata with blue band, body white, sculptured rhinophores . . . . . . . . Favorinus tsuruganus – Cerata with orange, blue, and yellow bands; central white line, head with orange patches near eyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caloria indica 66. Cerata long and thin, white with magenta bands, body with three stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Flabellina rubrolineata – Cerata long and thin, pink deepening to magenta, body also pale pink . . . . . . . . .Cuthona sibogae – Cerata blue or lilac, sometimes brownish, with a cream band . . . . . . . . . . . Phyllodesmium magnum

72 Key to species of Red Sea opisthobranchs

Species Accounts

Order Cephalaspidea

Family Hydatinidae

Order Cephalaspidea The Cephalaspidea are the most primitive order of opisthobranchs, containing burrowing forms which have a solid shell and an operculum. The more advanced forms still burrow, but have a reduced external or a small internal shell and no operculum. Most species are carnivorous; herbivory is a more evolved adaptation. Family Hydatinidae The Hydatinidae are a small family comprising few genera of rather large and brightly coloured species, especially in the Red Sea. The shell is large and globular, usually striated, and the animal is often brightly coloured and too large to retract into the shell. There is no operculum with which to close the shell. Hydatina is known to feed on sedentary cirratulid polychaete worms occurring in shallow tidal and intertidal waters.

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Hydatina physis (Linnaeus, 1758) Size: 65 mm shell (animal up to 90 mm) Diagnostic characters: The shell is cream or beige with thin dark brown lines running around the shell. There may be paler brown patches especially along the growth lines. The animal is large and fleshy in shades of orange, peach or pink, sometimes quite dark, with pale blue or green margins.

Hydatina physis

B&SK

74 Species accounts

Order Cephalaspidea

Family Hydatinidae

Biology and ecology: Hydatina physis can be found burrowing in soft sediments. The egg mass is a white convoluted ribbon crushed all together, laid on the mantle before being attached to the substrate. Occurrence: Dead shells are infrequently washed up in the Gulf of Eilat; numerous photographic records from the northern and central Red Sea. Geographical distribution: Widespread throughout the tropical Indo-West Pacific and the Atlantic Oceans; records from the latter are often cited as H. stromfelti Odhner and H. vesicaria Solander.

Hydatina zonata (Lightfoot, 1786) Size: 20 mm shell Diagnostic characters:

The shell is identical in shape to that of H. physis but smaller, globular and fairly well calcified. There are faint brown longitudinal lines with three white bands across them edged in thick brown lines: this is the colour variety ‘velum’ occurring in the Red Sea. The animal in the photograph is translucent with a luminescent pale blue band around the margins. The black and white photograph of the preserved animal in Eales (1938) is dark with white margins, implying variation in body colour as in H. physis. Her specimen had an 80 mm long cirratulid polychaete in its crop. Occurrence: Single photographic record from the Gulf of Eilat (1980’s). Geographical distribution: Tropical Indo-West Pacific.

Hydatina zonata

A. Diamant

Species accounts 75

Order Cephalaspidea

Family Diaphanidae / Family Aglajidae

Family Diaphanidae Members of the family Diaphanidae are generally small, with a fragile globular shell into which the animal’s body can retract completely. Numerous genera make up the family and are found from Iceland in the north, south through temperate and tropical zones to the Antarctic. These animals are rarely found and not much is known of their biology and ecology. The name comes from the Greek meaning transparent.

Colpodaspis thompsoni OL

Colpodaspis thompsoni Brown, 1979 Size: less than 5 mm Diagnostic characteristics: The body is dark brown to black with white patches containing one or more yellow pustules. There is an exhalent siphon on the right side and a long black tail. The outer surfaces of the tentacles on the head are paler than the rest of the body. The shell is completely enclosed. Biology and ecology: Lives epifaunally in brown algae communities and coral rubble. Occurrence: Two photographic records from the Gulf of Eilat. Geographical distribution: Tropical Indo-West Pacific. Family Aglajidae Aglajids are active predators of macro-invertebrates but none of the genera except Odontaglaja possesses a radula. The body comprises two body shields and a pair of parapodial lobes. The posterior processes are variously developed. A small internal shell is located in the posterior shield. Mating occurs in pairs or chains, with the animal in front acting as the female and the one behind as the male. Eggs are laid in a white string forming a cocoon around the slug which then slips out, leaving a hollow sausage-shaped egg mass attached to the substrate. The name Chelidonura refers to the tails of a swallow in Greek. Chelidonura flavolobata Heller and Thompson, 1983 Size: 70 mm Diagnostic characteristics:

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Chelidonura spp.

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The body colouration is deep blue black with creamy yellow ‘ears’. The caudal lobes are rounded and uneven, with the left lobe being longer and the right rather shorter. The sensory bristles at the front of the head can be seen clearly in this fantastic photograph (far right) and are typical of species of Chelidonura. Biology and ecology: Chelidonura flavolobata has a peculiar habit of waving its head and anterior body when disturbed. It is often found crawling on live coral, as does the similar Chelidonura sp. (left), due to the presence of tiny beige flatworms, Waminoa, on which they feed. Several animals may occur together, up to 50 on sandy sea bed with sea grasses during spring presumably for breeding (Dive master, Dahab, Egypt, pers. comm. 1991). They mate in a ring and lay a cocoon-like egg mass around their bodies before slipping out to leave the spawn attached below the substrate. 76 Species accounts

Order Cephalaspidea

Family Aglajidae

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Spawning near Heteroxenia B&SK

Chelidonura flavolobata

Occurrence:

Common on coral reefs, especially on brain corals, in coral rubble, on Heteroxenia colonies, and on the surrounding sand; not recorded from the southern Red Sea. Geographical distribution: Endemic to the Red Sea. Note: Two similar animals have been photographed in Eilat, but without specimens their identities must remain unknown (far left). See also p. 271 in the next chapter. Species accounts 77

OL

Order Cephalaspidea

Family Aglajidae

Chelidonura fulvipunctata

OL

Chelidonura fulvipunctata Baba, 1938 Size: 10 mm Diagnostic characteristics: The Red Sea animals are rather dark, brown to black, with small orange spots scattered on both shields, the parapodia, and the tail. The left tail extends into a very long filament which is not clearly visible in this photograph. There is a distinctive white “w” on the head, and the tip of the head shield and the top distal edge of the posterior shield are both marked with white pigment. Occurrence: Two photographic records from the Gulf of Eilat. Geographical distribution: First records for the Red Sea, otherwise West Pacific, South Africa, and Lessepsian migrant to the Mediterranean, where it grows to 30 mm. Chelidonura livida Yonow, 1994 Size: 50 mm Diagnostic characteristics:

Chelidonura livida

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The body colour is black or very dark brown (paler in juveniles) with electric blue rings or spots on the head and along the parapodial edges. The tails are uneven, the left longer than the right, and also have spots on them. The sensory bristles are often impossible to see in aglajids, but are visible on the anterior edges of both the juvenile and the adult of this species.

78 Species accounts

Order Cephalaspidea

Family Aglajidae

Biology and ecology: Chelidonura livida congregate in spring and early summer for breeding, mating in chains. One animal can be seen laying its cocoon-like egg mass (below right). Chelidonura livida can produce brownish yellow secretions when handled, which is toxic to fish (Underwater Observatory, Eilat, Israel, pers. comm. 1983). They can be found in shallow water on sandy/silty sea beds, in sea grass beds, most often at night. They can escape predators by burrowing into the soft sediment (T. Paulus, pers. comm.). Occurrence: Common especially in the Gulf of Eilat, less common in the main body of the Red Sea, not recorded from the southern half. Geographical distribution: Recorded from the Arabian seas and East Africa.

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Chelidonura livida

Spawning B&SK

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Juvenile OL

Chelidonura punctata Eliot, 1903 Size: 25 mm Diagnostic characteristics: The body is black with coral red spots on both the dorsal and ventral surfaces. The left tail is longer than the right, both fairly short in comparison to some of the other species in the Red Sea. There may be a faint white margin on the posterior end of the head shield and the edges of the parapodia. Species accounts 79

Order Cephalaspidea

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Family Aglajidae

Chelidonura punctata

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Occurrence:

Fairly common in the Gulf of Eilat, especially in April (B. & S. Koretz, pers. comm.) as evidenced from numerous photographs; no records from elsewhere in the Red Sea. Geographical distribution: New record for the Red Sea; otherwise western Indian Ocean (Yonow & Hayward 1991). Chelidonura sandrana Rudman, 1973 Size: 10 mm Diagnostic characteristics:

This tiny chelidonurid has a long left caudal lobe which can be very pointed, and a translucent zone in front of the eyes. The body colour can vary from pure black to black with speckles and mottles of white and/or orange.

10 mm

TP

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Chelidonura sandrana

80 Species accounts

OL

Order Cephalaspidea

Family Aglajidae

Biology and ecology: Amongst sea grasses, sand and silt, especially in spring and summer; Chelidonura sandrana is one of the prey items of Philinopsis cyanea (Yonow 1992). Occurrence: Common in localized areas, two to several animals crawling on the silty/sandy seabed. Limited to shallow waters 2–12 m depth in the Gulf of Eilat. Geographical distribution: Red Sea and western Indian Ocean: Zanzibar, Maldives, Chagos (Yonow et al. 2002), and Aldabra as C. babai Gosliner (syn. nov.).

Philinopsis cyanea (Martens, 1879) Size: 80 mm Diagnostic characteristics: A stumpy chelidonurid but the largest of the family which occurs in the Red Sea; fairly squared in shape and holding the tip of the head shield upright. The colours and patterns are extremely variable, from black usually with blue edging, to reddish brown or grey blue, sometimes with white and yellow markings and blue and black margins.

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Philinopsis cyanea

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Species accounts 81

Order Cephalaspidea

Family Aglajidae

Philinopsis cyanea

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Philinopsis cyanea is an active predator, feeding on other cephalaspideans such as Bulla ampulla, Chelidonura sandrana and Ringicula (see Yonow 1992). The prey item is engulfed, digested, and a clean shell is regurgitated. Breeding swarms occur during spring in Eilat (B. & S. Koretz, pers. comm.); the cocoon-like egg mass can be found on soft substrates (left, Réunion). Occurrence: Fairly common, especially in the spring, in shallow sandy or silty sea beds. Geographical distribution: Indo-West Pacific.

Biology and ecology:

P. Bidgrain

Philinopsis reticulata (Eliot, 1904) [= Aglaja phaeoreticulata Yonow, 1990] Size: 40 mm Diagnostic characteristics:

The body is moderately elongated, and the head shield bears a constriction anteriorly. The body is pale cream in colour with a brown to black network on both the dorsal and the ventral surfaces. There are violet edges to the front and tip of the head shield, the edges of the caudal lobes, and the tips of the posterior lobes, which are short and rounded. Sensory bristles are present anteriorly. There seem to be two colour forms in the Red Sea: the second (right, upper) is typical of East Africa and has bright blue and pale orange spots around the parapodial margins.

82 Species accounts

Order Cephalaspidea

Family Aglajidae

Occurrence:

Two specimens found in shallow coral rubble in Aqaba and in Eilat; several photographic records of the typical colour form from the Eilat and the central Red Sea; usually on or in coral sand and silt; numerous photos of the East African colour form from the Gulf of Eilat. Geographical distribution: Red Sea and East Africa.

East African form

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Ventral view

Philinopsis reticulata

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Odontaglaja guamensis Rudman, 1978 Size: 15 mm Diagnostic characteristics: The body varies from white to cream, with overtones of red, brown, and pale green reticulations and speckles. Some of the spaces within these reticulations have small yellow spots. There are two distinctive triangular patches near the eyes; similarly coloured patches can be found on the posterior half of the animal, both with black dots marking the edges. The left lobe is rather solid and long, while the right lobe is truncated. Species accounts 83

Spawning B&SK

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Order Cephalaspidea

Family Aglajidae Occurrence: Several photographic records from the Gulf of Eilat. Geographical distribution: First records for the Red Sea; otherwise tropical

West Pacific.

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Odontaglaja guamensis

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84 Species accounts

B&SK

Order Cephalaspidea

Family Gastropteridae

Family Gastropteridae Gastropteridae are small cephalaspideans which have paired parapodia, a pronounced dorsal hump, and a posterior funnel on the cephalic shield. A ctenidium is present on the right side of the midline, very obvious in some species. The shell is lacking in the genus Gastropteron and reduced to an internal, thin, cap-like scale in Sagaminopteron. Pacific species of Sagaminopteron appear to be associated with sponge habitats on coral reefs. The following species are the first records for this family in the Red Sea (Yonow 2000). Sagaminopteron cf. ornatum Tokioka and Baba, 1964 Size: 15 mm Diagnostic characteristics: The body varies from translucent violet to opaque lilac, with dense white markings and a thick orange band along most of the edges of the body and the edges of the parapodial lobes. Occurrence: Two photographic records from the Gulf of Eilat. Geographical distribution: This is similar to S. ornatum from Japan and Western Australia, which is deep magenta pink with fine white parapodial margins and golden orange tips on the sheilds. However, without specimens, no formal description can be made; the Red Sea species may be a colour or geographical variant, but most probably is a valid species. JH

Sagaminopteron cf. ornatum

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Species accounts 85

Family Gastropteridae

Order Cephalaspidea

Sagaminopteron cf. psychedelicum Carlson and Hoff, 1974 Size: 15 mm Diagnostic characteristics: The prominent dorsal hump is partially enclosed by the parapodia, but the translucent gills are visible. Also diagnostic is the upturned and greatly extended posterior end of the cephalic shield. Ground colour varies from cream to grey, with orange to brownish orange irregular patches surrounded by a dense black line.

OL

Sagaminopteron cf. psychedelicum

86 Species accounts

OL

PK

Family Gastropteridae

Order Cephalaspidea Biology and ecology: All records are from coral reef habitats; Sagaminopteron cf. psychedelicum seems to prefer coral rubble with fine algal growth although members of the genus are thought to be carnivorous. Occurrence: Numerous photographic records, one from the central Red Sea and the remainder from the Gulf of Eilat. Geographical distribution: Red Sea and Réunion (numerous colour photographs, P. Bidgrain). Notes: The Red Sea animals are similar to S. psychedelicum from sponge habitats in Guam and Papua New Guinea but with consistent differences in colour. In particular, the tip of the siphon is bright red in the Pacific species, even in juveniles, but this red pigment is absent in the individuals from the Red Sea and Réunion. Additionally, the ground colour between the patches has a definite green hue in the Pacific material. The Red Sea and Indian Ocean species appears identical and distinct from its West Pacific counterpart.

Siphopteron sp. Size: 5 mm Diagnostic characteristics:

The bulbous body has a long tail. It is lemon yellow with dark chocolate brown edging, some additional chocolate markings on the parapodia and across the posterior of the hump, and white areas composed of small speckles. The tail is yellow with no brown markings but has white streak centrally. Occurrence: Single specimen record from Quseir, found in coral rubble on the reef flats (April 2000). Geographical distribution: First record of the genus for the Red Sea; similar to three of several yellow species from coral habitats in the western Pacific, Siphopteron quadrispinosum Gosliner, S. brunneomarginatum Carlson & Hoff, and S. nigromarginatum Gosliner.

5 mm

Siphopteron sp. 5 mm

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Species accounts 87

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Order Cephalaspidea

Family Haminoeidae

Family Haminoeidae The external shell is large and inflated and can accommodate the body in defence. It is thin and fragile, transparent in life, rarely with stripes or bands. The animal has a large cephalic shield with posterior flaps and a pair of eyes medially. The parapodial lobes extend over some or all of the shell; a posterior mantle lobe, which makes the metapodium, may also have a lobe extending up over the shell. All species are herbivorous, with a broad radula and three grinding plates in the gizzard, and can be found grazing in turfing green algae.

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Atys naucum (Linnaeus, 1758) Size: up to 20 mm (juveniles) Diagnostic characteristics: These are inflated bubble shells which can grow almost as large as Bulla ampulla (see p. 95); the adult shells are glossy white but only juveniles have been photographed from the Red Sea. The juvenile shells bear from few to many brown longitudinal striations which are lacking in the adult. The animal itself is translucent or opaque white, with the eyes very visible at the base of the head lobes. Occurrence: Uncommon to rare in the Gulf of Eilat; few photographic records from elsewhere in the Red Sea. Geographical distribution: Indo-West Pacific to 45 mm shell (Eisenberg 1981).

Atys naucum

B&SK

88 Species accounts

Order Cephalaspidea

Family Haminoeidae

B&SK

Atys semistriatus Pease, 1860 Size: 13 mm alive (6.5 mm shell) Diagnostic characteristics: The bulbous shell is translucent white with incised striations or grooves at the apex and base but absent from the widest part of the shell. The animal within the shell can have brown speckling collected in patches; this speckling is lacking on the head shield, lobes, mantle, and foot. Other individuals may have a few dark flecks on the foot. The body also displays some glistening white patches, especially on the foot. Biology and ecology: Nocturnal, feeding on fine algae growing on dead coral and other substrata. Occurrence: Rarely recorded from coral rubble and sea grass beds; few photographic records from the Gulf of Eilat. Geographical distribution: Recorded only from the Red Sea and West Pacific. Species accounts 89

Atys semistriatus

OL

Order Cephalaspidea

B&SK

Family Haminoeidae

Diniatys dentifer (A. Adams in Sowerby, 1850) Size: 5 mm alive Diagnostic characteristics: The shell is globular, white, and with a distinctive tooth in the base of the columella. In life the shell is translucent and the animal is cream to pale green with opaque white flecks; the brown patch between the eyes appears to be distinctive. Occurrence: Diniatys dentifer is rarely recorded, presumably due to its small size and cryptic behaviour. Several photographic records from the Gulf of Eilat; previously recorded only from the Gulf of Suez. Geographical distribution: Red Sea and western Pacific Ocean.

Diniatys dentifer

OL

90 Species accounts

Order Cephalaspidea

Family Haminoeidae

B&SK

Diniatys dubius (Schepman, 1913) Size: 10 mm alive Diagnostic characteristics: The shell is very similar to that of Diniatys dentifer but without the tooth on the columella. The animal is more colourful, usually brownish with white and red speckling. The red speckling tends to be concentrated in patches. Biology and ecology: Forms breeding aggregations during the autumn in grassy areas on the reef; usually encountered at night. Occurrence: Uncommon in the Gulf of Eilat except during breeding; several photographic records. Geographical distribution: First records for the Red Sea, otherwise recorded from the West Pacific. Diniatys dubius

Haminoea cyanomarginata Heller and Thompson, 1983 Size: 6 mm alive Diagnostic characteristics: The translucent globular shell is smooth and shiny. The body is semi-translucent to opaque white, and the margins are purple blue (type specimen) to purple. There may be a purple patch present between the eyes extending forward and the purple ‘corners’ on the head are diffuse at their edges. The pigmented band around the foot (posterior mantle lobe) may be diffuse or distinct. Species accounts 91

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Order Cephalaspidea

OL

Family Haminoeidae

Haminoea cyanomarginata 6 mm

Occurrence: Several photographic records from the Gulf of Eilat. Geographical distribution: Endemic to the Red Sea, but established in

JH

the

Mediterranean Sea. Liloa curta (A. Adams in Sowerby, 1850) Size: 15 mm alive Diagnostic characteristics:

The shell is shiny and transparent, very cylindrical with a large aperture. The animal is creamy with brown speckles occurring in patches and varying in colour from beige to brown, pale green and pale orange. There are larger black flecks scattered on the mantle as well as small areas of glistening white, especially around the edges of the head and body lobes. The posterior end appears distinctive, with the lobe extending vertically beyond the shell. Biology and ecology: Generally nocturnal, living in coral reef habitats, grassy areas, coral rubble, sand, and silt. Occurrence: Numerous photographic records from the Gulf of Eilat. Geographical distribution: First records for the Red Sea, currently recorded only from the West Pacific. 92 Species accounts

Order Cephalaspidea

Family Smaragdinellidae

OL

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Liloa curta

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OL

Family Smaragdinellidae The family is small, comprising fewer than ten species distributed in two genera. The body form is similar to that of chelidonurid opisthobranchs, with two shields and parapodial lobes, and a reduced internal shell. All species are herbivorous with a broad radula consisting of many rows of hooked teeth. A gizzard is present. Habitat is variable, but specimens are usually found in eel grass beds and on rock and coral reefs. Phanerophthalmus means ‘visible eyes’ in Greek. Phanerophthalmus albocollaris Heller and Thompson, 1983 Size: 10 mm Diagnostic characteristics: The body is elongated and compact, translucent in colour with a lilac tinge. The most obvious marking is a dense white patch located on the posterior edge of the head shield, usually in the shape of a “W”. There is faint brown mottling on the rest of the animal’s body. Biology and ecology: Usually found at night, on coral reef slopes. Occurrence: Type specimen from the Sudanese coast, otherwise only few photographic records from the Gulf of Eilat. Geographical distribution: Endemic to the Red Sea. Species accounts 93

Phanerophthalmus albocollaris

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Order Cephalaspidea

Family Smaragdinellidae

Phanerophthalmus olivaceus (Ehrenberg, 1828) [= P. smaragdina (Rüppell & Leuckart, 1830)] Size: 50 mm but usually less than 20 mm in length Diagnostic characteristics: The elongated body has a distinctive head shield, with a small rounded notch in the posterior border. The parapodia are held very close to the body. An internal shell is present, cup-shaped with a point at one end. The body colour is translucent or opaque olive green with darker green speckles lacking along the edges. Biology and ecology: Found in rock pools, coral rubble with turfing algal growth, and eel grass beds. Occurrence: Three specimen records and several photographic records from northern Egypt and the Gulf of Eilat; probably not uncommon in its correct habitat. Geographical distribution: Tropical Indo-West Pacific. Notes: This species was described as new almost two centuries ago by Rüppell & Leuckart (1830) as Bulla smaragdina, nearly coinciding with Ehrenberg’s publication of a colour plate (1828) of Cryptophthalmus olivaceus. Vayssière (1912) subsequently recorded it as C. olivaceus Ehrenberg, which is the earliest name and the correct one (Yonow 2000). Ehrenberg’s and Rüppell & Leuckart’s illustrations of their new species are represented on pages 37 and 45. JH

Phanerophthalmus olivaceus

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94 Species accounts

Order Cephalaspidea

Family Bullidae

Family Bullidae These are the typical “bubble shells” of collectors, comprising the single genus Bulla. The smooth solid shell is ovoid with various markings of browns, white, purple greys, and reddish browns. The shell is heavily calcified, and can contain the whole animal when required. The animals are often brighter in colour than the shell, particularly the head shield. The eyes are conspicuous. All species are herbivorous, living in sea grass beds and intertidal algal mattes, emerging in vast numbers in the evening and at night to feed (Yonow & Hayward 1991). Bulla ampulla Linnaeus, 1758 Size: shell up to 60 mm in length Diagnostic characteristics: The animal is large and fleshy with a spatulate head. The body is usually beige with small light brown patches and tinges of orange medially and in front of the shell. The shell is light brown with darker brown patches of various intensities and sizes and can have purple tinges. Biology and ecology: Bulla ampulla lives buried in sand or sand with gravel, under stones, and in seaweed beds during the day, and is therefore difficult to find. It is a shallow water species. Eggs are laid in a zig zag fashion on sea grass blades; the sexually mature animal in this photograph was only 15 mm long. Occurrence: Most often recorded as dead shells, should be common in the correct habitat especially during the evening and at night. Geographical distribution: Common throughout the tropical Indo-West Pacific.

Bulla ampulla Spawning, 15 mm

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Species accounts 95

Order Anaspidea

Family Aplysiidae

Order Anaspidea These opisthobranchs are characterized by two pairs of rolled tentacles on the head, a pair of large parapodia often adapted for swimming, and a usually plump body which can attain a substantial size. The shell, if present, is a small structure found internally protecting the heart and the ctenidium, although it is larger and more substantial in Akera. All species have a spermathecal groove visible on the right side of the body; sometimes the eversible penis can be visible at the end of the groove below the right rhinophore. Family Aplysiidae Sea hares are fleshy opisthobranchs which can grow to a very large size. An internal shell is present, located dorsally between the parapodial lobes, and is sometimes visible through the mantle foramen in species of Aplysia. The elongated head bears rolled tentacles held upright, hence its common name the ‘sea hare’. Many species produce a smoke screen of violet-coloured ink when the animal is disturbed. Aplysiids are herbivorous, feeding on a range of algae and seaweeds, changing diet composition as they grow. Aplysia dactylomela Rang, 1828 Size: 400 mm (Eales 1960) Diagnostic characteristics: The body is coloured greenish brown, green, or yellowish green with irregularly sized but large smudged black rings on the parapodia, connected to each other with black reticulations. The parapodia are fairly upstanding and also have large black rings on the inner surfaces; they join each other low on the tail. In the photograph of the species swimming, the large ctenidium is clearly visible on the right side of the animal. Biology and ecology: The breeding season ranges from April to October in the northern Red Sea, where the sea hares can form long chains (J. Hinterkircher, pers. comm.).

Aplysia dactylomela

JK

96 Species accounts

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Order Anaspidea

Family Aplysiidae

Aplysia dactylomela Note gill

PK

Occurrence:

Uncommon, lives in the intertidal and shallow sublittoral in moderately sheltered bays; few photographic records. Geographical distribution: Circumtropical; it was illustrated from the Red Sea by Rüppell & Leuckart as Aplysia argus; see p. 41 for their beautiful illustration (Tab. 7 fig. 1). Aplysia oculifera Adams and Reeve, 1850 Size: 125 mm (Eales 1979) Diagnostic characteristics: Small

dark rings cover the body, which also bears fine reticulations. The basic colour is olive green or pale brown to dark brown. Additionally, there are clusters of opaque white dots covering the parapodia and head. The head is relatively large, forming approximately one third of the total length. The parapodia are joined low on the tail. Biology and ecology: Aplysia oculifera is known to feed on Ulva and Enteromorpha. The presence of adults coincides with algal blooms from December to June, demonstrating a seasonal life cycle, but is variable from year to year and place to place (Plaut et al. 1998). Both Aplysia oculifera and Notarchus indicus are common in sea grass beds during the spring algal blooms in the Gulf of Eilat; they often occur with Melibe rangii (p. 237). A tangled mass of rather sticky strands are the typical spawn of Species accounts 97

Egg mass

TP

Order Anaspidea

Family Aplysiidae

A. oculifera (left) and N. indicus (right)

Aplysia oculifera

JH

HS

TP

Aplysia; they may be shades of pink or yellow depending upon stage of development. The egg masses probably belong to A. oculifera; the photograph was taken in the Gulf of Eilat in March. Occurrence: Intertidal and subtidal in sea grass beds; appears to be the most common aplysiid species recently recorded in the Red Sea. Geographical distribution: Indo-Pacific Ocean. Aplysia parvula Guilding in Mörch, 1863 50 mm Diagnostic characteristics: The colour can be extremely variable in Aplysia parvula depending upon diet but is always in shades of pale brown or green, speckled with white. The upstanding parapodia are edged with purple darkening to black, and the ends of the tentacles and the edge of the foot are also often lined in darker pigment. Biology and ecology: Intertidal and shallow sublittoral, often at night, usually in sea grass beds. Appears to be able to attach itself by the tail and wave its head like Aplysia sp. Occurrence: Numerous photographic records, from the central Red Sea and Gulf of Eilat. Geographical distribution: Circumtropical. Size:

98 Species accounts

Order Anaspidea

Aplysia parvula

Family Aplysiidae

B&SK

B&SK

OL

OL

B&SK

Species accounts 99

Order Anaspidea

Family Aplysiidae

Aplysia sp. Size: 25 mm

OL

Diagnostic characteristics: Most usually black but sometimes a very dark brown, this tiny aplysiid has a red or pink rim (sometimes cream) to the parapodia, rhinophores, and oral tentacles. Sometimes there are white flecks on the body and parapodia. The upstanding parapodia are joined high up posteriorly, and the siphon is usually visible in the last convolution. The porcelain-like shell is clearly visible through the mantle foramen. Biology and ecology: Emits purple dye when disturbed, difficult to detach from substratum; most frequently on exposed reef faces and coral reef flats.

16 mm

G. Bemert

Aplysia sp. OL

B&SK

100 Species accounts

Order Anaspidea

Family Aplysiidae

Uncommon, but found more frequently than Aplysia parvula; appears to be restricted to the northern half of the Red Sea. Geographical distribution: A new species known only from the Maldives, Chagos, and Red Sea (Yonow et al. 2002). Occurrence:

Dolabella auricularia (Lightfoot, 1786) Size: 300 mm Diagnostic characteristics: The solid and chunky body has a flat posterior disc which is distinctive. The general appearance is knobbly, and the pointed papillae can be quite large. Dolabella auricularia is usually dull browns and greens in colour, extremely well camouflaged, but sometimes, as in Eilat, they can have lilac patches or hues. The large rolled rhinophores are characteristic of the order. The internal shell is large and heavily calcified, located in the flattened posterior disc. Biology and ecology: Dolabella auricularia is recorded as breeding from April to October in the northern Red Sea (J. Hinterkircher, pers. comm.). Under duress, it can squirt a jet of ink if it feels threatened (J. Dafni, pers. comm.). Occurrence: Uncommon in sandy and muddy substrates with boulders and sea grasses, and on shallow subtidal reef flats.

JK

Dolabella auricularia

Eilat B&SK

PK

GS

Species accounts 101

Order Anaspidea

Family Aplysiidae Geographical distribution: The two species, D. auricularia (Lightfoot) and D. gigas Rang are similar externally, currently only distinguishable by their shells. In fact, only the shell of D. gigas exists in the museums; no one has found a living animal. Dolabella auricularia is certainly the more common species, recorded under a number of synonyms, and has a wide tropical Indo-Pacific distribution (Yonow & Hayward 1991).

Dolabrifera dolabrifera (Cuvier, 1817) Size: 50 mm Diagnostic characteristics:

JH

This species is well camouflaged, usually coloured in drab browns and greens, but ranging to beiges and even pale pinks. The body is much flattened with a posterior hump and a broad foot, but does not have the ‘cut-off ’ disc appearance of Dolabella auricularia, so juveniles of the larger Dolabella cannot be confused with this smaller species. Variation in colour pattern, presumably for camouflage, is quite spectacular, as is flexibility of the body. Rarely, the mantle can bear small branching papillae. The small internal shell is bat-shaped. Biology and ecology: Hiding under rocks in the intertidal and very shallow waters of sandy or muddy substrates with rocks and stones in daytime; active during the night. Dolabrifera dolabrifera curls up like a chiton when disturbed. Occurrence: Common. Geographical distribution: Circumtropical.

Dolabrifera dolabrifera

JH

102 Species accounts

Order Anaspidea

Family Aplysiidae

JH

Petalifera ramosa Baba, 1959 Size: 30 mm Diagnostic characteristics: Petalifera ramosa is characterised by a greenish brown body, sometimes rather translucent, covered in firm conical tubercles with white tips. The smaller ones can be purple or pink with a darker basal ring and the larger ones are the same colour as the body but with a white ring edged in brown at the base. Some of the larger tubercles have a terminal papilla which has long filamentous branches. The eyes are set in clear circular patches near the bases of the rhinophores. The vestigial shell is internal, allowing for great body flexibility.

Petalifera ramosa

B&SK

Species accounts 103

Dolabrifera dolabrifera

JH

B&SK

Order Anaspidea

B&SK

Family Aplysiidae

Petalifera ramosa

B&SK

Occurrence:

Numerous colour photographs from the Gulf of Eilat; common in sea grass beds during the spring and summer (B. & S. Koretz, pers. comm.). Geographical distribution: First records for the Red Sea; otherwise only known from the western Pacific (Baba 1959). Phyllaplysia sp. Size: ? 10 mm Diagnostic characteristics:

The body is small and flattened, very similar to that of Phanerophthalmus (see p. 94), and is bright green with a row of small black spots along the parapodial edges. There is a faint linear pattern of green along the parapodia. The tentacles have black pigment on the distal portion; however, the rhinophores are well camouflaged, much paler and almost invisible, presumably protecting these vital sensory organs from predators. The eyes are visible in a diffuse brown area beside each rhinophore base. Occurrence: Single photographic record from Jeddah, Saudi Arabia; second record of the genus from the Red Sea (see table p. 55). Geographical distribution: Unknown; the genus is not well studied and numerous species are thought to co-exist in the Indo-West Pacific.

Phyllaplysia sp.

104 Species accounts

JK

Order Anaspidea

Family Notarchidae

Family Notarchidae Body form in this family is variable; however, all are without an internal shell and with contractile papillae. The size of these can vary considerably but their function is not known. There is a wide mantle cavity with a relatively large gill. The radula is well developed. Some species may have coloured eyespots. All species are herbivorous. Notarchus indicus Schweigger, 1820 Size: 55 mm Diagnostic characteristics: The plump body is globular in shape, translucent white, orange, brown, or green in colour, with black, brown, and white speckling. The brown and black speckles can form faint rings or reticulations, while the white concentrates on the papillae. The parapodia are almost completely fused together, enabling the species to swim with a crazy somersaulting action. The body is papillate, and the size of these can be variable.

B&SK

Notarchus indicus

B&SK

Species accounts 105

Order Anaspidea

B&SK

Family Notarchidae

Notarchus indicus

TP

Biology and ecology:

Common during February to May, when algal food is at its maximum abundance and breeding occurs. Aplysia oculifera and N. indicus are often found with Melibe rangii (p. 237). Swarms can occur on soft sediments in the early spring, with up to 100 animals per square meter (B. & S. Koretz, pers. comm.). When disturbed, it uses jet propulsion to launch itself into the water column in a series of somersaults (see figure on p. 30). Occurrence: Occurs in calm shallow waters in algal beds and sea grasses around coral heads, especially in the Gulf of Eilat. Geographical distribution: Circumtropical; N. indicus has probably entered the Mediterranean from the Red Sea through the Suez Canal. Stylocheilus longicauda (Quoy and Gaimard, 1824) [= S. citrina (Rang, 1828)] Size: ? 60 mm Diagnostic characteristics: Animals in the Red Sea are bright orange yellow, slightly translucent, with few to many large pink spots ringed with red. The body is rounded, soft and smooth, and despite its name, the long tail is not visible here. The rolled oral tentacles and rhinophores are very long, uniformly coloured with the body. 106 Species accounts

Order Anaspidea

Family Notarchidae

Stylocheilus longicauda lives on drifting algae in the open ocean, and can be found washed up on shores. Occurrence: Three photographic records, one from Hurghada in the northern Red Sea and two from Jeddah in the central Red Sea. Geographical distribution: Circumtropical. Biology and ecology:

JK

Stylocheilus longicauda

N. Kenward

Species accounts 107

Order Anaspidea

Family Notarchidae

Stylocheilus striatus (Quoy and Gaimard, 1832) Size: 75 mm Diagnostic characteristics: The body is translucent beige or brown with long thin brown lines and blue violet eye spots. There are two long pairs of tentacles, and the eyes are visible near the base of the rhinophores. The body is covered with simple and compound papillae. Their function and the reason for their variability are not known. Occurrence: Rarely observed, but seasonally common during spring algal blooms; many photographic records from the Gulf of Eilat, Egypt, and Saudi Arabia. Geographical distribution: Tropical Indo-Pacific and Atlantic oceans.

JH

B&SK

Stylocheilus striatus

108 Species accounts

Head JH

JK

JH

Order Pleurobranchida

Family Pleurobranchidae

Order Pleurobranchida The Pleurobranchida are a small order with members bearing a bipinnate gill on the right side of the body between the mantle and the foot, giving rise to the name. The oral veil is trapezoidal with rolled oral tentacles at its edges. The rolled rhinophores are often joined at their bases, protruding in front of the mantle or through a groove in the mantle above the oral veil. Family Pleurobranchidae Pleurobranchs are opisthobranchs with soft oval bodies. There may or may not be tubercles on the mantle. A reduced shell is present internally in most species. Ventrally, the foot has a posterior pedal gland which is very visible in a number of species. The skin is able to secrete sulphuric acid as a defence mechanism. Species of Pleurobranchus are thought to feed on ascidians but Berthellina feeds mostly on sponges and occasionally on corals. Berthellina citrina (Rüppell and Leuckart, 1828) Size: 65 mm, usually approximately 25 mm Diagnostic characteristics: The body colour varies from pale yellow orange to bright yellow and bright orange. Animals are often translucent enough to see the gut beneath the smooth notum. They can be rather slimy to handle. There are no markings on the dorsum as in Berthella stellata. The foot is usually visible, being as large as or slightly larger than the mantle, and the oral veil is obvious.

Berthellina citrina

JK

Species accounts 109

Order Pleurobranchida

Berthellina citrina

Family Pleurobranchidae

JK

JK

Biology and ecology:

Eggs are laid during the spring and summer, an orange red coiled ribbon up to 25 mm across (Gohar & Aboul-Ela 1957a). Experiments have shown that the skin secretions are acidic at pH 1 and that B. citrina was rejected as prey by its potential predators such as sea anemones, fish, crabs, and hermit crabs (Marbach & Tsurnamal 1973). Berthellina citrina is generally nocturnal. Occurrence: Very common, usually in the intertidal and shallow sublittoral under rocks on sandy sea beds, often more than one individual together. Geographical distribution: Indo-Pacific and now Mediterranean (Lessepsian migrant through the Suez Canal). The original figure of the type species for the genus is reproduced here on p. 40 (Tab. 5 figs. 1a – c). Pleurobranchus albiguttatus (Bergh, 1905) [= P. oblongus Audouin, 1826] Size: 35 mm Diagnostic characteristics:

The oblong body is translucent with brown and white pigment on the rounded pustules, which have angular bases. Larger and darker pustules are present in a paired series along the dorsum and have a triangular white pigment patch on the surface facing the midline, making them appear pointed. The large rhinophores are white on their distal upper surfaces. The foot is usually visible beyond the mantle margin. Biology and ecology: Juveniles are paler in colour and have a border of orange around the mantle which disappears as the animals grow: it is most persistent anteriorly behind the rhinophores (opposite, middle right). Occurrence: Numerous photographic records from the Gulf of Eilat and the central Red Sea; one specimen from Quseir, Egypt; the cluster of specimens was found under a rock in shallow water on the reef flat at Quseir (April 2000) (opposite, centre left). Geographical distribution: Indo-West Pacific. Notes: This is almost certainly the species illustrated by Savigny and recognized by O’Donoghue (1929) as Pleurobranchus oblongus (syn. nov.) (p. 35). Rüppell & Leuckart also have it listed amongst their collection retained at the Senckenberg Museum (Frankfurt, pers. obs.). It is unfortunate that P. oblongus has no type specimen or description, and therefore the name cannot be reinstated. The oblong shape, the slightly larger foot, the dorsal pustules, and the size are a perfect match for P. albiguttatus; no other species in the Red Sea has this combination of characters. 110 Species accounts

Order Pleurobranchida

Family Pleurobranchidae

B&SK

Pleurobranchus albiguttatus

JH

JK

Juvenile JK

Paraplanocera sp. JK

Thysanozoon sp. JK

Species accounts 111

Order Pleurobranchida

Family Pleurobranchidae

Pleurobranchus forskalii Rüppell and Leuckart, 1828 Size: 100 mm Diagnostic characteristics: The mantle is irregularly distributed with evenly sized tubercles: it may be white or cream with red semi-lunar markings or red with white semi-lunar markings. The foot is similar to the mantle, and often much larger. Posteriorly the mantle may be held to produce a small siphon as in P. grandis but there are usually several of these little siphons produced all around the mantle. See p. 40 for Rüppell & Leuckart’s painting of this species (Tab. 5 fig. 2). Very few photographs of small individuals thought to be juveniles may be an unrecorded species, P. tesselatus Pease. Occurrence: Rarely recorded; few photographic records from the Red Sea and Gulf of Eilat. Geographical distribution: Red Sea and tropical Indo-West Pacific.

JK

Juvenile or P. tesselatus ? JK

Pleurobranchus forskalii 70 mm

112 Species accounts

B&SK

Order Pleurobranchida

Family Pleurobranchidae

B&SK

B&SK

Juvenile or Pleurobranchus tesselatus ? 30 mm

Pleurobranchus forskalii 150 mm

B&SK

JH

JH

Species accounts 113

Order Pleurobranchida

B&SK

Family Pleurobranchidae

Pleurobranchus grandis Pease, 1868 Size: 200 mm Diagnostic characteristics: Pleurobranchus grandis, as its name suggests, is very large and fleshy, with red and white polygonal patterns. Each polygon contains a large central rounded tubercle surrounded by smaller rounded tubercles. The white areas are usually arranged in two series, one each side of the midline, and around the margin. The white may be yellow or even brown in some specimens. There is an obvious, very large posterior crenulation or siphon, which is lacking in specimens of Pleurobranchus cf. grandis, and the foot is often bicoloured. Biology and ecology: Juvenile specimens (50 mm) are similar to the adults (B. & S. Koretz, pers. comm.). A close-up of the ctenidium, found on the right side of the animal, reveals its tubercle and pinnule arrangement. The mantle is often flapped up and down to aerate the gill. This appears to be the first record of the shrimp Periclimenes imperator Bruce living on a species of pleurobranch; it is usually found on Hexabranchus (see p. 144), sea cucumbers, and cushion stars. Pleurobranchus grandis is generally nocturnal. Occurrence: Numerous photographic records from shallow waters, often seen at night. Geographical distribution: Tropical West Pacific Ocean and the Red Sea; there are no published records from the Indian Ocean (Yonow 2000).

Pleurobranchus grandis

B&SK

114 Species accounts

Order Pleurobranchida

Gill

Pleurobranchus grandis

A. Double

Family Pleurobranchidae

With shrimp

A. Double

B&SK

B&SK

Pleurobranchus cf. grandis Pease, 1868 Size: 110 mm Diagnostic characteristics: The body is large and soft, oval in shape, with the dorsum divided into regular polygons with a central nipple-like tubercle. The foot is larger than mantle but similar in colour, deep orange red to dark red or purple, with white flecks. Both the mantle and the foot can have a white edge. There is a distinctive pedal gland on the sole of Species accounts 115

Order Pleurobranchida

Family Pleurobranchidae

the foot near the posterior end. The mantle is not held in the distinctive siphon posteriorly as in P. grandis. Occurrence: Not uncommon on coral reefs 8–15 m deep, most often found at night; numerous photographs especially from the Gulf of Eilat. Geographical distribution: Endemic to the Red Sea (Yonow 2000 as the Red Sea form of P. forskalii).

120 mm

TP

B&SK

Pleurobranchus cf. grandis

116 Species accounts

B&SK

Order Pleurobranchida

Family Pleurobranchidae

Pleurobranchus mamillatus Quoy and Gaimard, 1832 Size: 30 mm Diagnostic characteristics: The mantle is covered in groups of tubercles which are pale on a reddish background; the arrangement is encircled by a dark red or purple line followed by a white line. The central tubercle is very long in adults. The space left between the tubercle groups is very pale. The juvenile is paler than the adults, and the central tubercle in each group is not yet elongated. The rhinophores are pale for their distal halves in juveniles, otherwise orange. Occurrence: Four photographic records from the Gulf of Eilat, including two adults. Geographical distribution: First records for the Red Sea; originally described from Mauritius and possibly recorded from South Africa, it is more common in the West Pacific.

Pleurobranchus mamillatus Juvenile, 10 mm

B&SK

Species accounts 117

Order Sacoglossa

Family Oxynoidae

Order Sacoglossa The Sacoglossa are a very large order comprising more than a dozen families and hundreds of species. Almost every species is herbivorous, some may have a rounded or bivalved shell, and all have a uniseriate radula comprising a single row of dagger-like teeth with which to pierce algal cells or egg cases. Family Oxynoidae Members of the Oxynoidae are small opisthobranchs with a reduced shell into which only the head can be retracted. Four genera contain fewer than a dozen species, of which only one species can be found in the Red Sea. Oxynoe has two parapodia, which are sometimes papillate, and a very long muscular tail that can be autotomised when the animal is threatened. All species are vegetarian, feeding on species of Caulerpa, where they are well camouflaged. Oxynoe viridis (Pease, 1861) Size: approximately 10 mm Diagnostic characteristics: Oxynoe viridis occurs in a wide range of colours and morphology. The colour varies from bright or pale green to yellowish, with differing concentrations of white pigment along the edges of the parapodia, rhinophores, and tail. Equally, specimens can be muddy olive green with white papillae on the parapodial margins and oral tentacles. Both colour variations may have deep blue spots that can be surrounded by diffuse blue rings. The general appearance of the body may be smooth or spiky, but globular anteriorly with large rolled rhinophores and a large muscular tail posteriorly. Occurrence: Two specimen records from the Egyptian coastline from shallow subtidal reef flats, and one photographic record from Eilat, Israel. Geographical distribution: Wide Indo-West Pacific distribution, to 45 mm in Australia (Jensen 1997).

Oxynoe viridis

JH

118 Species accounts

S. Schrenkel

Order Sacoglossa

Family Plakobranchidae

Family Plakobranchidae (= Elysiidae) Generally small opisthobranchs, the elysiids are the largest family of sacoglossans. The body is elongated with variously developed parapodial lobes. These contain branches of the digestive gland which are visible on the inside of the parapodia. The shell is absent, as are cerata and oral tentacles. The rhinophores are rolled. All species are herbivorous, and many collect and harvest the chloroplasts from the algae for their own use. Elysia tomentosa Jensen, 1997 Size: approximately 25 mm Diagnostic characteristics: The ‘hairy’ appearance of this opisthobranch is due to white papillae branching from the parapodial margins. There are pointed papillae covering the remainder of the body, both sides of the parapodia, the head, and the rhinophores. The body is pale to mid green with white streaks and black spots; the parapodial margins are pink, although this can be very faint. The parapodia are held high and open, and when they are closed the undulating lobes form openings. Biology and ecology: Animals feed on Caulerpa in shallow sublittoral and intertidal reef flats. Occurrence: Single specimen record from Egypt (October 1995) and several photographic records from the northern half of the Red Sea and the Gulf of Eilat. Geographical distribution: First Red Sea records; Elysia tomentosa is recorded from the tropical Indo-West Pacific. JH

Elysia tomentosa

OL

OL

JH

OL

Species accounts 119

Order Sacoglossa

Family Plakobranchidae

B&SK

Elysia trilobata Heller and Thompson, 1983 Size: 15 mm Diagnostic characteristics: Elysia trilobata has a lumpy appearance due to the round tubercles scattered all over the parapodia, the body, and the rhinophores. The parapodia are unequally lobed, both in size and shape. These lobes carry more white pigment than the rest of the animal, and the tubercles each have a yellow centre. Ground colour is pale green with darker green speckles, and maroon or brown patches on the parapodia, head, and rhinophores. Additionally, blue pigment of different shades is found on the tubercles, rhinophores, head, foot corners, and body. Occurrence: Few photographic records from the Gulf of Eilat and Hurghada, Egypt; first records since the original description. Geographical distribution: Endemic to the Red Sea.

Elysia trilobata

B&SK

M. Habib

Elysia sp. 1 Size: 25 mm Diagnostic characteristics:

Elysia sp. 1

PK

The body is pale green with white streaks on the exterior parapodial surfaces, head, rhinophores, and the body generally. The large undulating parapodia have black edges and an orange submarginal band separated by very thin bright white line. There are few black spots inside parapodia (but not outside) and dense black streaks on the head extending onto the rhinophores. The orange band is absent on the rhinophore tip and the black margin can be quite extensive. The juveniles bear the colour pattern of the adults, but with fewer undulations of the parapodia (opposite, left). Rarely, individuals are found without white streaks. Biology and ecology: Localized distribution, as abundant as 1–3 animals per square meter during copulation in Sharm el Sheikh, Egypt (June 1995, J. Hinterkircher, pers. comm.). Occurrence: Not uncommon, numerous photographic records from reef flat habitats in shallow water (see notes p. 122). Geographical distribution: Endemic to the Red Sea. 120 Species accounts

Order Sacoglossa

Family Plakobranchidae

G. Reinicke

JK

Juvenile

Elysia sp. 1

PK

T. Metcalfe

K. Metcalfe

A. Valdés

Elysia sp. 2 Size: 30 mm or larger? Diagnostic characteristics:

The colour of this sacoglossan is muddy olive green, covered in small black spots and streaks which extend onto the head and rhinophores. The black marginal band is continuous along the parapodia. The yellow submarginal band may be faint or bright on the parapodia. The rhinophores have a black rim as well as a diffuse lighter submarginal band. Occurrence: Less frequently encountered than Elysia sp. 1 (see also notes over) but several photographic records; specimen record from Egypt (October 1995). Geographical distribution: Endemic to the Red Sea. Species accounts 121

Elysia sp. 2

DPS

Order Sacoglossa

Family Plakobranchidae

The precise identities of Elysia grandifolia (Kelaart) and E. ornata (Swainson) remain uncertain. Both coexist in the Indo-West Pacific but with colour variations. Additionally, their radulae and possibly their food items differ, but as so many records have been lumped together, it is very difficult to distinguish the differences between the two species. However, the two Red Sea species differ from both named species. Elysia sp. 1 is closer to what some authors call ‘ornata’ but the excess black pigment on the head does not occur in records from the Indo-West Pacific.

Notes on the previous two species:

Elysiella pusilla Bergh, 1872 Size: approximately 10 mm Diagnostic characteristics: The

body is very compact, dark green with white speckling, especially on head and parapodial margins. The parapodia are held very close to the body and do not extend over the tail, closely resembling Phanerophthalmus olivaceus (p. 94). The rolled rhinophores are mottled with beige. Biology and ecology: Feeds on the alga Halimeda, on which it is very cryptic. Occurrence: Single specimen from the Egyptian coast (September 1995). Geographical distribution: First record from the Red Sea. Elysiella pusilla is otherwise recorded from the tropical Indo-Pacific, better known as E. halimedae (Macnae).

Elysiella pusilla

A. Valdés

122 Species accounts

Order Sacoglossa

Family Plakobranchidae

Thuridilla decorata (Heller and Thompson, 1983) Size: 20 mm Diagnostic characteristics: The intricate undulating pattern of pale and dark green regions, white lines, and black spots along the edge of the parapodia is distinctive. The body and rhinophores are dark green, and the rhinophores are similarly banded with black, pale green, and white. The eyes are clearly visible in their green triangle at the base of the rhinophores. Occurrence: A common species, but difficult to find due to its small size and cryptic habit, in amongst coral rubble in shallow water; many photographic records throughout the Red Sea. Geographical distribution: Endemic to the Red Sea.

B&SK

B&SK

OL

JK

Thuridilla decorata

B&SK

11 mm

NY

Species accounts 123

Order Sacoglossa

Family Plakobranchidae

Thuridilla ? indopacifica Gosliner, 1995 Size: 20 mm Diagnostic characteristics: The body is typical of the genus, elongated in shape, and brightly coloured. It is pale and dark green on the parapodia and pale grey on the head and rhinophores, all covered in white speckles. The margins of the parapodia are orange, followed by a black line. Near this black line are bright iridescent blue patches, forming a discontinuous band along the submargin. The rhinophores are grey basally with an orange band bordered on each side by a black line and a diffuse turquoise area extending to the tip.

OL

Thuridilla ? indopacifica

B&SK

B&SK

Occurrence:

This species appears to be common in the Gulf of Eilat during late winter and early spring; photographic records only. Geographical distribution: Thuridilla indopacifica is known only from the Indian Ocean, despite its name. It differs in lacking the abundant white speckles of the Red Sea species. Unfortunately, without specimens, the precise identity of the Red Sea species will remain unknown. Plakobranchus ocellatus van Hasselt, 1824 Size: 35 mm Diagnostic characteristics: The body is dorso-ventrally flattened with the fleshy parapodia held tightly against body. It is pale green covered with ocelli varying in colour from cream, brown, beige, and white to pink and purple. Ocellations are also present ventrally. Biology and ecology: Easily found in the intertidal and very shallow subtidal, crawling and hiding in coral rubble, rock pools and lagoons, and in sand or silt. 124 Species accounts

Order Sacoglossa

Family Plakobranchidae

Occurrence:

Probably more common than records show; Vayssière’s (1912) plate including this species is reproduced on p. 47. The several specimen and photographic records are from the northern half of the Red Sea. Geographical distribution: Tropical Indo-West Pacific, with many colour variations and as many names.

H. Schmid

PK

Plakobranchus ocellatus

PK

T. Metcalfe

Species accounts 125

Order Sacoglossa

Family Polybranchiidae

Family Polybranchiidae These sacoglossans have a broad body covered in easily autotomised cerata. The cerata can be flat and leaf-shaped, angular, or inflated and cushion-like. They contain branches of the digestive gland in Polybranchia. The rhinophores are rolled and bifid at the tips. There is no shell. All species are herbivorous and some species are brilliantly coloured.

DPS

PK

Cyerce elegans Bergh, 1870 Size: 30 mm Diagnostic characteristics: The body is translucent in colour, beige to a faint orange. The large inflated cerata are oval, the scalloped edge is gold or white, and they are very easily cast off. Pale blue or green spots present on the cerata are connected to each other and the edge by faint lines, making the animal appear iridescent. The distinguishing features of the family, the rolled oral tentacles and rhinophores, the latter bifid at the tips with eyes at the bases, can be seen in this close-up of the anterior part of the animal (upper left). Occurrence: Single specimen record from the Egyptian coast and several photographic records from the central Red Sea. Geographical distribution: First Red Sea records; otherwise recorded only from the West Pacific to Hawaii.

Cyerce elegans

JK

126 Species accounts

Order Sacoglossa

Family Polybranchiidae

Swimming

Polybranchia orientalis

B&SK

JH

JK

JH

Polybranchia orientalis (Kelaart, 1858) Size: approximately 70 mm Diagnostic characteristics: The angular uneven cerata are upstanding all over the body, and look very similar to the sea grasses amongst which the species lives. The colour of the body and cerata is usually opaque, beige or pale brown to orange. The cerata have white serrated edges and are papillate. The bifurcating rolled tentacles are clearly visible in the photograph of the animal swimming (under duress; this is not a natural behaviour). Biology and ecology: Abundant during the breeding season during early spring. This may be the egg mass of Polybranchia orientalis, photographed close to the specimen (April 2000). Occurrence: Several photographic records from Quseir, Egypt, on a shallow reef flat, Jeddah, Saudi Arabia, and the remainder from Eilat, Israel, in sea grass beds at night. Geographical distribution: New species record for the Red Sea; otherwise P. orientalis has an Indo-West Pacific distribution. Species accounts 127

Suborder Doridina

Family Goniodorididae

Order Nudibranchia : Suborder Doridina The order Nudibranchia is the largest and most varied of all the opisthobranch orders. All species have lost the shell as adults. The nudibranchs are formed of four suborders, of which the Doridina is the largest. Most dorids have lamellated rhinophores and a circlet of gills around the anal papilla. Here they are loosely divided into the phanerobranch families and cryptobranch families; these are not accepted taxonomic groupings but serve a useful purpose. The Porostomata differ in lacking a radula altogether: they have a muscular buccal pump instead. Phanerobranch dorids In the Red Sea this grouping includes the more elongated species which often have a reduced or absent mantle skirt. The gills do not retract into a singe pocket, and the rhinophores may have a pocket in which to retract. Often there are elongated processes near the gills and near the rhinophores, and sometimes all around the mantle ridge. Family Goniodorididae Goniodorids are generally small dorids with a high body profile and a curled mantle skirt (Goniodoris), a fringe of pallial processes (Okenia), or conditions in-between with fewer pallial processes (Ancula, Trapania). Only the genera Goniodoridella, Goniodoris, and Trapania are found in the Red Sea. The head is enlarged with flattened lobes or long tentacles, and long tapering rhinophores with few lamellae. Species are known to feed on bryozoans and compound ascidians. Goniodoris joubini Risbec, 1928 Size: approximately 22 mm Diagnostic characteristics: The

6 mm

JH

body is high in profile, dark blue black or brown with white speckles and larger spots. The rolled mantle margin is translucent with orange or yellow spots. The head is enlarged with prominent tentacles, and the rhinophores also look disproportionately large. The metapodium is almost as long as the mantle, and covered with white speckles. The large feathery gills are brown or translucent with white midribs.

Goniodoris joubini 6 mm

JH

128 Species accounts

22 mm

B&SK

Suborder Doridina

Family Gymnodorididae

Occurrence:

Single specimen recorded from Aqaba, Jordan (June 1994) and three photographic records from under stones in shallow water near sea grass beds, Gulf of Eilat. Geographical distribution: First records for the Red Sea; otherwise only known from the western Pacific. Family Gymnodorididae These are soft phanerobranchs which have only a mantle ridge and no skirt. The name is derived from the Greek for bare or naked, and the semi-translucent species do appear insubstantial and bare. The frontal veil in species of Gymnodoris may bear pointed papillae which can be long or short. Species are generally white, pale yellow, or orange with yellow, orange, or red markings; some have no markings at all and others are wildly different in colour pattern. The gills are simply pinnate, arranged in a circlet in Gymnodoris and in an arch in front of the anal papilla in Analogium. At least three species are known to eat other opisthobranchs, but many feed on bryozoans. Analogium striatum (Eliot, 1908) Size: 50 mm Diagnostic characteristics: The body is pale creamy yellow with bright yellow orange to orange longitudinal ridges; these ridges are characteristic of this species. The gills are present in an arch across the middle of the back. The bulbous rhinophores are the same colour or darker than the stripes on the body. The edge of the foot is orange, and the sides of the body also bear orange ridges. JK

Analogium striatum

PK

Species accounts 129

Suborder Doridina

Family Gymnodorididae

JK

In the Red Sea Analogium striatum is most commonly found on coral sand, presumably hunting opisthobranch prey. In the western Pacific they feed on the sacoglossan Plakobranchus ocellatus (p. 124). Occurrence: Common in the main Red Sea, shallow subtidal, openly crawling on sand and coral rubble; only one specimen record from the Gulf of Eilat. Geographical distribution: Indo-West Pacific. Biology and ecology:

Analogium striatum

JK

Analogium sp. Size: 35 mm Diagnostic characteristics:

Analogium sp.

B&SK

The colouration of this gymnodorid is not typical: the basic colour appears translucent lilac or violet, with numerous longitudinal ridges deep red in colour. The gills, rhinophores, and frontal margin are also red. Scattered on the dorsum in-between the ridges and at the bases of the gills are opaque white patches, with a large one behind and between the rhinophores. The gills are present in an arch across the middle of the back. The edge of the foot is not coloured but its tip, the edges of the gills, and the rhinophores are red. Biology and ecology: This species occurs on coral sand and is the food item of another gymnodorid in the Red Sea, Gymnodoris alba (B. & S. Koretz, pers. comm.). Occurrence: Analogium sp. is seasonally common in the Gulf of Eilat and the very northern part of the Red Sea; numerous photographic records only. Geographical distribution: A new species endemic to the Red Sea. 130 Species accounts

Suborder Doridina

Family Gymnodorididae

Analogium sp.

B&SK

Gymnodoris alba (Bergh, 1877) Size: 50 mm Diagnostic characteristics: The body is translucent white with numerous small yellow to orange pustules all over the body. There is a keel on the posterior portion of the tail, and the tail also bears pustules. Another diagnostic feature is the enormous oral ‘lappets’ of the oral veil; these are the same colour as the pustules. The frontal margin is marked by a line of short pointed papillae, also the same colour as the pustules on the body. The gills are present in a circlet approximately half-way along the body.

JK

Cannibalism

Gymnodoris alba

B&SK

Species accounts 131

JK

Suborder Doridina

Family Gymnodorididae

Gymnodoris alba

JK

Genital papilla

JK

Biology and ecology: In the Gulf of Eilat, Gymnodoris alba feeds on Analogium sp. and appears to be active at night (B. & S. Koretz, pers. comm.). Near Jeddah, central Red Sea, the species is cannibalistic (J. Kuchinke, pers. comm.) (previous page). Occurrence: Single specimen record from Sharm el Sheikh, Egypt, and numerous photographic records from the northern and central Red Sea. Geographical distribution: Recorded from the Red Sea and West Pacific only.

Gymnodoris ceylonica (Kelaart, 1858) Size: approximately 80 mm Diagnostic characteristics: The

translucent animal bears few raised dark orange pustules on the body and around the head. The circlet of large gills is pale with the outer axes marked in dark orange. The rhinophores

Gymnodoris ceylonica Mating B&SK

132 Species accounts

JK

Suborder Doridina

Family Gymnodorididae

Gymnodoris ceylonica

JH

and edge of foot are also orange but can be paler in the darker specimens. The rhinophores are unusual for species of Gymnodoris in that they are small, held vertically and facing outwards like small horns, and have only the distal portion pigmented. Biology and ecology: The egg masses of G. ceylonica are not the neat spiral ribbons of typical dorid egg masses, but a tangled string of orange eggs similar to those of anaspids (sea hares). The developing eggs are clearly visible as a double row in a clear mucous jelly string (right). Occurrence: Found in the shallow subtidal on fine coral rubble and sand; two specimens and several photographic records from the northern and central Red Sea. Geographical distribution: Indo-West Pacific. Species accounts 133

Egg mass

JH

Suborder Doridina

Family Gymnodorididae

Gymnodoris citrina (Bergh, 1875) Size: 21 mm Diagnostic characteristics: The body is variable in colour from white to cream or pale yellow or orange. It is covered with scattered yellow orange papillae, which are often pointed. There is a margin of pointed papillae around head on the pallial rim, and characteristically a V-shape behind the gills marking the mantle margin. The rhinophores, oral tentacles, and gills are usually orange but variable in shade and intensity, often darker than the body. Variation in colour is not unusual, and one individual has pale green gills and rhinophores. Biology and ecology: This species can be found in the intertidal and on very shallow shores, on silty substrata with rocks covered in algae. Occurrence: Three specimen records and few photographic records. Geographical distribution: Records have been confused with G. bicolor but probably G. citrina has an Indo-West Pacific distribution.

PK

Gymnodoris citrina

134 Species accounts

PK

Suborder Doridina

Family Gymnodorididae

Gymnodoris impudica (Rüppell and Leuckart, 1828) [= G. rubropapulosa (Bergh, 1905)] Size: 60 mm Diagnostic characteristics: The creamy or white body is covered in orange pustules which may be completely orange or just orange at their bases. The mantle margin is vaguely marked with orange pustules, and the bulbous rhinophores are a similar shade of orange or darker. The foot bears an orange margin. The plumose gills have orange on their larger axes. Occurrence: In the literature, G. impudica has been reported from the Red Sea twice since Rüppell & Leuckart’s description but there are no recent records (see table p. 57). Geographical distribution: Widespread distribution throughout the Indo-West Pacific as G. rubropapulosa (Bergh) (syn. nov.), Gymnodoris impudica has been completely ignored in the more recent taxonomic literature. However, there is no mistaking Rüppell & Leuckart’s description and illustration (Tab. 10 fig. 2; see p. 44). The species is illustrated here with an animal from the Gulf of Oman almost identical to Rüppell & Leuckart’s painting.

Gymnodoris impudica Oman GS

Gymnodoris inornata (Bergh, 1880) Size: 15 mm Diagnostic characteristics: The body is uniformly orange yellow in colour with weakly raised pustules. The frontal pallial rim is marked by a ridge. The numerous gills form a complete circlet around the anal papilla and may be little darker than the body. Occurrence: Two specimen records from near Hurghada, Egypt, in shallow water; few photographic records from the Gulf of Eilat and Egypt. Geographical distribution: New Red Sea records; otherwise known from South Africa and the West Pacific, where it grows much larger, 50 mm. Species accounts 135

Suborder Doridina

Family Gymnodorididae

Gymnodoris inornata

JH

JH

Gymnodoris sp. Size: ? 35 mm Diagnostic characteristics: The body is translucent white and covered with raised rounded or conical tubercles of various sizes. These are dense white and cover the body; only around the head are some tubercles arranged to mark the margin. Even the foot and sides are tuberculate. The gills and rhinophores are both white, but the rhinophores may have an ochre tinge; the tips are opaque white. The (approximately 12) gills are arranged in a complete circlet around the anal papilla, and have white axes. Biology and ecology: This species lays a string of yellow orange eggs very similar to those of G. ceylonica (Schmid 1996). Occurrence: Three photographic records from Jeddah, Saudi Arabia, and one from the Gulf of Aqaba. Geographical distribution: A new species of Gymnodoris found throughout the Indo-West Pacific.

JK

Gymnodoris sp. JK

136 Species accounts

Suborder Doridina

Family Polyceridae

Family Polyceridae The Polyceridae form a polymorphic family. Generally the body is long and thin and the mantle is reduced. The relatively large gills retract with great difficulty but not into pockets (known as phanerobranch dorids). Anterior processes may be present near the rhinophores and near the gills, presumably as a deterrent to predators to protect the vital organs. The rhinophores are lamellate and each has a pocket into which it can retract. Species are usually brightly coloured, although some can be cryptic, and found hiding under stones. Species of the genus Nembrotha feed on ascidians but Tambja and Polycera feed on bryozoans and nudibranchs. Nembrotha megalocera Yonow, 1990 Size: 75 mm Diagnostic characteristics: The body is black with striking yellow, orange, white, and violet markings. The species receives it name from the Greek for its long rhinophores, which are black. The gills are shades of red with purple or violet bases. There is a clear patch behind each rhinophore, as in Chromodoris charlottae (p. 181).

Nembrotha megalocera

JK

Species accounts 137

Suborder Doridina

Family Polyceridae

Nembrotha megalocera

T. Metcalfe

H. Sjöholm

Feeds on the violet solitary ascidian Diazona (i.d. N. Shenkar), and swims with lateral flexions of the body when disturbed (pers. obs.). Occurrence: Common on coral reef faces in the central Red Sea; not recorded from the Gulf of Eilat. Geographical distribution: A species endemic to the Red Sea. Biology and ecology:

Tambja affinis (Eliot, 1904) GBR, Marshall & Willan 1999) The body has a black ground colour with longitudinal orange lines, quite thin, fading into wider green stripes between the rhinophores and up the gills. The edge of the foot is more yellow than the dorsal stripes. The gills and rhinophores are black. Occurrence: Uncommon on open slopes on coral reefs. Geographical distribution: Tropical Indo-West Pacific in range. Size: 85 mm (Australia, Diagnostic characteristics:

JK

Tambja affinis

138 Species accounts

DPS

Suborder Doridina

Family Polyceridae

Tambja affinis

JK

Tambja limaciformis (Eliot, 1908) Size: 12 mm Diagnostic characteristics: A small nudibranch, the long narrow body broadens at the head and tapers to the pointed tail. The lobed oral tentacles are very obvious. Tambja limaciformis is brick red in colour with scattered creamy spots approximately marking the mantle rim. The three creamy gills are located in the centre of the body. The bulbous rhinophores and gills are tipped with purple. Biology and ecology: Tambja limaciformis can be found in shallow water in coral reef flats, under or amongst coral rubble Occurrence: Originally described from the Red Sea; single specimen from the Gulf of Eilat, one specimen from the Suakin Expedition, and numerous photographic records. Geographical distribution: Known from the Red Sea and tropical Indo-Pacific.

Tambja limaciformis 10 mm

JH

11 mm

JH

Species accounts 139

Suborder Doridina

Family Polyceridae / Family Triophidae

Thecacera pacifica

H. Göthel

Thecacera pacifica (Bergh, 1883) Size: 60 mm (South Africa, Gosliner 1987) Diagnostic characteristics: This species is unmistakeable, bright orange in colour with blue tips to the partial rhinophore sheaths (“theca” means encased or sheathed in Greek) and gill processes. The blue colour can vary from light blue to turquoise. There is a black band between the blue and orange pigmentation. The rhinophores and bipinnate gills are orange with black and blue markings. Occurrence: Shallow reef top, three photographic records from the northern and central Red Sea. Geographical distribution: First records for the Red Sea; otherwise tropical Indo-West Pacific. Family Triophidae Species of Triophidae are phanerobranch dorids with a much reduced mantle skirt. The edges of the oral veil and mantle have tubercles which may be branched or papillate. There is no pocket for the gills but the lamellated rhinophores withdraw into pockets. Species feed on encrusting and arborescent bryozoans. Plocamopherus indicus Bergh, 1890 Size: 35 mm Diagnostic characteristics:

Previously unknown, this species appears variable in colour, but with orange spots around foot and mantle and scattered brown patches (similar to P. ceylonicus). There are two papillae as a single pair near the gills which terminate with a dark knob. There are three pairs of larger lateral papillae and other arborescent papillae on the body. Posterior to the gills, the tail bears a keel.

140 Species accounts

Suborder Doridina

Family Triophidae

JK

Plocamopherus indicus 35 mm

B&SK

JK

Other species of Plocamopherus have the ability to swim by lateral flexions of the body, aided by the prominent keel on the tail. Plocamopherus imperialis can emit flashes of blue light when disturbed, while the pairs of knobs on P. ceylonicus are thought to create luminescence. Occurrence: Four photographic records from Jeddah, Saudi Arabia, and Eilat, Israel. Geographical distribution: First recent record for the Red Sea; otherwise single record from the tropical Indo-West Pacific but previously considered a nomen dubium (Thompson 1975). Note: This species may have an older name, P. ceylonicus Kelaart (1868): Vayssière was able to distinguish between the two, so until we have specimens available for study, the species must remain valid (see notes on Vayssière’s plate, p. 48). Biology and ecology:

Species accounts 141

Suborder Doridina

Family Triophidae / Family Aegiridae

Plocamopherus ocellatus Rüppell and Leuckart, 1828 Size: 85 mm (Eliot 1908) Diagnostic characteristics: The body is normally chocolate brown or black with bright yellow to orange round spots (see Rüppell & Leuckart’s figure of the type on p. 40). This record is much paler, but still retains the large orange spots and the three paired conical processes on the dorsum. The frontal margin is decorated with a series of branching processes. The body appears softer and more translucent than that of P. indicus, and lacks a mantle margin. Biology and ecology: Previous authors have found the Red Sea specimens with the bryozoan Bugula neritina, on which it is presumed to feed. Eliot (1908) reported that specimens faded in colour when food was not available, and that they luminesced freely, not only when disturbed. Occurrence: Single photographic record from the central Red Sea. Geographical distribution: Originally described from the Red Sea, P. ocellatus is a Lessepsian migrant into the Mediterranean, where it appears to be more frequently encountered.

Plocamopherus ocellatus

JK

Family Aegiridae Elongated dorids which have no mantle skirt and a high body profile. They are usually knobbly and stiff. The tough leathery epidermis has sponge spicules imbedded in the body wall. The smooth rhinophores are long and tapering. All species which have been studied are known to feed on calcareous sponges. The genus Aegires has recently been synonymised with Notodoris but this is not adopted here.

142 Species accounts

Suborder Doridina

Family Aegiridae

Aegires citrinus Pruvot-Fol, 1930 Size: 15 mm Diagnostic characteristics: The hard bright yellow body is covered with pale grey patches composed of tiny dots. There are mushroom-shaped knobs all over body but especially near the mantle margin. The margin on the tail is marked by rounded tubercles. The smooth tapered rhinophores and the gills are each protected by three knobs. Occurrence: Three specimen records, found under coral heads on sand in shallow water, from the Gulf of Eilat and Quseir, Egypt. Geographical distribution: First records for the Red Sea; otherwise recorded from South Africa and the West Pacific.

6 mm

JH

Aegires citrinus 15 mm

JH

Aegires villosus Farran, 1905 Size: 10 mm Diagnostic characteristics: The body is typical of aegirids – firm, knobbly, and colourful. The knobs are club-shaped and relatively long. The rhinophores are protected by several papillae. The gills are protected by a row of branching papillae over-arching them. These papillae are the only ones which appear branched or confluent at their bases; all the others are solitary. The body is white, and the papillae are multicoloured, white, Species accounts 143

Suborder Doridina

Family Aegiridae / Family Hexabranchidae

Aegires villosus 10 mm

B&SK

orange, and lilac, mostly with deep violet tips. The very long rhinophores are translucent at the base fading rapidly into white, orange, and deep violet for the distal third. Occurrence: Single specimen record (Suakin Expedition 1980) and one photographic record from the Gulf of Eilat (June 2005). Geographical distribution: First record for the Red Sea; otherwise known only from the tropical Indo-West Pacific. Family Hexabranchidae Species of the single genus Hexabranchus have an ample dorsum which is soft and flexible, with the mantle edge rolled up when the animal is crawling. It can be unfurled to startle predators and for swimming (see figure on p. 30). The oral tentacles are large flattened lappets, and the six gills can partially retract into six separate pockets in the mantle. Recorded as feeding on a variety of invertebrates, including sponges and ascidians. Hexabranchus sanguineus (Rüppell and Leuckart, 1830) Size: 220 mm Diagnostic characteristics: Hexabranchus sanguineus is best known as the Spanish Dancer. The Red Sea species is deep red with white markings just inside the mantle edge, which is held curled up when the animal is crawling. The rhinophores are deep red and the gills are paler with white axes. Biology and ecology: This beautiful dorid swims by undulating the edges of its mantle in two waves which run posteriorly (p. 30). The flash of colours, the increase in size, and the sudden movement warn off predators. In addition, Hexabranchus swims without stimulus or disturbance: a 144 Species accounts

Suborder Doridina

Family Hexabranchidae

Hexabranchus sanguineus Swimming; note shrimps

TP

group was observed to swim together in 2 m of water (Gat & Fainzilber 1984). A commensal shrimp lives with H. sanguineus, usually found amongst the gills. Periclimenes imperator Bruce feeds on detritus adhering to the mucus on the nudibranch’s body (Schuhmacher 1973a). They are even able to feed when the Hexabranchus is swimming, a rare adaptation in cleaner shrimps. Two cleaner shrimps, one red and one white, are visible in the photograph (above). The egg mass is characteristic, a large ‘rose’ varying in colour from pink to red and measuring up to 50 mm in diameter (over). Breeding peaks between April and November, although the egg masses can be found all year round (pers. obs. and Gohar & Soliman 1963). They are conspicuous due to chemical protection: the nudibranch obtains potent chemical defences from its sponge prey, which it passes on to its egg ribbons. The macrolides isolated from Hexabranchus, its egg mass, and the sponge Halichondria are strong inhibitors of fish feeding (Pawlik et al. 1988). Occurrence: Common on shallow coral reef plateaus, 10–20 m, all year; a population explosion was recorded during the summer of 2006 in Eilat (B. & S. Koretz, pers. comm.). Species accounts 145

Suborder Doridina

Family Hexabranchidae

Pair of Periclimenes TP

Egg mass

Hexabranchus sanguineus

NY

A. Double

Geographical distribution:

Endemic to the Red Sea. Notes: There are beautiful illustrations in Rüppell & Leuckart (see p. 42) and in Ehrenberg (p. 37). Many scientists consider that there are only one or two species world-wide; however, this “colour form” is only ever seen in the Red Sea to the exclusion of the other colour forms, which overlap in the rest of the Indo-West Pacific. The Indian Ocean Hexabranchus marginatus (Quoy & Gaimard) is consistently different in coloration and pattern and has not been recorded in the Red Sea, which further supports the theory that the two are different. 146 Species accounts

Suborder Doridina

Family Hexabranchidae

Hexabranchus sp. Size: 520 mm

Diagnostic characteristics: Hexabranchus sp. has the same body form as H. sanguineus but is much larger, more than twice as large, with the dorsum covered in soft pustules. The body is pink in colour with tiny yellow rhinophores, yellow oral tentacle lappets, and some yellow patches along the mantle edge. A face-on view shows the lamellae on the yellow rhinophores, and the peculiar texture of the skin unlike that of the smaller species of Hexabranchus (over). Biology and ecology: Despite its enormous size, this species is capable of swimming when disturbed, and unfurls its mantle edge in warning. The commensal shrimp Periclimenes imperator also occurs on this species (over). However, it is bright yellow, whereas on H. sanguineus it can be red, white, and speckled red and white, but not yellow.

Hexabranchus sp.

A. Double

Species accounts 147

Suborder Doridina

Family Hexabranchidae

Note yellow Periclimenes A.

Double

Hexabranchus sp. A.

Double

Occurrence:

Abundant in its single known locality, a platform at the bottom of a cliff, 35 metres depth, in the southern Red Sea. Geographical distribution: Red Sea and western Pacific: a population of equally large specimens was found in Hong Kong and one specimen was preserved for examination. Photographs of animals from Fiji also appear identical to these in external morphology. Cryptobranch dorids This loose grouping applies to the majority of dorids, those which have a single pocket, often with an elaborate rim, into which the gills retract. If the gill pocket is decorated, often the rhinophore pockets are also. The species are listed alphabetically by family, excepting the Chromodorididae which can be found at the end (p. 169). 148 Species accounts

Suborder Doridina

Family Actinocyclidae

Family Actinocyclidae Two genera are included in this family, both recorded from the Red Sea, Hallaxa with small species and Actinocyclus with very large species. The body is very humped or domed in Actinocyclus, firm to the touch in both, often with low rounded pustules and hollows or pits to resemble sponge osculae. Most characteristic in both genera is the way the gills are held, a small clump held closed with all the tips pointing inwards and almost touching, forming a globe. Hallaxa feeds on siliceous sponges while Actinocyclus feeds on non-spiculose sponges. Actinocyclus verrucosus Ehrenberg, 1831 Size: 100 mm Diagnostic characteristics: The rubbery mantle is olive green to brown with lighter and darker mottling. The high dorsum is lumpy with white and dark specking on the mounds. There are also brown ocelli scattered on the dorsum. The small clump of greenish gills is characteristically held in a closed circlet forming a goblet. Both gills and rhinophores are proportionally smaller than usual and the same colour as the mantle, presumably to protect them from predation. Occurrence: Four photographic records from the northern and central regions of the Red Sea; these are the first records since Ehrenberg described the species from the Red Sea in 1831. Geographical distribution: Red Sea and tropical Indo-Pacific to Hawaii.

Actinocyclus verrucosus

JK

JH

Species accounts 149

Suborder Doridina

Family Actinocyclidae / Family Discodorididae

Hallaxa indecora (Bergh, 1905) Size: 15 mm Diagnostic characteristics: This is a very small species, dark purplish to brown, with prominent gills and rhinophores. The base and tip of each bulbous rhinophore is white. There can be additional white flecks on the dorsum and around the rims of the rhinophore pockets. Occurrence: Single specimen record from the northern Red Sea; several photographic records from the Gulf of Eilat. Geographical distribution: Ranging throughout the Indo-West Pacific.

Hallaxa indecora

B&SK

OL

Family Discodorididae This family is suffering from lack of recent taxonomic work: many species have been described, and numerous specimens are rarely found for comparisons of external and internal characters. Species of this family are characterized by a much flattened mantle extending beyond the foot on all sides. The dorsum is usually rough to touch due to small evenly sized tubercles containing sponge spicules. Many species autotomise parts of their mantle in defence. The rhinophores and gills are simple, and the six gills are simply pinnate. Ventrally, the markings can be important, as spots, patches, or bands of pigment. The four species described here are all distinctive; two other species recorded from the Red Sea have not been found recently (p. 58). Discodoris fragilis (Alder and Hancock, 1864) Size: 150 mm Diagnostic characteristics:

The stiff mantle can be variable in colour, white, grey, or beige, covered in somewhat symmetrically arranged darker brown, grey, or black patches of irregular shapes and sizes. Ventrally, there are also patches on the hyponotum generally concentrated towards the sole of the foot, which has smaller spots and flecks. The gills and rhinophores are greyish with white speckling. Occurrence: Specimen records from southern Egypt and Sudan; shallow water on reef flat under stones and slabs; several photographic records from the Gulf of Eilat, northern, and central Red Sea. Geographical distribution: Widespread throughout the Indo-Pacific oceans; Lessepsian migrant to the Mediterranean. 150 Species accounts

Suborder Doridina

Family Discodorididae

JK

JK

Discodoris fragilis 32 mm

JH

Species accounts 151

Suborder Doridina

Family Discodorididae

Discodoris schmeltziana Bergh, 1880 Size: 50 mm Diagnostic characteristics: The oval dorsum is covered in large tubercles. The basic ground colour is cream or beige, splashed with purple, brown, cream, and yellow patches. The rhinophores are yellowish with white tips and brown and white speckled lamellae. The gills are sparsely pinnate, translucent to pale yellow and brown in colour. Biology and ecology: Active at night (B. & S. Koretz, pers. comm.). Occurrence: Single specimen found under dead coral in lagoon, Hurghada, Egypt, and several photographic records from the Gulf of Eilat and Egypt. Geographical distribution: First records for the Red Sea; otherwise western Pacific Ocean.

B&SK

B&SK

JH

Discodoris schmeltziana

152 Species accounts

JH

Suborder Doridina

Family Discodorididae

Paradoris erythraeensis (Vayssière, 1912) Size: 50 mm Diagnostic characteristics: The body is oval and flattened, finely covered in small tubercles with a sprinkling of larger tubercles, and the margin is slightly crenulated. Colour can vary from pale grey to white, the larger tubercles may be pale yellow, and there are additional black flecks. Around the margin is an irregular, sometimes faint, band of yellow. The rhinophores are pale to dark brown, and the six bipinnate gills are translucent watery yellow, and not readily extended.

Paradoris erythraeensis 11 mm

B&SK

Biology and ecology: Found in very shallow sublittoral reef flats, under rocks and stones; the white egg ribbon is a spiral, standing on edge with the eggs visible in transverse rows (Gohar & Aboul-Ela 1959). Occurrence: Two photographic records from Eilat, Israel and Jeddah, Saudi Arabia; it was frequent at Al-Ghardaqa (Hurghada), Egypt, from February to June 1952 but rare thereafter (Gohar & Aboul-Ela 1959). Geographical distribution: Red Sea and Indo-West Pacific.

Sebadoris nubilosa (Pease, 1871) Size: 250 mm Diagnostic characteristics: The oval body is covered in soft papillae up 5 mm long, longest around the edges of the mantle. The body colour is cream with variously sized patches in shades of grey brown, the central ones more or less symmetrical about the midline (and therefore the species can be confused with Discodoris fragilis). The rhinophores and six large gills are greyish yellow with intensely white midribs. There is white speckling on the rhinophores and the distal portions of the gills. The ventral surface is mottled, concentrated in a sub-marginal band around the hyponotum. There are spots on the foot which decrease in size towards the edges. Species accounts 153

JK

Suborder Doridina

Family Discodorididae Biology and ecology: Sebadoris nubilosa is one of the few dorso-ventrally flattened dorids which can swim. The coiled egg ribbon is dense white in colour turning to reddish violet before hatching; it is laid flat rather than on its edge (Soliman 1980). Occurrence: Several photographic records, in shallow water on the reef flats; often found crawling on coral sand in reef areas. Geographical distribution: Recorded from the Red Sea, the western Indian Ocean, and the West Pacific.

B&SK

Sebadoris nubilosa Swimming

Anterior margin and rhinophores

B&SK

A. Stern

Ventral surface

154 Species accounts

A. Stern

Suborder Doridina

Family Dorididae

Family Dorididae The Dorididae is an all-embracing family of many genera and species of dorids; the taxonomy, definitions, and relationships between the numerous genera are still in need of revision. Generally, the species are oval, sometimes coarsely tuberculate, but always with a gill circlet posteriorly and lamellated rhinophores anteriorly. The rhinophores retract into pockets and the usually tripinnate gills retract into a single pocket (cryptobranchiate). Most species are cryptic, and the dull mottled colour patterns reflect this life style; when they are brightly coloured, it may be because they are cryptic on their colourful sponge prey. Many species have distinctive markings underneath, so it is always worth checking. In species which have been studied, the food item is sponge, with each opisthobranch species preferring one or few species. Aldisa sp. 1 Size: 22 mm Diagnostic characteristics:

The body is deep red with two large rimmed ‘pits’ resembling the osculae of sponges. There are numerous small spits with spiculose rims. The dorsum, the pits, and the rhinophores are red, while the six pinnate gills are whitish green, resembling a sprinkling of coral sand. Biology and ecology: Feeds on sponges and is very well camouflaged to such an extent that the pits on the dorsum resemble the exhalent osculae of the sponge ‘host’. Occurrence: First record from the Red Sea: single specimen from Seghala Island, southern Red Sea. Geographical distribution: A new species with unknown distribution: a very similar if not identical species is found in Réunion and presented below.

Aldisa sp. 1 Réunion

22 mm

NY

P. Bidgrain

Species accounts 155

Suborder Doridina

Family Dorididae

Aldisa sp. 2

B&SK

B. Picton

Aldisa sp. 2 Size: 20 mm Diagnostic characteristics:

The body is elongated oval in shape, pale or dark pink and black as in species of Phyllidiella (see p. 220). The pink areas are faintly tuberculate and the black areas low-lying and smooth. The rhinophores are black for the whole of their lengths. The gills are located far posteriorly on the mantle, and are translucent white in colour. They are rarely extended, making them look even more like a toxic phyllidiid. Occurrence: A single specimen record, and several photographic records from the northern Red Sea and the Gulf of Eilat. Geographical distribution: This is a new species which may prove to be endemic to the Red Sea. Note: This species is tentatively placed in the genus Aldisa only because several other phyllidiid mimics belong to this genus. Atagema ornata (Ehrenberg, 1831) [= Trippa intecta (Kelaart, 1858)] Size: 55 mm Diagnostic characteristics:

The humped mantle is generally reddish or reddish brown with a central ridge of tubercles in contrasting white or ‘coral sand’ colour. There are compound tubercles arranged in very discrete groups on the remainder of the dorsum resembling solitary corals. White spots are usually present on the mantle, gills, and rhinophores. The rhinophores and gills are dark except in juveniles (B. & S. Koretz, pers. comm.).

Atagema ornata

JK

156 Species accounts

B&SK

Suborder Doridina

Family Dorididae

Biology and ecology: Found in the spring in the Gulf of Eilat, and nocturnal in habit. The egg ribbon is creamy yellow, slightly undulated on the free edge. At hatching the ribbon is light pinkish violet (Gohar & Soliman 1967g). Occurrence: Several photographic records from the central Red Sea and Gulf of Eilat; in shallow reefs with sand or coral rubble. Geographical distribution: Tropical Indo-West Pacific.

Atagema spongiosa (Kelaart, 1858) [= Trippa areolata (Alder & Hancock, 1864)] Size: 140 mm Diagnostic characteristics: This large dorid is irregular and knobbly, with a very high central hump. Distinctive features are the deep black pits on the sides, visible even in the juvenile. The dorsum is variable in colour, shades of mottled creams, beiges, browns, etc. The foot is dark purple in colour. Juvenile PK

Atagema spongiosa

JK

Biology and ecology: Spawning starts in May in the northern Red Sea: the egg ribbon is white with regular undulations along the free edge but violet prior to hatching (Gohar & Soliman 1967g). Occurrence: Several photographic records from the central Red Sea, on encrusted coralline rocks or rubble of tide pools and shallow reefs. Geographical distribution: Tropical Indo-West Pacific oceans.

JK

Species accounts 157

Suborder Doridina

Family Dorididae

Hoplodoris grandiflora (Pease, 1860) Size: 55 mm Diagnostic characteristics: The mantle is vaguely oblong in shape, with the posterior end barely wider than the anterior. There are large tubercles all over the dorsum, some with white patches at the base, the majority with dusty white markings on the top. The dorsum is beige to brown with darker brown patches on the hump; the mantle skirt is broad. The gill pocket is large in comparison to H. pustulata, with six lobes, and the breadth of the gills spans almost all the posterior of the mantle when fully extended. Occurrence: Two photographic records and one specimen record from the northern Red Sea and the Gulf of Eilat. Geographical distribution: First Red Sea records; otherwise Indo-West Pacific.

Hoplodoris grandiflora 30 mm

JH

Hoplodoris pustulata Abraham, 1877 [= H. bifurcata (Baba, 1993)] Size: 30 mm Diagnostic characteristics: The mantle is pale brown to fawn or beige. It is covered in evenly sized and spaced tubercles which are often ringed with white except those in the centre of the dorsum, making the central region appear darker. There are tiny black spots present on the mantle. The rhinophores sheath is closed by two valves and the gill pocket is tuberculate. The anterior foot margin is bilaminate. 158 Species accounts

Suborder Doridina

Family Dorididae

Occurrence:

These are the first two specimen records, found under stones in the lagoon at Sharm el Sheikh, Hurghada, and Quseir, Egypt; additionally, there are numerous photographic records from Saudi Arabia, Egypt, and the Gulf of Eilat. Geographical distribution: First Red Sea records; otherwise Indo-West Pacific. Note: Well known in the Pacific Ocean as Hoplodoris nodulosa (Angas) before a recent review, it now appears to be called H. bifurcata (Baba) (syn. nov.). This is in error since H. pustulata is not a synonym of H. nodulosa (see Yonow et al. 2002).

B&SK

Ventral view, 15 mm

Hoplodoris pustulata

B&SK

JH

B&SK

15 mm

16 mm

JH

JH

Species accounts 159

Suborder Doridina

Family Dorididae

Sclerodoris apiculata (Alder and Hancock, 1864) Size: 40 mm Diagnostic characteristics: This dorid is very well camouflaged on its sponge prey and the associated invertebrate community. The body is highly domed with a central area of irregular compound tubercles. The body colour is drab beige with lighter and darker spots and patches. The gills and rhinophores are similarly coloured. There are characteristic long extensions on the tips of some of the tubercles, visible in three of the photographs. The ventral surface clearly shows the structure of the mouth, the bilaminate foot, and the long oral tentacles (far right). Occurrence: Several photographic records from Sharm el Sheikh, Egypt, and Jeddah, Saudi Arabia. Geographical distribution: Indo-West Pacific, not commonly found presumably due to its excellent cryptic colouration.

JH

PK

Sclerodoris apiculata JK

160 Species accounts

Suborder Doridina

Family Dorididae

Sclerodoris apiculata

GS

Ventral surface

JH

Sclerodoris tuberculata Eliot, 1904 Size: 25 mm Diagnostic notes: This cryptic dorid is a wonderful red or orange colour, with scattered patches resembling coral sand. There is a large central pit in the middle of the mantle in front of the gills, and several smaller ones around the sides, resembling sponge osculae. Ventrally, the species is completely red: the hyponotum, the sole of the foot, and the head and tentacles, visible under the damaged mantle (below, right). Biology and ecology: Nocturnal in habits (B. & S. Koretz, pers. comm.). Occurrence: Numerous photographic records from the Gulf of Eilat and the central Red Sea. Geographical distribution: Tropical Indo-West Pacific oceans.

JK

Sclerodoris tuberculata

JK

JK

Species accounts 161

Suborder Doridina

Family Halgerdidae

Family Halgerdidae A family not clearly defined taxonomically, comprising several genera which vary considerably in body form. Generally, a sculptured mantle covers the foot, and the rhinophore and gill pockets are protected by a series of flaps. Like most dorids, species of the family feed on sponges. Asteronotus cespitosus (van Hasselt, 1824) Size: 270 mm Diagnostic characteristics: The massive body is smooth and firm, almost gelatinous. There is a median ridge of tubercles and scattered tubercles forming the middle area of the dorsum. Rings of tubercles run parallel to the mantle edge around the median region. The mantle and foot can be brown, beige, or grey, often with white on the tubercles. The gills and rhinophores are well protected in this species: the rhinophore rim is large and fleshy, resembling the other tubercles when the rhinophores are retracted and the fleshy gill lappets close over the gills when they are retracted. The ventral view shows the characteristics of the hyponotum, foot margin, the head and long oral tentacles.

TP

Asteronotus cespitosus

Ventral view

CT

JK

JK

Biology and ecology: Spawning reaches a maximum between May and September in the northern Red Sea (Gohar & Soliman 1967e). Occurrence: Few records exist of this species; found in the very shallow sublittoral, possibly intertidal, under rocks and stones throughout the Red Sea. Geographical distribution: Tropical Indo-West Pacific oceans.

162 Species accounts

Suborder Doridina

Family Halgerdidae

Halgerda willeyi Eliot, 1904 Size: 90 mm Diagnostic characteristics: The rubbery dorsum is thrown into peaks connected by ridges. The body colour is white with yellow lines on the ridges and yellow tubercles where the ridges meet. In the ‘hollows’ are irregular dark brown to black lines and smaller ones around the mantle margin. Considerable variation appears to occur in the pigmentation of this species, but the most common is illustrated in the main photograph. These other forms occur rarely in comparison to the usual colour; it may be that with such an abundant and successful species, aberrations and variations are more likely to occur. Biology and ecology: Often found in groups of three or more, Halgerda willeyi seems to prefer shaded areas under overhanging corals and walls. Occurrence: Very common, sublittoral to 20 m depth, on flat areas of caves and coral slopes, in amongst and on the coral rubble accumulated there; more common in the southern Red Sea. Geographical distribution: Indo-West Pacific oceans.

JK

TP

TP

Halgerda willeyi

JK

JK

Species accounts 163

Suborder Doridina

Family Kentrodorididae

Family Kentrodorididae A polymorphic family, the name is based on the genus Kentrodoris which is atypical not only of dorids but of the family as well: body form in the single species Kentrodoris rubescens (Bergh) is elongate, gelatinous, and very soft; it has not been recorded from the Red Sea. Jorunna is more typically dorid in shape, generally oval and humped, white, beige, or yellow with spots and patches in shades of brown and black. The mantle is spiculose in Jorunna and therefore rough to touch. Gills are usually tri-pinnate. Members of the family feed on sponges.

OL

Jorunna funebris (Kelaart, 1858) Size: 70 mm Diagnostic characteristics: The oval elongate body is white with round or oval patches containing black papillae; these are spiculose, giving a spiky appearance to the animal. The pattern and density of black spicules is variable. The animal is stiff and rough to touch. Rhinophores are black. Occurrence: A common species found in the shallow subtidal, under rocks and boulders; several specimen and photographic records from the northern and central Red Sea and the Gulf of Eilat. Geographical distribution: Widely distributed in the tropical Indo-West Pacific.

JK

Jorunna funebris 35 mm

NY

164 Species accounts

Suborder Doridina

Family Kentrodorididae

Jorunna funebris

JK

Jorunna pantherina (Angas, 1864) Size: 25 mm Diagnostic characteristics: The mantle is evenly spiculose, translucent beige or pinkish brown in colour with darker patches. There are many extended spicules on the ends of the tubercles, which are ‘glistening’ white. The gills are located far back, held in a rather closed tulip-shape. The ochre rhinophores are tipped in white. Occurrence: Single specimen record from Quseir, Egypt, and few photographic records from Eilat, Israel. Geographical distribution: First Red Sea record; otherwise, described from NSW, Australia only.

Jorunna pantherina

JH

Species accounts 165

Suborder Doridina

Family Kentrodorididae

Jorunna sp. Size: ? 20 mm Diagnostic characteristics:

JK

The elongate body bears scattered isolated black or dark brown spicules evenly distributed on the mantle. The body may be coloured white or yellow; when white, the mantle bears small patches of yellow brown. Consistent in the photographs examined are the isolated black spicules and the dark patches at the tip of the foot, which extends beyond the mantle and has additional black spots around its sides. The gills are more numerous and more dense than in J. funebris. The rhinophores are dense black for at least the distal half of the lamellate portion but this may be a function of size. Occurrence: Single specimen record and several photographic records from the central and northern Red Sea and Gulf of Eilat. Geographical distribution: Red Sea; photographs from Réunion and South Africa indicate its presence in the western Indian Ocean. This is probably a new species.

18 mm

JH

Jorunna sp. JK

OL

166 Species accounts

Suborder Doridina

Family Platydorididae

Family Platydorididae Very flattened oval dorids with the mantle extending far beyond the foot all around. The mantle is rough to the touch due to the sponge spicules incorporated in the dorsum. The gills are foliaceous, retractile into a distinctive pocket with a scalloped edge. Platydoris scabra (Cuvier, 1804) Size: 100 mm Diagnostic characteristics: The mantle is yellowish brown with darker brown patches arranged more or less symmetrically either side of the midline. There is an orange edge to the mantle and the foot. The rhinophore sheaths and gill pocket are also marked with an orange line. Biology and ecology: Breeds from May to September, laying an orange yellow ribbon in a spiral approximately 100 mm across (Soliman 1978). Occurrence: Platydoris scabra can be found attached to the under surface of rocks on reef platforms; it should be common but has been rarely recorded. These photographs are from Jeddah, Saudi Arabia. Geographical distribution: Indo-West Pacific.

JK

JK

Platydoris scabra

PK

Species accounts 167

Suborder Doridina

Family Platydorididae

Platydoris striata (Kelaart, 1858) Size: 100 mm Diagnostic characteristics: The body is white with very fine undulating brown lines. The rhinophores and gills are covered in fine brown lines and can be very dark brown. The sheaths of the gills and rhinophores are raised and bear brown lines on the outer surfaces; the inside of the sheaths and the ventral surfaces are dense white. Occurrence: First specimen record and several photographic records from Egypt and Saudi Arabia. Geographical distribution: First record of Platydoris striata for the Red Sea; otherwise recorded only from the western Indian Ocean. Note: A similar West Pacific Ocean species, Platydoris cruenta (Quoy & Gaimard), has blotches of orange red on the dorsum and hyponotum in addition to the thin undulating lines. They have been synonymised by some authors.

JK

Platydoris striata

JK

168 Species accounts

A. Valdés

Suborder Doridina

Family Chromodorididae

Family Chromodorididae The chromodorids are a large family with numerous genera which all feed on sponges, from which they acquire their toxins. These are stored in mantle glands located around the mantle, sometimes in special projections in some genera. The rhinophores and gill circlet are often a contrasting colour to the mantle, which is usually brightly marked with various colours and patterns. Chromodorids such as Chromodoris and Hypselodoris have a soft, smooth, elongated body with the foot projecting beyond the mantle posteriorly. In Cadlinella, the mantle is firm and spiculose with club-shaped or elongated tubercles. The body is high and firm and the mantle is greatly reduced in Ceratosoma, but produced into rigid projections beside and behind the gills containing the defensive glands. In the genera Glossodoris, Ardeadoris, and Thorunna the edges of the mantle are thrown into permanent folds and the mantle profile is high. The body is firm and broad in Miamira and Orodoris but has ridges extending to the edges, producing a scalloped effect.

B&SK

Ardeadoris egretta Rudman, 1984 Size: 15 mm Diagnostic characteristics:

The mantle is generous with many undulations and one permanent fold. The body is white with a yellow band at the margin. The foot is white. The rhinophores are disproportionately large, translucent white, and the white gills are tiny and digitiform. Occurrence: Three photographic records from Eilat, Israel. Geographical distribution: First record for the Red Sea; otherwise, tropical Indo-West Pacific where it grows to 50 mm.

Ardeadoris egretta

B&SK

B&SK

Species accounts 169

Suborder Doridina

Family Chromodorididae

Glossodoris electra Rudman, 1990 Size: 20 mm Diagnostic characteristics: The body is semi-traslucent white with a yellow margin and the foot is also white. There is at least one permanent undulation midway between the gills and rhinophores. The latter are orange or tipped with orange, and with white lines along their axes. The gills are held open, and spiral in at each end, typical of the genus. Occurrence: Several photographic records, all from the Gulf of Eilat. Geographical distribution: New record for the Red Sea; known from New Caledonia and the Great Barrier Reef, Australia. .

OL

OL

Glossodoris electra

170 Species accounts

OL

Suborder Doridina

Family Chromodorididae

Thorunna africana Rudman, 1984 Size: 15 mm Diagnostic characteristics: The body profile is high, and the reduced mantle is thrown into three permanent undulations. It is opaque white with a yellow margin. The tail is also white with a yellow margin and extends far beyond the mantle. The simply pinnate gills and rhinophores have red midribs and may have red edges to the lamellae. Occurrence: Several photographic records from the Gulf of Eilat only. Geographical distribution: Recorded only from the northern Red Sea and East Africa.

OL

Thorunna africana

OL

Species accounts 171

Suborder Doridina

Family Chromodorididae

Cadlinella ornatissima (Risbec, 1928) Size: 20 mm Diagnostic characteristics: The body is deep orange yellow in colour with very long white rhinophores; the rhinophores have large obvious lamellae, approximately 16 or 17 in number. The dorsum is covered in slightly pointed club-shaped tubercles which are white with magenta tips. The mantle glands are located around the edge of the mantle and are clearly visible as glistening pale yellow patches. Elongated white spots form a ring around the mantle margin inside the band of mantle glands. The five to seven simple gills are white. Occurrence: Several photographic records from the Gulf of Eilat. Geographical distribution: First record for the Red Sea; otherwise, tropical Indo-West Pacific.

B&SK

OL

Cadlinella ornatissima

172 Species accounts

B&SK

Suborder Doridina

Family Chromodorididae

Cadlinella sp. Size: ? 20 mm Diagnostic characteristics:

The translucent white body bears elongated tapering papillae, most of which have a distal red band; the actual tips of these papillae are white. The base of each papilla lies in a somewhat hexagonal yellow line. There is a band of complex mantle glands around the margin with a denser concentration on the sides of the animal, visible as glistening pale yellow patches. The rhinophores are long and white; the the gills are translucent white, and protected by a ring of small papillae. Biology and ecology: The three animals are probably together because they may be feeding on the sponge. Occurrence: Three photographic records from Jeddah, central Red Sea, and the Gulf of Aden. Geographical distribution: A distinctive new species presently known only from the Red Sea and Gulf of Aden.

Cadlinella sp.

JK

H. Schmid

Species accounts 173

Suborder Doridina

Family Chromodorididae

Chromodoris africana Eliot, 1904 Size: 90 mm Diagnostic characteristics: The ground colour is black, with two white stripes down the centre – which may be thick or thin – surrounded by a white line followed by a broad orange edge. The gills and rhinophores are also orange, sometimes deeper in colour than the mantle. This species should not be confused with C. quadricolor, which has two blue bands, three black ones, and a white edge. Chromodoris africana tends to be larger and have a more irregular margin. Biology and ecology: The breeding season begins in March lasting until September in the northern Red Sea (Gohar & Aboul-Ela 1959). Occurrence: Found in the sub-littoral to 15 m on coral reefs; uncommon in the northern half of the Red Sea, but not recorded from the southern part. Geographical distribution: Red Sea and western Indian Ocean.

Chromodoris africana

PK

On the sponge Latrunculia OL

174 Species accounts

65 mm

NY

Suborder Doridina

Family Chromodorididae

PK

65 mm

Chromodoris africana

NY

JK

Pseudoceros sp.

Species accounts 175

JK

Suborder Doridina

Family Chromodorididae

Chromodoris quadricolor (Rüppell and Leuckart, 1830) Size: 70 mm but usually smaller Diagnostic characteristics: The three longitudinal black and two blue bands on the dorsum are characteristic, although there may be some extra black marks behind the rhinophores, for example. This central striped region is followed by a thick or thin white line, a wider orange (or yellow orange) band, and white at the very edge. The gills and rhinophores can be the same colour as the marginal band or deeper orange.

On the sponge Cliona, 17 mm

NY

Chromodoris quadricolor In front of the soft octocoral Xenia biseriata

176 Species accounts

On Latrunculia JH

JK

Suborder Doridina

Chromodoris quadricolor

Family Chromodorididae

G. Bemert

JH

Biology and ecology: Chromodoris quadricolor is most often found on the branching sponges Latrunculia corticata and Negombata magnifica, and the encrusting sponge Cliona vastifica, all red or reddish brown, and common in the Red Sea. Occurrence: Very common, known as the “pyjama nudibranch”, on coral reef tops and shallow sublittoral coral reefs and slopes to 20 m; it has been observed at depths of 70 m in the Gulf of Eilat (S. Einbinder, pers. comm.). It is more common in the northern Red Sea. Geographical distribution: Red Sea and Tanzania only.

Chromodoris strigata Rudman, 1987 Size: 50 mm Diagnostic characteristics: This species is similar to Chromodoris quadricolor, but with extra black lines extending behind the rhinophores: all five black lines may be broken. There are five translucent but dark patches on the dorsum: a pair behind the rhinophores, a pair in front of the gills, and a broader patch in the centre. These patches may appear blue. Additionally, the gills appear to be orange but actually have white lamellae. Biology and ecology: Chromodoris strigata is fairly common in the spring in the Gulf of Eilat (B. & S. Koretz, pers. comm.) and can be found in the daytime.

Chromodoris strigata

B&SK

B&SK

Species accounts 177

Suborder Doridina

Chromodoris strigata

Family Chromodorididae

B&SK

B&SK

Occurrence:

Moderately common, generally occurring in deeper waters than Chromodoris quadricolor; only in the northern Red Sea and Gulf of Eilat. Geographical distribution: First record for the Red Sea; more variable in colour across the Indo-West Pacific. Chromodoris annulata Eliot, 1904 Size: 57 mm Diagnostic characteristics:

The body is white with large, raised, orange yellow pustules and two purple rings enclosing the rhinophores and gills. These orange yellow spots are also present on the foot and can be variable in number. The margin is purple, the gills and rhinophores are white edged in purple. A juvenile specimen was identical to the adults but with fewer gills, the number of which appears to increase with size.

Chromodoris annulata

OL

178 Species accounts

Suborder Doridina

Family Chromodorididae

JK

Chromodoris annulata

Ventral view, 40 mm

GS

JH

T. Metcalfe

Biology and ecology: Like a number of chromodorids, C. annulata has a habit of flapping the front of its mantle, showing the deep purple crescent on the underside, although this is not a constant feature. Breeding begins in July on the Egyptian coastline (Gohar & Aboul-Ela 1957e). A 3000 square meter area south of Quseir, Egypt, sampled ten times in eight days revealed a total of 43 individuals: Lüttman et al. (2006) concluded that individual movement ranged from 1–20 meters per day. Occurrence: Not uncommon in the Red Sea: adults are generally found on current-swept cliff faces to 10 m deep, juveniles on the shore if it consists of coral sand and rubble with rocks and boulders. Geographical distribution: Found only in the Red Sea, Arabian Gulf, and western Indian Ocean.

Chromodoris aspersa (Gould, 1852) Size: 50 mm Diagnostic characteristics: The body is pale yellow orange with a deeper orange edge. There are scattered purple spots on slightly raised pustules. The yellow gills and rhinophores are somewhat translucent or watery in colour, as are the prominent oral tentacles on the head underneath. There are purple spots and an orange margin on the foot. Biology and ecology: Breeding occurs from May to November on the Egyptian coastline (Gohar & Soliman 1967b). Species accounts 179

Suborder Doridina

Ventral view

Family Chromodorididae

CT

JK

Chromodoris aspersa

JK

JH

Occurrence:

Moderately common, hiding under and among coral blocks and boulders on reef tops, in very shallow water. Geographical distribution: Indo-West Pacific; however, this form differs from the Red Sea material in having smaller, almost smooth, spots which ‘bleed’ into the white mantle to form a halo. Chromodoris charlottae (Schrödl, 1999) Size: 82 mm (Eliot 1911) Diagnostic characteristics: The

broad mantle is bright orange with a central network of brown reticulations. There are translucent patches behind the rhinophores. The margin is banded by orange, light blue and deep blue, followed by white on the edge. Ventrally the animal is pure white. A juvenile can be discerned by the lack of mantle undulations, although colour pigmentation is well developed (upper right). Occurrence: Common, numerous photographic records in recent years. Geographical distribution: Chromodoris charlottae is endemic to the Red Sea. Eliot (1911) described a nudibranch from shallow water, 1 metre depth, on sand with sea grasses in Dongonab harbour, Sudan, which is identical to the species illustrated here; he did not recognise it as new and called it Chromodoris ? inopinata Bergh, a species belonging to the C. tinctoria colour group (see p. 184). 180 Species accounts

Suborder Doridina

Family Chromodorididae

B&SK

Three individuals

B&SK

Juvenile

JK

Sub-adult

Chromodoris charlottae

PK

JK

H. Schmid

Species accounts 181

Suborder Doridina

8 mm

Family Chromodorididae

JH

Chromodoris fidelis (Kelaart, 1858) Size: 22 mm Diagnostic characteristics: The creamy dorsum has an undulating white line bordered by magenta crescents around the edge; it has a broad orange margin. The rhinophores are black, as are the gills, and the foot is white. The juvenile in the photograph is only 8 mm and already shows the adult characteristics (left). Occurrence: Uncommon, on well colonized coral reefs which are fairly exposed. Geographical distribution: Indo-West Pacific with several colour forms.

J. Walker

JK

Chromodoris fidelis

JK

182 Species accounts

OL

Suborder Doridina

Family Chromodorididae

OL

Chromodoris geminus

GS

JK

Chromodoris geminus Rudman, 1987 Size: 55 mm Diagnostic characteristics: Chromodoris geminus is almost luminescent under water, with a broadly oval creamy yellow orange mantle with dark violet spots surrounded by bright lilac rings. These spots are also present on the foot, which has a glistening white line along the edge. The mantle is bordered by yellow, cream, and grey bands with a white line along the margin. Occurrence: Found on coral reefs to 15 m depth, not uncommon throughout the Red Sea. Geographical distribution: Red Sea and Indian Ocean. Species accounts 183

G. Bemert

Suborder Doridina

Family Chromodorididae

Chromodoris obsoleta (Rüppell and Leuckart, 1830) Size: 50 mm Diagnostic characteristics: The animal has an irregular white dorsum with fawn reticulations, a thick purple border, and a thin orange margin. The margins of the mantle are undulated in larger specimens. The relatively large rhinophores are orange with the lamellae edged in white, as are the simply pinnate gills. A juvenile only 14 mm long, found in 6 m of water on a platform of coral rubble and sand, was identical to the adults (pers. obs.). Occurrence: Fairly common on coral reefs; specimen record (Suakin Expedition) and several photographic records from the whole of the Red Sea. Geographical distribution: Endemic to the Red Sea; this is probably the species described as new by Pruvot-Fol (1933; see p. 50).

Pseudoceros sp.

JK

JK

Chromodoris obsoleta

JK

GS

Chromodoris tinctoria (Rüppell and Leuckart, 1830) Size: 80 mm Diagnostic characteristics: The broad white mantle bears a red reticulate pattern which stops just short of the margin, where it presents as a series of red spots in a white area. There is a bright yellow line around the margin of the mantle. The mantle bears small pustules. The foot is white with red spots on its upper surface and without a yellow margin. The gills and rhinophores are red and white. 184 Species accounts

Suborder Doridina

Family Chromodorididae

Occurrence:

Not uncommon in the Gulf of Eilat; two photographic records from the Egyptian coastline. Geographical distribution: Red Sea, Gulf of Oman, and western Indian Ocean. Note: The similar species C. inopinata Bergh, C. marindica Yonow, and C. reticulata (Quoy & Gaimard) all exist in the Indo-Pacific regions.

Pseudoceros cf. irretitus JK

OL

Chromodoris tinctoria

OL

Oman

Chromodoris verrieri (Crosse, 1875) Size: 25 mm Diagnostic characteristics: This is one of the few Red Sea chromodorids whose dorsum is pustulose and irregular. It is creamy with opaque speckling and small patches of yellow orange. It is enclosed by two thin marginal bands, the inner of orange yellow, and the outer of deep red. The inner yellow orange band may be broken up into patches. The foot is uniformly dense white. The rhinophores and gills have a red core with white edges to the lamellae. The approximately ten gills are held in a cup-shape as in Hallaxa (p. 150) and Thorunna (p. 171). Occurrence: Occurring in the shallow subtidal, amongst coral rubble and boulders; numerous photographic records from the Gulf of Eilat. A similar species has been recorded from Jeddah, Saudi Arabia (right). Geographical distribution: Indo-West Pacific.

Chromodoris verrieri

A. Stern

Species accounts 185

GS

Chromodoris cf. verrieri JK

OL

Suborder Doridina

Family Chromodorididae

Chromodoris sp. Size: 15 mm Diagnostic characteristics:

The translucent white or pale orange body has tiny scattered magenta spots and wavy white lines meandering across it; they stop short of the edge, producing a white band. The margin is orange with white patches. The tail is translucent with a white streak down the middle. The gills and rhinophores are translucent orange with white markings near the ends, and the very tips of some of the gills are orange. Occurrence: Shallow water on reef tops with rubble and sand, several specimen records and numerous photographs from the northern Red Sea. Geographical distribution: Endemic to the Red Sea, almost certainly a new species; Chromodoris decora (Pease) is a sympatric species found in the Pacific Ocean.

GS

PK

Chromodoris sp.

OL

JK

186 Species accounts

Suborder Doridina

Family Chromodorididae

Durvilledoris lemniscata (Quoy and Gaimard, 1832) Size: 15 mm Diagnostic characteristics: The translucent lilac body bears three thick creamy or white stripes with a red line in-between them. There is a white margin with a lilac submarginal band, which deepens in colour anteriorly and posteriorly. The foot is pale violet with a darker margin. The gills and rhinophores are translucent or white basally, then reddish orange, and the ends are a deep purple. Occurrence: Occurs in very shallow water, amongst coral rubble; few photographic records from the central Red Sea. Geographical distribution: Tropical Indo-West Pacific.

PK

JK

Durvilledoris lemniscata

JK

Species accounts 187

GS

Suborder Doridina

Family Chromodorididae

Durvilledoris pusilla (Bergh, 1874) Size: 10 mm Diagnostic characteristics: The dorsum of this small species has a broad creamy margin enclosing a complicated central area of pink with violet or magenta edging. This broadens anteriorly and posteriorly, and in two places along the dorsum on each side, behind the rhinophores and before the gills. At these wider places, there is a dense white patch centrally. The foot is pink fading into violet at the edge. The gills and rhinophores are watery orange in colour, the rhinophores with a central white core and the gills with white lamellae. Occurrence: Found sub-tidally on coral reefs; three photographic records from the central and northern Red Sea. Geographical distribution: Tropical Indo-West Pacific.

JK

Durvilledoris pusilla

OL

JK

Glossodoris cincta (Bergh, 1888) Size: 70 mm Diagnostic characteristics: The broad dorsum is high in profile with many permanent convolutions of the margin. It is reddish or brownish with creamy spots and mottling, with a distinctive tri-coloured margin on both surfaces: white, blue green, and ochre. The rhinophores are dark with white edges to the lamellae. Several colour forms exist in the Indo-West Pacific, of which two are found in the Red Sea. Biology and ecology: Egg masses can be found between May and September (Gohar & Soliman 1967d). Occurrence: Moderately common underneath coral and stones on rock and coral reefs; numerous photographic records. Geographical distribution: Widespread throughout the Indo-West Pacific. 188 Species accounts

Suborder Doridina

Family Chromodorididae

Typical

JK

JK

JH

Glossodoris cincta

JH

Species accounts 189

JK

Suborder Doridina

Family Chromodorididae

Glossodoris hikuerensis (Pruvot-Fol, 1954) Size: 60 mm Diagnostic characteristics: This is a large chromodorid with a high body profile and many permanent undulations in the mantle which increase with size. The gills are numerous and arranged in an arc which spirals inwards at each end. The dorsum is a muddy reddish brown colour with reticulations fading towards the margin, which is banded with cream or white, grey brown, and white. The foot is coloured like the mantle, sometimes a little paler. Occurrence: An uncommon species, several photographic records. Geographical distribution: Found thoughout the Indo-West Pacific.

JH

Glossodoris hikuerensis

JK

190 Species accounts

JK

Suborder Doridina

Family Chromodorididae

10 mm

JH

JK

JK

JK

Glossodoris pallida (Rüppell and Leuckart, 1830) Size: 15 mm Diagnostic characteristics: The body is creamy white or yellow with two white semi-lunar markings on the dorsum, sometimes connected by white patches along the midline. There is a thickened yellow margin, and the mantle has at least one permanent fold; the number of folds increases with size. The foot is also creamy yellow with a yellow margin; note the displaced gills in the lowest individual of the group. Occurrence: Uncommon, found in shallow waters; several specimen records (including Suakin Expedition) and several photographic records. Geographical distribution: Indo-West Pacific ocean. Species accounts 191

Glossodoris pallida

PK

Suborder Doridina

Family Chromodorididae

Glossodoris symmetricus Rudman, 1990 Size: 25 mm Diagnostic characteristics: The mantle is semi-translucent, beige or cream with opaque white markings meandering along the central region. The margin is very rumpled, with an opaque white band and an orange red edge which contains denser spots. The foot is beige or cream like the mantle, fading into a white edge. The rhinophores and gills are orange red. Occurrence: Two photographic records from the Gulf of Eilat and the Egyptian coastline. Geographical distribution: New record for the Red Sea; most records are from the Indian Ocean but G. symmetricus is also recorded from the western Pacific (Yonow et al. 2002).

Glossodoris symmetricus

B&SK

B&SK

Hypselodoris dollfusi (Pruvot-Fol, 1933) Size: 40 mm Diagnostic characteristics: This chromodorid is startling in its colour pattern. The body is creamy white with bright magenta pink rings surrounding lilac spots on the dorsum. These rings are also present around the margins of the rhinophore and gill openings. The edge of the mantle

R. Cohen

Hypselodoris dollfusi Eilat

R. Cohen

192 Species accounts

Suborder Doridina

Family Chromodorididae

Hypselodoris dollfusi Suez

M. Hassan

is bright yellow or yellow orange. The white foot is similarly marked, but without a coloured margin. The gills and rhinophores are orange red, sometimes quite dark, with a darker red line up the midribs. Occurrence: Several photographic records from the Gulfs of Eilat and Suez. Geographical distribution: Gulf of Eilat, Gulf of Suez, and Gulf of Oman. Hypselodoris infucata (Rüppell and Leuckart, 1830) Size: 70 mm Diagnostic characteristics: The body is long and thin, white with an assortment of differently sized black and yellow (or orange) spots. There are also blue grey areas arranged around the sides, generally triangular in shape. The rhinophores are bright orange red with a white core and the gills are white with red midribs and sometimes a tinge of red on the lamellae. The juvenile animal was 25 mm, found on a sandy substrate at 5 m depth. It could be mistaken for a species of Chromodoris from its body shape, but comparison with adult specimens clearly shows that they belong to the same species (over, far right). Occurrence: Relatively common on wrecks, in harbours, and other locations related to human habitation e.g. marine laboratories; often on coral sand; numerous records. Geographical distribution: Indo-West Pacific; has migrated from the Red Sea into the Mediterranean. Species accounts 193

B&SK

Hypselodoris infucata

JK

Suborder Doridina

Family Chromodorididae

A. Stern

B&SK

JH

Hypselodoris infucata

Juvenile, 25 mm

JH

TP

194 Species accounts

Suborder Doridina

Family Chromodorididae

Hypselodoris maculosa (Pease, 1871) Size: 25 mm Diagnostic characteristics: The translucent body is blotched with red spots and patches arranged in bands. Usually there are orange patches around the edges. There are longitudinal white lines (also on the tail) with smaller pink red spots in between. The rhinophores are striped with three white and two orange bands. Some colour variations occur (bottom row). Occurrence: In fine coral sand and coral rubble; uncommon, records of juveniles especially from the northern Red Sea. Geographical distribution: Indo-West Pacific ocean.

A. Stern

Adult DPS

OL

A. Stern

Hypselodoris maculosa B&SK

CT

Species accounts 195

Head CT

Suborder Doridina

Family Chromodorididae

Hypselodoris maridadilus Rudman, 1977 Size: 35 mm Diagnostic characteristics: The elongated body is white, cream, or pale yellow with five magenta stripes and a magenta border. The head is spatulate and edged in magenta which thickens at the anterior margin. The rhinophores and gills are orange. The distinctive spatulate head is common of this colour group of hypselodorids. Biology and ecology: Hypselodoris maridadilus is often found in pairs, both in the Red Sea and Indian Ocean (pers. obs.). Occurrence: Moderately common, shallow water (10 m) in coral rubble; many photographic records. Geographical distribution: Indo-West Pacific ocean.

JH

10 mm

Hypselodoris maridadilus

JK

196 Species accounts

NY

JK

Suborder Doridina

Family Chromodorididae

Hypselodoris nigrostriata (Eliot, 1904) Size: 30 mm (Chagos, Yonow et al. 2002) Diagnostic characteristics: The body shape is typical for hypselodorids, long and thin with a spatulate head. The ground colour is creamy to pale blue with longitudinal, diagonal, irregular, deep violet lines. These are also present on the foot, and both the foot and the mantle additionally have bright yellow spots. The mantle margin is yellow, especially anteriorly; this band may be broken up. The gills and rhinophores are red orange and white. Occurrence: Recorded only once by the Suakin Expedition (1980, unpublished); the second photograph was taken in the Gulf of Oman. Geographical distribution: Red Sea and western Indian Ocean (Yonow et al. 2002). Vayssière (1912) described and illustrated Chromodoris tenui-linearis (pl. 1 fig. 13, p. 47), a species described by Farran (1905) from Sri Lanka; both records are of Eliot’s species.

Hypselodoris nigrostriata Oman

GS

CT

Head CT

Mexichromis katalexis Yonow, 2001 Size: approximately 30 mm Diagnostic characteristics: The

three photographs depict a creamy white body with compound tubercles terminating in a pointed purple tip. The margin bears patches of burnt orange, and the rhinophores and gills are white basally and violet distally. Ventrally, the foot is edged in violet until the foot corners; the head and anterior margin are white but the oral tentacles are tipped in violet.

Mexichromis katalexis

HS

Species accounts 197

Suborder Doridina

Family Chromodorididae

30 mm

B&SK

Mexichromis katalexis 13 mm

CT

Occurrence: Four photographic records from the Gulf of Eilat and Geographical distribution: First record from the Red Sea; otherwise

Egypt. West Pacific. Additionally, photographs from Réunion constitute the first records for the Indian Ocean (P. Bidgrain). Notes: This species is similar to M. multituberculata Baba described from the western Pacific with simpler tubercles and a purple margin. Specimens recorded by Rudman (1983) as Mexichromis mariei from the Red Sea (13 mm) and Indian Ocean (13–17 mm preserved) are juveniles. Noumea alboannulata Rudman, 1986 Size: 11 mm Diagnostic characteristics:

Noumea alboannulata

CT

The body is translucent to deep pink with a distinctive opaque white line extending from between the rhinophores and splitting to form two lines down the dorsum and joining up behind the gills. The margin of the mantle is cream, and inside this is a line of magenta spots. The gills and rhinophores are pale red orange. Occurrence: Single specimen from Sudan (Suakin Expedition 1980), no other records. Geographical distribution: Red Sea and Coral Sea, West Pacific. 198 Species accounts

Suborder Doridina

Family Chromodorididae

Noumea flava (Eliot, 1904) Size: 15 mm Diagnostic characteristics: The whole animal is a beautiful lemon yellow: the mantle, the foot, the rhinophores, and the gills. The edge of the mantle, but not the foot, has a bright red line which thickens intermittently. The mantle is constricted at the centre of the body to form an hour-glass shape. It is reduced in this species, high in profile, and the foot is large. Occurrence: Single photographic record from Eilat, Israel. Geographical distribution: First record for the Red Sea; otherwise recorded from the western Indian Ocean to the western Pacific.

Noumea flava 15 mm

OL

15 mm

OL

Noumea norba Marcus and Marcus, 1970 Size: 10 mm Diagnostic characteristics: This tiny chromodorid is translucent pinkish orange, with an opaque white mantle margin lined by large, diffuse magenta spots. An opaque white line extends from between the rhinophores along the dorsum and splits around the gill pocket to join behind. The gills and rhinophores are a watery orange red. Occurrence: No further records since Yonow 1989 (as Noumea purpurea Baba). Geographical distribution: Tropical West Pacific and photographic record from Réunion (P. Bidgrain).

Noumea norba 10 mm

NY

10 mm

Species accounts 199

J. Dafni

Suborder Doridina

Family Chromodorididae

JH

Noumea simplex (Pease, 1871) Size: ? 15 mm Diagnostic characteristics: This small chromodorid is pale pink fading into diffuse white towards the margin. The edge is a continuous bright orange line in three individuals, and a broken orange line in the fourth. The foot is white. The rhinophore stalk and the lower half of the lamellate portion are translucent with a white core; the distal end is tipped with orange. The fewer than ten gills are also translucent white and some have orange tips. Occurrence: Four photographic records from the central and northern Red Sea. Geographical distribution: First record for the Red Sea (not recorded from the Indian Ocean); otherwise tropical western and central Pacific Ocean.

Noumea simplex

JK

Noumea sudanica Rudman, 1985 Size: 15 mm Diagnostic characteristics:

The mantle is a rather odd shape, broad at the head region, with an irregular margin, and narrowing irregularly to the tail. The body is white with a thin yellow margin and small scattered yellow patches. The foot is white with no coloured margin. The rhinophores and gills are translucent, and the rhinophore lamellae are edged in white.

200 Species accounts

Suborder Doridina

Family Chromodorididae

DPS

Noumea sudanica

DPS

GS

Occurrence:

Single holotype specimen from Sudan (Suakin Expedition 1980); two photographic records from the northern and central areas of the Red Sea. Geographical distribution: Endemic to the Red Sea. Risbecia pulchella (Rüppell and Leuckart, 1830) Size: 130 mm Diagnostic characteristics:

The colour pattern varies tremendously in this species, but animals are always cream with orange yellow spots, a purple margin, and a spatulate head. There are many small spots, which may be raised on pustules. Faint purple patches can extend from the margin towards the centre of the dorsum, and the numerous gills are arranged in an arc with spiralling ends. The foot is spotted and has a faint purple margin. The rhinophores are dark purple with white axes and dots on the edges of the lamellae, while the pinnate gills are white with strong purple lines on two sides. Species accounts 201

Risbecia pulchella R. Cohen Note shrimp

Suborder Doridina

Family Chromodorididae

JH

OL

Risbecia pulchella

JK

202 Species accounts

OL

Suborder Doridina

Family Chromodorididae

Risbecia pulchella

OL

Biology and ecology: Breeding was recorded in March and April in Hurghada by Gohar & Aboul-Ela (1957c). The trailing behaviour illustrated is very characteristic for species of Risbecia (upper left). This is the first record of the cleaner shrimp Periclimenes living with Risbecia pulchella (p. 201). Occurrence: Common on coral reefs, most frequently between May and September in the Gulf of Eilat (B. & S. Koretz, pers. comm.). Geographical distribution: Red Sea and western Indian Ocean; Rüppell & Leuckart’s engraving is reproduced on p. 43 (Tab. 9 fig. 5). Notes: The identity of this species is confused with Risbecia ghardaqana (Gohar & Aboul-Ela). Risbecia ghardaqana has fewer larger spots and a more Chromodoris-like shape. There is a row of orange spots ventrally where the hyponotum meets the foot. The foot is white with only the tip pigmented with purple (right).

Ceratosoma tenue Abraham, 1876 Size: 96 mm preserved (White 1951) Diagnostic characteristics: The body is stiff

and elongated with a reduced mantle margin marked by violet spots and dashes. There is a horn-like process over-hanging the gills from behind, hence the Greek generic name “horn body”, and the mantle bears two pairs of small lobes in front of the gills. Colour is variable, but in the Red Sea appears to be very similar to C. trilobatum, cream with violet margins. There are pale brown patches on the body, and bright orange spots. The foot bears purple around its margin, but the gill and rhinophores pockets do not (in contrast to C. trilobatum). The rhinophores stalks are translucent white, and the lamellate portion vivid orange with a violet tip. Occurrence: Single specimen record (Suakin Expedition 1980, Rudman 1988). Geographical distribution: Red Sea and Indo-West Pacific. Species accounts 203

Risbecia ghardaqana 30 mm

CT

Suborder Doridina

Family Chromodorididae

Ceratosoma tenue

CT

Ventral view

CT

Ceratosoma trilobatum (Gray, 1827) Size: 200 mm Diagnostic characteristics: The elongated body is stiff with a curved projection behind the gills and a large one on either side of the gills. The mantle edge is reduced to only a ridge around the rest of the body, but coloured by a continuous purple line which breaks up at the ‘neck’. The animal is blotchy red with bright yellow spots and a violet edge around the gill and rhinophore pockets. The horn is spotted with larger red spots on both sides, as are the lateral projections. The foot has purple patches around its margin. Biology and ecology: Found exposed on hard sea bed floors and coral reefs. Occurrence: Specimens from the Gulf of Aden; several photographic records; not uncommon in the Gulf of Eilat in the spring. Geographical distribution: New records for the Red Sea; otherwise IndoWest Pacific. Notes on the above two species:

Both species are variable in colour and easily confused; Ceratosoma tenue has smaller lateral projections but more importantly has a pair of secondary ones. The gill and rhinophore pockets are not edged with a purple line in Ceratosoma tenue.

204 Species accounts

Suborder Doridina

Family Chromodorididae

JH

Ceratosoma trilobatum 130 mm

B&SK

Species accounts 205

Suborder Doridina

Family Chromodorididae

Miamira magnifica Eliot, 1910 Size: 40 mm Diagnostic characteristics: The solid body is in the shape of the Cross of Lorraine, with a central ridge and four or six distinct lobes. The whole of the mantle is covered in nodules. There are two colour forms in the Red Sea: one is pale green with white nodules, the second has a pinkish orange ground colour with purple reticulations. Both have larger orange spots, and blue spots or rings. Ventrally, the foot and hyponotum are white with raised orange spots. Occurrence: Single specimen from Eilat (Israel), from a shallow reef flat with rubble; several photographic records from Aqaba (Jordan), Egypt, and Eilat. Geographic distribution: Northern Red Sea and tropical western Indian Ocean (Yonow 1994).

12 mm

B&SK

35 mm

Miamira magnifica 30 mm

J. Dafni

OL

206 Species accounts

Head OL

Suborder Doridina

Family Chromodorididae

Orodoris miamirana Bergh, 1875 Size: 90 mm Diagnostic characteristics: The solid oval body has numerous permanent scallops around the margin. The central median ridge extends as four or five perpendicular ridges to the margin. The colour of the animal is complicated shades of brown, red, purple, and green as reticulations; the pustules are cream, the blue spots are ringed with red, and everything appears out-of-focus. Eye-like spots are present in the spaces between the lateral ridges, near the median ridge. Ventrally there are pale to dark green reticulations and blue ocelli can be present in the junction between foot and mantle. Occurrence: Several photographic records, mostly from Eilat, Israel, one from Egypt, and few from Jeddah, Saudi Arabia. Geographical distribution: Tropical Indo-West Pacific. Note: The three genera described above were recently amalgamated into one genus, Ceratosoma, a move which muddies the taxonomic waters a little more and is not adopted here.

JK

Orodoris miamirana

JK

OL

Species accounts 207

Suborder Doridina

Family Dendrodorididae

Porostome dorids The two families Dendrodorididae and Phyllidiidae were combined into one grouping by Bergh in 1876 due to their pore-like mouth leading to a long buccal pump and the lack of radula in both families. The dendrodorids are most similar in body shape to the soft cryptobranch dorids, such as the Chromodorididae. The Phyllidiidae differ completely in being stiff and rubbery, covered in tubercles which are often capped in yellow, and with no dorsal gills. Family Dendrodorididae Most species of Dendrodoris are typically dorid in shape, oval with anterior rhinophores and posterior gills. They are usually soft, somewhat slimy, and usually have an ample skirt and broad margin. Some species can have pustules or large soft tubercles. The rhinophores and gills are located very close to the anterior and posterior edges respectively, and the rhinophores have a distinctive “bent-back” shape. Both the gills and the rhinophores withdraw completely into pockets. Lacking radula and jaws, dendrodorids have a long oral tube which they insert into the sponge prey and use secretions to digest it externally. They then suck up the juices. The genus Dendrodoris has never been completely revised, and some confusion surrounds the older species names. Dendrodoris coronata Kay and Young, 1969 Size: 35 mm Diagnostic characteristics: The body shape is humped with a central transverse ridge or ‘crown’ of tubercles (hence the name coronata). The mantle colour is translucent yellow to orange, covered in black and white flecks. The visceral hump additionally has patches of grey or brown. The arborescent gills are orange with grey axes and the rhinophores are orange with grey stalks. The foot is speckled with black.

Dendrodoris coronata

208 Species accounts

JK

Suborder Doridina

35 mm

Family Dendrodorididae

JH

JK

PK

Pseudobiceros sp.

Dendrodoris coronata

JK

Occurrence:

Appears to be rather common; found underneath coral heads covered in epiphytes and epifauna, in shallow waters; several specimens and additional photographic records from Jeddah, Saudi Arabia, and the northern Red Sea. Geographical distribution: First Red Sea records; otherwise tropical western Pacific. Species accounts 209

JH

JK

Suborder Doridina

Family Dendrodorididae

JK

Dendrodoris elongata Baba, 1936 Size: ? 35 mm Diagnostic characteristics: The body is very elongated, gently domed, with soft projections on the hump and smaller pustules on the sides. It is translucent white with larger brown spots arranged in an approximate line centrally and brown speckles all over. The pustules and conical projections are white as are the ends of the rhinophores. Occurrence: Several specimen and photographic records from the northern Red Sea; found in shallow water under stones. Geographical distribution: New species record for the Red Sea (unless one of the many Red Sea names available proves valid); recorded from Réunion and the West Pacific.

JH

Dendrodoris elongata

JK

JH

210 Species accounts

Suborder Doridina

Family Dendrodorididae

Dendrodoris fumata (Rüppell and Leuckart, 1830) Size: 65 mm Diagnostic characteristics: The body has a dorsal hump with a thin, flattened, undulating margin. It is variable in colour with two forms occurring in the Red Sea. There is a smoky grey form with grey green patches and a paler margin. The pale pink to red form has no coloured patches but has some darker markings. The rhinophores match the body colour but have white tips regardless of their colour. The six gills are large and plumose, often extending to or beyond the margins. Dendrodoris fumata is easily confused with some colour forms of D. n igra.

JK

JK

Dendrodoris fumata 45 mm

JH

Pseudobiceros uniarborensis JK

Biology and ecology: Reproductive activity begins in December to January and peaks between May and October, producing orange coloured egg ribbons (Gohar & Soliman (1967a). Occurrence: Rare in the intertidal and shallow sublittoral, found in calm bays, back reefs, underneath rock and coral rubble; several specimen and photographic records. Geographical distribution: Thought to have been endemic to the Red Sea but these colour forms are now known from the tropical Indo-Pacific.

Species accounts 211

Suborder Doridina

Family Dendrodorididae

Dendrodoris nigra (Stimpson, 1855) Size: 50 mm Diagnostic characteristics: Dendrodoris nigra is easily confused with D. fumata, but has a longer and thinner body shape. It also has a flattened and crumpled margin. The mantle is soft but may have low soft pustules. In colour it is much less variable in the Red Sea, jet black or very dark grey, usually with few or many white speckles which may be grouped together. The margin may have a red band near the edge. The gills are located very far back and held in a close clump, the spread being much smaller than in D. fumata. The rhinophores are black with white tips. Biology and ecology: Intertidal and shallow sublittoral; more common than Dendrodoris fumata, but care must be taken with identification of the two species. Occurrence: Some specimens and many photographic records. Geographical distribution: Tropical and warm temperate waters, excluding the American continent.

JH

Dendrodoris nigra

212 Species accounts

JK

Suborder Doridina

Family Dendrodorididae

Dendrodoris nigra

Pseudobiceros gloriosus

JK

Pseudobiceros ? ocellatus

JK

JH

Dendrodoris tuberculosa (Quoy and Gaimard, 1832) Size: approximately 230 mm Diagnostic characteristics: The soft dorsum is completely covered with large complex tubercles which become polygonal around their edges. A specific feature is the series of beautiful white spots outlined in black on the hyponotum. Measuring approximately 18 mm when alive, the juvenile shows developing characteristics of adult D. tuberculosa but the preserved specimen has no white spots on its hyponotum (over).

Dendrodoris tuberculosa

JK

Ventral surface

Species accounts 213

JK

Suborder Doridina

Family Dendrodorididae

Dendrodoris tuberculosa

Dendrodoris tuberculosa Juvenile, 18 mm

V. Neumann

Occurrence:

Single juvenile specimen from Aqaba; several photographic records from the northern and central Red Sea; intertidal and shallow subtidal on reef plateaus and sand platforms. Geographical distribution: Indo-West Pacific; two species can co-exist in similar habitats at the same locality, and hence records are confused with Dendrodoris carbunculosa (Kelaart), which has tuberculate discs on the dorsum with spaces between them, and no spots on the hyponotum (Yonow et al. 2002). Dendrodonis carbunculosa has not been recorded from the Red Sea to date. Dendrodoris sp. Size: ? 40 mm Diagnostic characteristics:

The dorsum is patched in shades of brownish red and covered in what appears to be raised pimples. These little pustules are brown or black with a white ring around them. They are of a similar size and evenly distributed all over the mantle. The gills and rhinophores are brown; the gills have white midribs, and the rhinophores have white tips. The edge of the mantle is ruffled, and the dorsum is high in profile. Occurrence: Three photographic records from Jeddah, Saudi Arabia. Geographical distribution: Red Sea and Réunion. Notes: Close-up photographs of the rhinophores and ventral surface of an almost identical individual from Réunion (P. Bidgrain) indicate that they may be a species belonging to the family Discodorididae. 214 Species accounts

Family Phyllidiidae

Suborder Doridina

Dendrodoris sp.

JK

T. Metcalfe

Family Phyllidiidae Family members are very distinctive; the rubbery body is elongate oval, most often tuberculate, and without a dorsal gill circlet: the secondarily developed gills are present as a series of leaflets lying between the foot and the mantle. The anus is located on the dorsum, in the normal dorid position, in all genera except Fryeria, where it can be found at the end of the body between the foot and the mantle. Species are usually black, white, and orange in colour, although species of Phyllidiella lack the orange and white and have a pinkish tinge. Some species of Phyllidia and Fryeria have a blue tinge. Species of Phyllidiopsis are extremely variable and have no colour rule. Phyllidiidae feed on sponges by extruding their mouthparts as they have no jaws or radula. They digest the sponge externally, and suck up the juices. The terpenoids from the sponge are concentrated in dorsal glands; many species exude a white secretion when disturbed and some also smell terribly. As a result, fish and crab predators avoid them (Gunthorpe & Cameron 1987). Fryeria rueppelii (Bergh, 1869) Size: 60 mm Diagnostic characteristics: The oval black central region extends to the margin as rays. The central line of tubercles is tipped with orange, and the rhinophores are bright orange. Unique to this species is an orange edge to the mantle, which (very rarely) breaks up into segments.

TP

Fryeria rueppelii

JK

Species accounts 215

Family Phyllidiidae

Suborder Doridina

25 mm

NY

Fryeria rueppelii

OL

60 mm

B&SK

JK

Occurrence:

Common on coral reefs, and present nearly all year round throughout the Red Sea. Geographical distribution: Endemic to the Red Sea, but also present in the Gulf of Aden. The similar Indian Ocean F. marindica Yonow & Hayward has no orange margin. Phyllidia multifaria Yonow, 1986 Size: 45 mm Diagnostic characteristics:

The central area is composed of tubercles tipped in orange yellow and black lines forming semi-ellipses around the margin. The rhinophores are bright orange. A black line is present between the hyponotum and the foot near the gills, and a solid black line marks the sole of the foot.

Phyllidia multifaria Foot

B. Picton

216 Species accounts

PK

Family Phyllidiidae

Suborder Doridina

Phyllidia multifaria

JH

Occurrence:

Common on coral reefs and slopes in the main Red Sea; more common further south. Geographical distribution: Endemic to the Red Sea; the similar Phyllidia elegans Bergh exists in the Indo-Pacific. Phyllidia schupporum Fahrner and Schrödl, 2000 Size: approximately 30 mm Diagnostic characteristics: Most

easily confused with Fryeria rueppelii, this species does not have an orange border around the mantle. The body is white with rather high conical tubercles, the largest of which are tipped in yellow. Two black lines extend along the median ridge from behind the rhinophores to the posterior mantle margin. There are rays extending from these two lines to the margins, and additionally there are some lines

Phyllidia schupporum 25 mm

B&SK

Species accounts 217

Holotype M.

Schrödl

PK

Family Phyllidiidae

Suborder Doridina

Ventral, 25 mm

B&SK

which cross the central ridge. The rhinophores are orange, a little different in colour from the tubercles, and there is no line on the sole of the foot. The conical oral tentacles are tipped with orange in the holotype. The anus is difficult to see, in the black area beyond the last large central tubercle, adding to the confusion with Fryeria rueppelii. Occurrence: These are the first records since this species was described from a single specimen; two photographic records from Eilat and Jeddah indicate a distribution limited to the northern Red Sea. Geographical distribution: Endemic to the Red Sea. Phyllidia undula Yonow, 1986 Size: 70 mm Diagnostic characteristics:

18 mm

NY

The mantle varies from bright orange to yellow, with a black scalloped pattern down each side. The black is edged in white. The largest tubercles are contiguous with the white band, along the midline and within the scallops. The rhinophores are the same colour as the mantle. There may be black flecks on the sides coalescing to form patches. Occurrence: Common on coral reefs and coral rubble ledges, Phyllidia undula is more frequently encountered in the southern Red Sea. Geographical distribution: Red Sea and eastern to southern Africa.

Phyllidia undula

OL

218 Species accounts

Family Phyllidiidae

Suborder Doridina

Phyllidia undula

JH

18 mm

NY

Phyllidia varicosa Lamarck, 1801 Size: 100 mm Diagnostic characteristics: The body bears three rows of orange-tipped tubercles and four black lines on the dorsum. A broken black line is present on the sole of the foot, and the head has orange-tipped oral tentacles. Biology and ecology: Like many phyllidiids, Phyllidia varicosa produces an extremely unpleasant odour when handled. Spawning was only observed in the laboratory from July to October: the flat tight coil was creamy white up to 30 mm in diameter (Soliman 1986). Occurrence: Very common on coral reefs and coral rubble; not yet recorded from the Gulf of Eilat and more frequently encountered in the southern Red Sea. Geographical distribution: A very common Indo-West Pacific species.

JH

Phyllidia varicosa

JK

Species accounts 219

Family Phyllidiidae

Suborder Doridina

Phyllidiella pustulosa (Cuvier, 1804) Size: 70 mm Diagnostic characteristics: A very common species variable in colour and pattern. Generally, the ground colour is black with groups of greenish (under water) or pinkish (out of water) tubercles. Sometimes the black can be very thick or present as thin lines. In most species of opisthobranchs, the juveniles resemble the adults, but in this species it is not identical, being much smoother and the groups of tubercles not divided with black. The rhinophores are black and there is no black line on the sole. Occurrence: Very common on coral reefs, present all year round throughout the Red Sea. Geographical distribution: Widespread Indo-West Pacific species.

PK

Juvenile, 25 mm

Phyllidiella pustulosa

PK

220 Species accounts

JH

JK

Family Phyllidiidae

Suborder Doridina

Phyllidiopsis cardinalis Bergh, 1873 Size: 70 mm Diagnostic characteristics: This is a very ‘untypical’ phyllidiid. Two to four rows of large complex tubercles are located behind the rhinophores. The general colour effect is one of browns and yellows with tinges of red, orange, olive, purple, and pink. The rhinophores appear green. A diagnostic feature is the bright yellow foot and hyponotum; even the gill lamellae are yellow. The upper edge of the foot bears dark spots which are indistinct around their edges. Occurrence: Several photographic records; under dead coral in reef lagoon at 1 m depth at Hurghada and deeper than 20 m at Eilat. Geographical distribution: First records from the Red Sea; otherwise recorded from the Indo-West Pacific. It is less common in the Indian Ocean, known only from Western Australia, Aldabra, Réunion, and Chagos (Yonow et al. 2002).

R. Cohen

Phyllidiopsis cardinalis

Ventral surface

B&SK

Species accounts 221

B&SK

Family Phyllidiidae

Suborder Doridina

GS

Phyllidiopsis dautzenbergi (Vayssière, 1912) Size: 20 mm Diagnostic characteristics: One of the smallest of the Red Sea phyllidiids, this nudibranch has a smooth white body, sometimes almost translucent, with a central black ellipse and five paired black lines radiating from it. The rhinophores are translucent white, as is the foot. Occurrence: Found under coral heads and rocks; rarely recorded but five photographic records. Geographical distribution: Endemic to the Red Sea and Gulf of Aden; recorded only from the southern half of the Red Sea.

G. Bemert

Phyllidiopsis dautzenbergi

H. Schmid

PK

222 Species accounts

Family Phyllidiidae

Suborder Doridina

Phyllidiopsis monacha (Yonow, 1986) Size: 20 mm Description: The central region of irregular black markings with five rays extending to the margin and a posterior one is identical to the holotype, as is the central irregular tuberculate patch and the two oval patches surrounding the rhinophores. There are few white tubercles around the edges of the black area, which are lined with white crescents and orange or yellow semi-circles. The rhinophores are coloured orange/yellow, a novel character for species of Phyllidiopsis which usually have half black, half white, ochre, or white rhinophores. Occurrence: Two photographic records from Jeddah, Saudi Arabia, and Eilat, Israel. Geographical distribution: Endemic to the Red Sea. Note: This species was originally described from a preserved specimen which had no colour remaining. Phyllidiopsis monacha is the first species of Phyllidiopsis to be recorded as having bright yellow or orange pigmentation but examination of further specimens will confirm the precise taxonomic placement and possibly redefine criteria of the genus.

T. Metcalfe

B&SK

Phyllidiopsis monacha

B&SK

Species accounts 223

Family Phyllidiidae

Suborder Doridina

Phyllidiopsis sinaiensis (Yonow, 1988) Size: 85 mm Diagnostic characteristics: This is the only pink and black species which could be confused with Phyllidiella pustulosa. The tubercles and black lines are arranged in a longitudinal pattern, not in groups, and the thickness of the black is variable. The black is present as two main longitudinal lines with many rays extending to the margins and sometimes secondary lines across the median ridge of tubercles. The tubercles are very complex in the largest specimens. The rhinophores are half black and half pink. Biology and ecology: Phyllidiopsis sinaiensis produces a very unpleasant smell when handled. It can be found all year on rocks and boulders; its behaviour is not at all cryptic. Occurrence: Numerous photographic and specimen records, present all year round in the Gulf of Eilat. Geographical distribution: Endemic to the Red Sea, common in the northern part, but absent from the southern half of the Red Sea.

48 mm

NY

Phyllidiopsis sinaiensis

GS

PK

224 Species accounts

Suborder Dendronotina

Family Tritoniidae

Suborder Dendronotina A relatively small suborder of the Nudibranchia, the body form is as varied as in the larger suborders. Characteristic of all families are the secondary gills aligned in a paired series on the pallial margin. These may be large and bushy as in the Tritoniidae, protected by flaps as in the Bornellidae, Scyllaeidae, and Tethyidae, tuberculate as in the Dotoidae (and some Tethyidae), or completely reduced to small flaps as in Lomanotus. There is always a frontal veil which is variously developed. Family Tritoniidae The body is high in profile, approximately quadrilateral in cross section in most species. The mantle edge is produced in a series of tree-like secondary gills along the sides. The oral tentacles on each side of the veil can be very reduced. The rhinophore sheaths are obvious, usually flared, and the rhinophores themselves are complicated and often branch distally. The diet comprises of alcyonarian soft corals and gorgonians. In species which have been well studied, each species of nudibranch is species-specific on a food item. Marionia glama (Rüppell and Leuckart, 1828) Size: 60 mm Diagnostic characteristics: The dorsum is sculptured with raised pustules and reticulated valleys in between. The body colour is almost iridescent in shades of pale blue, pale lilac, and pale green. The creases are dark green and the pustules are capped in pale pink. There are at least 12 very bushy gills on each side which alternate in size along each side. They also alternate in orientation, vertically and horizontally. The oral veil has simple digitiform processes, and the rhinophore sheaths are flared and scalloped.

JK

Marionia glama

JK

Species accounts 225

Suborder Dendronotina

Family Tritoniidae

Marionia glama 50 mm

B&SK

JK

Occurrence:

Several photographic records from Eilat, Israel and from Jeddah, Saudi Arabia; the original figure (Tab. 4 fig. 2) is reproduced on p. 39. Geographical distribution: Only known from the Red Sea although the species recorded by Gosliner (1987) from South Africa may be Marionia glama (fig. 207, 11 mm). Marioniopsis cyanobranchiata (Rüppell and Leuckart, 1828) Size: 45 mm Diagnostic characteristics:

The dorsum is golden yellow tending to orange with blue green reticulations in the crevices between the soft pustules. There are dark markings on the dorsum behind the rhinophores and sometimes between pairs of gills. The rhinophores are yellow and blue green; the club has arborescent processes and the sheath is simply flared.

Marioniopsis cyanobranchiata

JK

226 Species accounts

Suborder Dendronotina

Family Tritoniidae

The gills are unequal in number on each side but evenly sized. The yellow basal stalk is chunky and the pinnate branches are blue green with white. Occurrence: Found in the shallow subtidal on reef tops with coral rubble; numerous photographic records from the central and northern Red Sea. Geographical distribution: Endemic to the Red Sea; other records (Tanzania, South Africa, and Indonesia) appear to represent a different species (Yonow 2000). Rüppell & Leuckart’s plate (Tab. 4 figs. 3a, b) is reproduced on p. 39.

JK

Marioniopsis cyanobranchiata

GS

Head DPS

Species accounts 227

Suborder Dendronotina

Family Tritoniidae

JK

Marioniopsis levis

B&SK

OL

Marioniopsis levis (Eliot, 1904) [= M. fulricola Avila et al., 1999] Size: 40 mm Diagnostic characteristics: The body is long and thin with smaller gills than the other Red Sea species. It is variable in colour but usually with a brown and white transverse linear pattern and green overtones if not olive green or grey all over. The bases of the gills and rhinophores are all spotted with darker reddish brown. The rhinophore sheaths are simply flared; the gills alternate along each side from facing semi-vertically to semi-horizontally. A very pale form also exists. Biology and ecology: Common all year in Eilat (B. & S. Koretz, pers. comm.); feeds on the soft coral Parerythropodium fulvum fulvum (Avila et al. 1999); the egg mass is a yellow brown flat spiral found in the spring (B. & S. Koretz, pers. comm. and Avila et al. 1999). Occurrence: Numerous photographic records from Eilat and the Gulf, but fewer records from Jeddah and the central Red Sea. Geographical distribution: Red Sea and East Africa. 228 Species accounts

Suborder Dendronotina

Family Tritoniidae

Marioniopsis rubra (Rüppell and Leuckart, 1828) Size: 100 mm Diagnostic characteristics: The body is pale pink or orange to deep orange with pink over-tones. The plumose gills are of a similar size, and each has a pink stalk with a pinkish white to golden crown. The dorsum is pustulose, and the oral veil is almost semi-circular with dividing digitations. Rüppell and Leuckart’s illustration (Tab. 4 fig. 1) is reproduced here on p. 39. Occurrence: Shallow reef tops, generally nocturnal; most frequent in spring in Eilat (B. & S. Koretz, pers. comm.); several photographic records from the northern and central Red Sea. Geographical distribution: Endemic to the Red Sea; the single record from Mauritius by Odhner (1936) is probably an error (Yonow 2000).

B&SK

DPS

B&SK

DPS

Marioniopsis rubra

OL

Species accounts 229

Suborder Dendronotina

Family Tritoniidae

Marioniopsis viridescens (Eliot, 1904) Size: 25 mm Diagnostic characteristics: The body is quite long and slender, the 10 or 11 paired gills sparse in comparison to M. rubra, with which it could be confused, but are held on long stalks. They alternate the direction they face along each side, but are evenly sized. The colour is somewhat variably marbled with beige and brown, sometimes with orange or pink overtones. There is always a distinct symmetrical white pattern down the

Marioniopsis viridescens Eilat

OL

230 Species accounts

Suborder Dendronotina

Juvenile, Eilat

Family Tritoniidae

OL

JK

Marioniopsis viridescens Aqaba

TP

dorsum which branches to the gills bases. The frontal veil has long processes which divide at the ends. The rhinophore sheaths are flared and crenulated, and may have spots. The rhinophores are much branched and resemble the gills. A white variety is common in the Gulf of Eilat. Biology and ecology: Reported to be nocturnal; two photographs show it near Heteroxenia colonies on which it may feed (B. & S. Koretz, pers. comm.). The white form is usually found a white soft coral (as yet unidentified). The sparse gills of M. viridescens are identical to the coral polyps. Occurrence: Found in shallow water; numerous photographs from the Gulf of Eilat, one photographic record from Jeddah. Geographical distribution: Western Indian Ocean and Red Sea. Species accounts 231

Suborder Dendronotina

Family Tritoniidae

Tritoniopsis elegans (Audouin, 1826) Size: 50 mm Diagnostic characteristics: This tritonid is rather squat and oval in outline. It has a broad dorsum with almost horizontal gills. The ‘classic’ colour form (painted by Savigny from the Red Sea, see p. 35) is red orange and white, but an all-orange form also exists in the Red Sea. The gills are held horizontally and form an almost complete fringe around the body; as a result, the watery orange rhinophores and their sheaths are nearly indistinguishable. The oral veil is broad but with short simple processes. The tail protrudes as a triangular appendage to the feathery mass. Occurrence: Several photographic records from Eilat and Jeddah. Geographical distribution: Widespread in the Indo-West Pacific and occurring there in many colour forms.

Two individuals

Tritoniopsis elegans

OL

JK

232 Species accounts

B&SK

Suborder Dendronotina

Family Bornellidae

Family Bornellidae The small family is monotypic; species have sinuous, elongated bodies with no mantle margin. There are complicated papillae or flaps protecting the gills and rhinophores. The rhinophores additionally have digitate sheaths. The oral veil is divided into two distinct lobes, each digitate like small hands. The species are thought to feed on hydroids. Bornella stellifer (Adams and Reeve in Adams, 1848) Size: 40 mm Diagnostic characteristics: The body is cream to white and covered with orange marbling and reticulations. The ends of the cerata, rhinophore processes, and oral processes are usually ringed with a line of orange pigment. Not to be confused with Flabellina bicolor (p. 244). Occurrence: Originally recorded from the Gulf of Aden, these are the first recent records from the northern (same specimen as in Yonow 2000) and central Red Sea; additional specimen record from Sudan (Suakin Expedition 1980). Geographical distribution: Red Sea and Indo-Pacific.

JK

Bornella stellifer 20 mm

JH

20 mm

JH

Species accounts 233

Suborder Dendronotina

Family Dotoidae

Family Dotoidae The Dotoidae are a very small family in the Red Sea, with only one species to date. The body has paired swollen cerata, which can be evenly or irregularly tuberculate. The tubercles may have coloured spots on the ends. A gill may be present at the base of the cerata in some species. There is a rounded oral veil, the smooth rhinophores are long and tapering, and issue from tubular or flared sheaths. All species feed very specifically on hydroids, usually on one or few species belonging to a single genus. Doto orcha Yonow, 2000 Size: 10 mm Diagnostic characteristics: Doto orcha has a narrow creamy body with greenish beige longitudinal central band; the body is very tuberculate. There are eight pairs of cerata which are irregularly tuberculate. The black spot at the base of each ceras may not always be obvious and the central green band can be faint. Biology and ecology: Probably an obligate feeder on the sertulariid hydroid Dynamena disticha (Bosc), growing on sea grasses and other substrata. Occurrence: One specimen from the northern Red Sea (Yonow 2000) and one photographic record from Eilat, Israel. Geographical distribution: Doto orcha may be endemic to the Red Sea; however, the hydroid is widespread in the tropical Indo-Pacific.

Doto orcha Holotype, 10 mm

234 Species accounts

JH

Suborder Dendronotina

Family Lomanotidae

Family Lomanotidae A small family with few temperate and tropical species. They have a mantle margin produced into a single row of flattened or concave cerata. The rhinophores are lamellate (not typical of dendronotaceans) and there is no digitate or branching veil at the front, only two or four tentacles. All species feed on hydroids. Lomanotus vermiformis Eliot, 1908 Size: 25 mm Diagnostic characteristics: This nearly invisible opisthobranch is long and tapering, almost square in cross-section. It is coloured in beige, brown, and olive green. The pallial margin is just visible as translucent white lines along each side of the body with short cerata. There may be a brown or white linear pattern on the dorsum. The rhinophore sheaths

Lomanotus vermiformis Philippines

T. Wanner

Species accounts 235

Suborder Dendronotina

Family Lomanotidae / Family Scyllaeidae

are flared and scalloped around the margins; the rhinophores are usually darker, with few oblique lamellae on the posterior face and a straight projection on the anterior side. Biology and ecology: Obligate predator of the aglaopheniid hydroid Macrorhynchia philippina (Kirchenpauer) (i.d. W. Verwoort) on which it is extremely cryptic. Occurrence: None since Eliot (1908; 45 specimens on “plumularian hydroid”) and Pruvot-Fol (1933; one specimen from bottles containing especially hydroids and sponges) but the hydroid occurs in the Red Sea so we can expect Lomanotus vermiformis. Geographical distribution: Red Sea and Pacific (east and west); no records from the Indian Ocean. Note: Lomanotus vermiformis was originally described from the Red Sea; there are no recent records but the species is illustrated here with a photograph from the Philippines. Family Scyllaeidae Scyllaeids are elongated but laterally compressed dendronotaceans. There is no mantle ridge or skirt, but one or two pairs of lobes mark the margin. Each lobe bears gills on its medial surface. Species feed on epiphytic hydroids growing on brown algae and sea grasses. Scyllaea pelagica Linnaeus, 1758 Size: unknown for the Red Sea Diagnostic characteristics: The body

is pale green to brown with blue spots and white patches. The body has lateral papillae, two pairs of irregularly squared lobes, and a short tail with a prominent keel. The beautiful blue spots are clearly visible in the photograph of the darker individual (below). Biology and ecology: Associated with drifting oceanic brown algae and in the shallow sublittoral with Sargassum, Scyllaea pelagica can be beached intertidally with the weed. It feeds on the hydroids growing epiphytically on Sargassum.

Scyllaea pelagica

236 Species accounts

N. Kenward

Suborder Dendronotina

Family Scyllaeidae / Family Tethyidae

Scyllaea pelagica On Sargassum

H. Schmid

Occurrence: Two photographic records from the central and northern parts of the Red Sea. Geographical distribution: Worldwide temperate and tropical seas, utterly cosmopolitan; maximum size 60 mm (Western Australia, Wells and Bryce 1993).

Family Tethyidae These dendronotaceans have a very extended or dilated oral veil fringed with tentacles, and flattened or swollen cerata along the lateral edges of the mantle. Gills may be present on the inner face at the base of each ceras. Jaws are variable, may be absent in some species. Known to feed on plankton and small crustaceans. Melibe rangii Bergh, 1875 [= Melibe engeli Risbec, 1937] Size: 90 mm Diagnostic characteristics: The long thin body can be translucent cream or brown to opaque light brown, with many swollen cerata. The cerata are tuberculate and have elongate papillae at the ends. The body is also covered in papillae and tubercles. The rhinophores have a flared sheath with a large flap bearing elongated processes on the margin, clearly visible in the photographs (but often contracted or missing in preserved specimens). The enlarged buccal hood bears two to five rows of large and small tentacles inside the margin; the hood may be incised on its upper and lower margins. Species accounts 237

Melibe rangii

HS

Family Tethyidae

Suborder Dendronotina

Biology and ecology: The dozen or so cream spheres inside the body (diagnostic of this species) are the ovotestis (or gonad), and the amazing egg mass they ‘produce’ is visible in the photograph of the disturbed spawning animal. Species of Melibe swim unwillingly by folding the foot and flexing the body laterally. The expanded hood is used in feeding by stripping algal filaments (Schuhmacher 1973b). Juveniles are very transparent indeed, but darken with age (I. Burghardt, pers. comm.). Occurrence: Abundant in the spring in suitable habitats, such as calm shallow subtidal algal beds. Often occur with Aplysia oculifera (p. 97) and Notarchus indicus (p. 105).

HS

Melibe rangii

238 Species accounts

JH

Family Tethyidae

Suborder Dendronotina

HS

B&SK

B&SK

Melibe rangii Spawning

HS

Geographical distribution: Red Sea and West Pacific. Notes: The genus Melibe in the Indo-West Pacific continues

to be problematic. Contraction of preserved specimens varies greatly, making it almost impossible to define boundaries between species until live material is available. Melibe rangii was recorded from a single specimen from the Red Sea and designated an uncertain species in subsequent reviews. It has been recorded (erroneously) as M. bucephala; both were species originally described from and endemic to the Red Sea. However, it is quite clear that the common West Pacific species Melibe engeli Risbec (syn. nov.) is the same species as M. rangii. The characters of M. rangii fall well within the range of those observed and described for M. engeli, especially as the original descriptions and illustrations were based on preserved material. Melibe rangii precedes M. engeli by sixty – two years. The only true record of M. bucephala Bergh appears to be by O’Donoghue (1929). This species is more opaque, greenish brown with a wider foot, and less extensive rhinophore sheaths. The records misidentified as M. bucephala (see table p. 63) confirm the presence of a species common at certain times of the year in the Red Sea, which should be known as M. rangii. Species accounts 239

Suborder Arminina

Family Arminidae

Suborder Arminina The Arminina are a very diverse group, encompassing species which do not belong to the other suborders. As a result, families are combined into groups unrelated to each other, and morphology is extremely variable. Species can be ridged or tuberculate as in the Arminidae, covered in cerata like Madrella, or bear an undulating extension of the margin in some temperate and tropical families. Family Arminidae The flattened body is teardrop in shape, smooth or with longitudinal ridges or tubercles. There are no dorsal gills; secondary gill leaflets are found between the mantle and foot in Armina. The retractile rhinophores are located far anteriorly, separated from the mantle by a ridge in Armina. The rhinophore lamellae are vertical in this family and not horizontal as in the dorids. Species are usually found on or in sandy substrata, and feed on pennatulaceans. The most common species in the Red Sea appears to be un-named!

Pseudoceros sp. JK

Dermatobranchus albus (Eliot, 1904) Size: 15 mm Diagnostic characteristics: The ridged body is white with patches of black and lines of white and yellow. The mantle margin is edged with yellow as is the white foot. The oral veil is white bordered by a yellow line. The rhinophore stalks are white and the vertically lamellate portion is black. Occurrence: Two photographic records from the Gulf of Eilat. Distribution: First record for the Red Sea; otherwise, western Indian Ocean (Yonow et al. 2002).

B&SK

Dermatobranchus albus

B&SK

240 Species accounts

Suborder Arminina

Family Arminidae

Dermatobranchus gonatophorus van Hasselt, 1824 Size: unknown for the Red Sea Diagnostic characteristics: The body is white with longitudinal ridges originating near the rhinophores and splaying out to the margin: in the central area, the ridges are black-topped while in the next region ridges may have black outgrowths. The outermost section of ridges is white, and there are black outgrowths just inside the orange margin. There may be grey shading centrally. The frontal veil is white with at least two black patches and an orange margin. The rhinophores are black. Occurrence: Few photographic records. Geographical distribution: There is one recent record from Indonesia and Okinawa (Gosliner et al. 1996) since the original description but no size was given; Bergh (1905) had a 35 mm specimen from Indonesia.

H. Schmid

Dermatobranchus gonatophorus

E. Lieske

Species accounts 241

Suborder Arminina

Family Arminidae

Dermatobranchus ornatus (Bergh, 1874) Size: approximately 50 mm Diagnostic characteristics: The body is opaque white with rounded tubercles, sometimes compound, and each with an orange ring at its apex. The margin of the mantle, oral veil, and foot are red. The foot additionally has black lines and spots around its edge. The rhinophores are black and set very close together. The edges of the longitudinal lamellae and the tips are white. Occurrence: Two photographic records: 15–20 m depth from near Hurghada, Egypt and near Dahab, Sinai. Geographical distribution: First record for the Red Sea; otherwise tropical Pacific.

Dermatobranchus ornatus

M. Jozon

Dermatobranchus sp. Size: 25 mm Diagnostic characteristics:

The dorsum is longitudinally ridged with the ridges splaying out to the edges. The ridges are lightly tuberculate. Colour is variable, generally pale with patches of lighter pigment on the ridges and black spots. Most distinctive is the blue grey band around the margins of the veil and the mantle containing creamy orange patches alternating with a row of black dots. The rhinophores have yellow longitudinal lamellae. Occurrence: Common on coral reefs, with numerous photographic records from Eilat, Israel and Jeddah, Saudi Arabia. Geographical distribution: Probably a new species, possibly endemic to the Red Sea.

Dermatobranchus sp.

JK

242 Species accounts

JK

Family Madrellidae

Suborder Arminina

Family Madrellidae The Madrellidae are a very small family of nudibranchs, containing only one genus and probably one species. Cerata are present in a fringe all around the body. Species feed on encrusting bryozoans, on which they are well camouflaged. They release a fluid when they are disturbed and can also cast off the cerata. The common Indo-West Pacific species is recorded from the Red Sea for the first time; a second unidentified species has also been photographed (see “Incertae Sedis”). Madrella ferruginosa Alder and Hancock, 1864 Size: 6 mm, probably larger Diagnostic characteristics: This rare species is distinguished by a red body and semi-translucent red cerata with contrasting ‘coral sand’ rhinophores and patch between them. The cerata are soft and elongated, pointed at the tips, and are arranged all around the outer half of the body: the middle is bare of cerata. The rhinophores are rather odd, elongated and pointed with a swelling near the tip, and there is an oral veil. There can be speckles and patches of grey white elsewhere on the body and cerata. Biology and ecology: The species is known to feed on bryozoans, from which it obtains its pigment and upon which it is extremely well camouflaged. Occurrence: A single individual is recorded from the Gulf of Eilat in an area of coral reef and sea grasses (June 2007). Geographical distribution: This is the first record of this widespread IndoWest Pacific species in the Red Sea; a similar animal, M. sanguinea (Angas) is only known from Australia, and is probably the same species.

6 mm

Madrella ferruginosa 6 mm

B&SK

Species accounts 243

B&SK

Suborder Aeolidina

Family Flabellinidae

Suborder Aeolidina These nudibranchs are generally elongated with long tentacles anteriorly and the body symmetrically covered in projections termed cerata. These cerata contain ramifications of the digestive gland and, in many species, contain a sac at the distal end in which the nematocysts of the cnidarian prey are stored for later use (see p. 25). They do not have cerata around the front of the head as do some arminids (see Madrella, previous and Janolus, p. 275). The criteria defining genera and families are still under discussion. Family Flabellinidae Aeolids which have long propodial tentacles and oral tentacles. The rhinophores may be smooth, papillate, or lamellate. The long thin cerata are grouped on stalks. Species are usually brightly coloured, and feed on hydroids. Flabellina bicolor (Kelaart, 1858) Size: 15 mm Diagnostic characteristics: The translucent body has opaque white patches, watery orange perfoliate rhinophores, and only two or three cerata per cluster. The cerata have a subterminal orange ring. The oral tentacles can be swollen near the tips, often making them club-shaped. Occurrence: Uncommon in coral rubble, the shallow sublittoral, and sometimes found in pairs; three photographic records from Eilat and the remainder from the central and southern Red Sea. Geographical distribution: First Red Sea record (Yonow 2000); otherwise tropical Indo-Pacific.

Flabellina bicolor

PK

244 Species accounts

DPS

Suborder Aeolidina

Family Flabellinidae

Flabellina bicolor

G. Bemert

“Flabellina” sp. Size: ? 15 mm Diagnostic characteristics:

This species is very similar in external morphology to the one above, mainly differing in the rhinophores, which are long and smooth in this species. The body is translucent with opaque white patches, and there are two yellow spots surrounded by an opaque white irregular region on the head. There appear to be two rows of cerata in each clump, making the animal appear more substantial than

“Flabellina” sp. 15 mm

OL

15 mm

Species accounts 245

OL

Suborder Aeolidina

Family Flabellinidae

Flabellina bicolor. The long rhinophores and oral tentacles are translucent at their bases and opaque white for most of their lengths. Occurrence: Single photographic record from the Gulf of Eilat. Geographical distribution: This is probably a new species, which may be endemic to the Red Sea. Flabellina bilas Gosliner and Willan, 1991 Size: 25 mm Diagnostic characteristics:

The translucent body can have a deep orange tinge and is covered with elongated opaque white markings. The cerata have red and white rings near the tips and opaque white or cream bands near the bases. The rhinophores are translucent with two opaque white or cream bands, and the very long oral tentacles are swollen at the ends. Occurrence: There are several photographic records from the northern and central Red Sea. Geographical distribution: First records for the Red Sea; otherwise recorded only from the West Pacific.

JK

Flabellina bilas

T. Metcalfe

246 Species accounts

JK

Suborder Aeolidina

Family Flabellinidae

Flabellina rubrolineata (O’Donoghue, 1929) Size: 35 mm Diagnostic characteristics: The creamy white body has three beautiful magenta violet stripes down its length. There are similar colour bands on the cerata, and the propodial and oral tentacles. The rhinophores are papillate only on their posterior sides, and magenta on the anterior faces. The dark contents of the cerata can be visible in some specimens, and the tips of the cerata are golden yellow or orange. The body may have a violet tinge in some individuals. Occurrence: Not uncommon, specimen record from Sudan (Suakin Expedition) and numerous photographic records. Geographical distribution: Widely recorded in the Indo-Pacific.

JH

Flabellina rubrolineata

HS

JK

Species accounts 247

Suborder Aeolidina

Family Facelinidae

Family Facelinidae These aeolid families are difficult to define, but facelinids are generally moderately narrow with a wide foot and a long pointed tail. The cerata can vary in shape, and are grouped in clusters. The rhinophores are lamellate, and the oral tentacles and propodial tentacles are long. The radula consists of a single row of multicuspid teeth.

A. Stern

Facelina rhodopos Yonow, 2000 Size: 25 mm Diagnostic characteristics: The translucent body has a yellow or orange tinge on the upper surfaces and on the tentacles. The cerata are swollen and have a characteristic terminal nipple. They are transparent with an opaque pink band fading towards the tip and the digestive gland is visible within. The metapodium is very long, and has a dense pink longitudinal band and tip. Occurrence: Single pair of specimens (Yonow 2000), but many photographic records from the northern Red Sea, under stones on sandy substrates in shallow water. Geographical distribution: Currently recorded from the Red Sea and Réunion (P. Bidgrain, pers. comm.) so possibly a species with a limited western Indian Ocean distribution.

Facelina rhodopos

PK

248 Species accounts

Suborder Aeolidina

Family Facelinidae

JH

JK

JK

JK

Facelina rhodopos

JH

Species accounts 249

Suborder Aeolidina

Family Facelinidae

“Facelina” sp. Size: 25 mm

The rhinophores are not lamellated, but this species is so similar to F. rhodopos that it is placed here for comparison. The translucent body can be orange or lilac. There are pink and lilac (or orange and purple) bands on the cerata and lilac lines on the head, tentacles, and tail. The digestive gland is visible in the cerata. Occurrence: Numerous photographic records from the Gulf of Eilat; this species is easily recognized and certainly appears to be more common than Facelina rhodopos (previous). Geographical distribution: A new species currently known only from the Gulf of Eilat; once specimens are available, the genus can be established. Diagnostic characteristics:

OL

B&SK

“Facelina” sp.

250 Species accounts

B&SK

Suborder Aeolidina

Family Aeolidiidae

Family Aeolidiidae Somewhat flattened aeolids which have numerous densely packed cerata arranged in rows or arches. The rhinophores can be smooth, papillate, or annulate. The oral tentacles are long and the propodial tentacles are usually well-developed. The majority of species feed on sea anemones and related cnidarians. Aeolidiella alba Risbec, 1928 Size: 10 mm Diagnostic characteristics: The flattened body and numerous cerata of this tiny aeolid are translucent with opaque white markings. The cerata are short and plump, absent from a central band along the body. The white head is large, almost round, and carries unusual propodial tentacles and rhinophores. The tentacles are swollen with two constrictions and the rhinophores are long with two swellings. The rhinophores are translucent with white markings concentrated distally and beige marbling increasing basally. There are additional beige flecks on the cerata. Occurrence: Several photographic records from Eilat. Geographical distribution: First record for the Red Sea; originally described from New Caledonia, but published records indicate an Indo-Pacific distribution.

Aeolidiella alba

OL

OL

Baeolidia fusiformis Baba, 1949 Size: 23 mm Diagnostic characteristics: The cream body bears a row of lemon yellow rings each encircling a white spot along its length. The cerata are cylindrical (not flattened) and long, with a very long ceras in alternate rows. There is a large gap between the second and third rows. The rhinophores have papillae on their posterior faces. Species accounts 251

Suborder Aeolidina

Family Aeolidiidae Biology and ecology: Cryptic with a burrowing anemone in coral rubble under rocks and stones; the cerata bristle when the animal is disturbed. Occurrence: Record of four specimens (Yonow 2000) and numerous additional photographic records from the central Red Sea and Gulf of Eilat. Geographical distribution: Japan (type locality) and Red Sea.

JK

Baeolidia fusiformis

23 mm

NY

JK

Note:

A similar species has white markings on the body and cerata, including white rings, and is designated Baeolidia cf. fusiformis until specimens are collected. The several photographic records are from the central Red Sea and Gulf of Eilat.

Baeolidia cf. fusiformis

252 Species accounts

JK

JK

Suborder Aeolidina

Family Aeolidiidae

Limenandra sp. Size: 35 mm Diagnostic characteristics:

The long opaque white body has short, curled cerata, like fingers. There are brown speckles on the cerata, the rhinophores, and the oral tentacles indicating the presence of zooxanthellae. The rhinophores have papillae only on the posterior sides, and some of the larger cerata are also papillate. There is a series of yellow rings along the dorsum which may contain blue or white flecks, and one large one on the head. A 17 mm specimen was laying eggs in September (below). Occurrence: Found on coral sand in shallow water, common in the Gulf of Eilat; several photographs from the central and northern Red Sea (recorded as L. nodosa by Yonow 2000). Geographical distribution: A new species currently identified from the Red Sea and Hawaii only.

TP

JK

B&SK

Egg mass

Limenandra sp.

HS

Species accounts 253

NY

Suborder Aeolidina

Family Aeolidiidae

Spurilla australis Rudman, 1982 Size: unknown for the Red Sea Diagnostic characteristics: The colour of the dark brown body is due to zooxanthellae. There is a blue spot on the head. Blue and orange banding decorates the somewhat swollen cerata; there is a large gap between the first and second groups. The darker rhinophores are papillate. Biology and ecology: Occurs on the shallow reef flat, feeding on anemones. Occurrence: Several photographic records only from the northern and central Red Sea. Geographical distribution: First Red Sea record, otherwise Australia, where it grows to 35 mm.

JK

JK

Spurilla australis

254 Species accounts

JK

Suborder Aeolidina

Family Aeolidiidae / Family Eubranchidae

Spurilla major (Eliot, 1903) Size: 25 mm Diagnostic characteristics: The body and cerata are cream with dark brown speckling due to symbiotic zooxanthellae. The flattened cerata are translucent with the digestive network visible within, easily cast off. White caps tip the cerata. The head is characteristically swollen with distinct vertical cleft anteriorly. The brown rhinophores are short and papillate. Occurrence: Single published record (Yonow 2000); additional specimen record from Egypt and photographic record from the Gulf of Eilat. Geographical distribution: Tropical Indo-Pacific oceans.

Spurilla major

NY

25 mm

NY

Family Eubranchidae These are small, rarely recorded aeolids, usually colourful, with long rhinophores and oral tentacles. The cerata are few in number, usually in simple rows, and inflated, often with tubercular swellings. Species are found on gymnoblastic and calyptoblastic hydroids. Dunga ocellata (Alder and Hancock, 1864) Size: approximately 6 mm Diagnostic characteristics: This unusual species has a translucent body with large inflated cerata which have three or four whorls of tubercles. The ends of the cerata and tubercles are marked with white and creamy orange, especially on the sides facing outwards. There are red rings on the cerata and larger tubercles. The rhinophores and oral tentacles are both very long, and irregularly banded with white and red. Biology and ecology: Shallow reef habitats. Occurrence: Three photographic records from Eilat, Israel. Geographical distribution: Western Indian Ocean, growing to 10 mm.

Dunga ocellata

OL

Species accounts 255

Juvenile, 6 mm

B&SK

Suborder Aeolidina

Family Eubranchidae

Eubranchus rubropunctatus Edmunds, 1969 Size: 7 mm Diagnostic characteristics: This rarely recorded species is distinguished by a translucent body which is covered in white and brown speckles. Both the cerata and body are tuberculate; the cerata are large and few in number, with a distinctive knob on the end. The knob is translucent distally with a pinkish band outlined in white. This is followed by a turquoise blue band containing black spots; after the blue band the cerata broaden at a pink band. Both the oral tentacles and rhinophores are long; the rhinophores are tuberculate. Occurrence: Two single individuals are recorded from the Gulf of Eilat in areas of sea grasses and patchy coral. Geographical distribution: This is the third record of this species, originally described from Tanzania (Edmunds 1969) and recorded from the Great Barrier Reef (Marshall & Willan 1999).

7 mm

B&SK

Eubranchus rubropunctatus

256 Species accounts

OL

OL

Family Glaucidae

Suborder Aeolidina

Family Glaucidae This family of aeolids usually have long slender bodies and clusters of cerata in rows or arches. The rhinophores are once again variable: simple, annulate, papillate or lamellate. Species are often brightly coloured. Many species have a symbiotic relationship with zooxanthellae obtained from their alcyonarian prey, but others feed on egg masses, hydroids, and other cnidarians. Some species can be aggressive. Caloria indica (Bergh, 1896) Size: 25 mm Diagnostic characteristics: The translucent body has a single opaque white line marking the dorsum, and two on the head extending from in front of the rhinophores. The head has orange markings and very long oral tentacles which are mostly white with a yellow band. The fusiform cerata are banded with orange, dark blue, and yellow, and have white tips. The smooth rhinophores are suffused with orange basally and lemon yellow distally. The recurved propodial tentacles and the end of the long tail are yellow. Biology and ecology: Found in shallow reef habitats, feeding on hydroids; generally nocturnal in habit (B. & S. Koretz, pers. comm.). Occurrence: Single specimen record and several photographs from Eilat, single photographic record from the central Red Sea. Geographical distribution: First Red Sea records; otherwise tropical IndoWest Pacific.

N. Ben-Eliahu

Caloria indica

JK

B&SK

Species accounts 257

Family Glaucidae

Suborder Aeolidina

Favorinus japonicus Baba, 1949 Size: 30 mm Diagnostic characteristics: The body is translucent with dense white pigmentation also present on oral tentacles, the rhinophores, and on the very long and narrow tail. There are five clusters of cerata which are variable in colour but all have some white speckling on the distal, usually recurved, ends. The rhinophores have two or three distinctive swellings, and are translucent basally and white apically. Biology and ecology: Favorinus japonicus feeds on the egg masses of other opisthobranchs, so the colour of the cerata can vary considerably from cream to pink, green, beige, and brown. Occurrence: Single specimen record from Egypt (October 1995) and photographic records from shallow waters in Quseir, Egypt and Eilat, Israel. Geographical distribution: This is the first Red Sea record; otherwise Favorinus japonicus is known in the tropical Indo-Pacific.

Favorinus japonicus 30 mm

JH

258 Species accounts

Family Glaucidae

Suborder Aeolidina

Favorinus tsuruganus Baba and Abe, 1964 Size: 12 mm Diagnostic characteristics: The body, tentacles, and tail are white, while the cerata are orange with blue banding below the white tips. Inside the cerata, at the level of the blue banding, is a dark sphere. The rhinophores are most unusual, with three dish-like swellings: above these, the rhinophores are white except for the tip, which is covered in a grey speckling like the ‘dishes’. The uppermost ‘dish’ has white pigmentation. Occurrence: Several photographic records from the Gulf of Eilat. Geographical distribution: New records for the Red Sea and photographic records from Réunion (P. Bidgrain, pers. comm.); originally described from Japan, but otherwise recorded in the western Pacific only.

OL

Favorinus tsuruganus

OL

Species accounts 259

Family Glaucidae

Suborder Aeolidina

Glaucilla marginata Bergh, 1868 Size: unknown for the Red Sea Diagnostic characteristics: The three to four clumps bear numerous rows of cerata in each cluster. There is a short tail, and the breadth of the body (including the cerata) approximately equals its length. This nudibranch swims upside down, and its ventral (uppermost) surface is brownish with iridescent blue green flecks on cerata. The body is very dark blue, and the bubble to aid floatation is clearly visible in the region of the first cluster.

Glaucilla marginata

JK

Biology and ecology: A pelagic species feeding on Physalia and other siphonophores; the bubble of air is swallowed to aid flotation, and can be seen in the stomach since the individual floats upside down. The cerata can wave up and down, but movement is slight and non-directional. Occurrence: Single photographic record from the central Red Sea. Geographical distribution: Circumtropical, up to 35 mm in size.

Phyllodesmium colemani Rudman, 1991 Size: ? 20 mm Diagnostic characteristics:

The body is very long and thin, with not many cerata per clump which are well spaced out along the body. Animals are specked brownish green, the pigment coming from the symbiotic

Phyllodesmium colemani

260 Species accounts

OL

Family Glaucidae

Suborder Aeolidina

Phyllodesmium colemani

OL

OL

zooxanthellae, and there are opaque white patches on both the cerata and the body. The rhinophores and oral tentacles are long and smooth. Biology and ecology: A western Pacific species known to feed on the Organ pipe Coral, Tubipora musica Linnaeus. Occurrence: Two photographic records from the Gulf of Eilat. Geographical distribution: First record for the Red Sea; otherwise P. colemani is known only from the West Pacific. Phyllodesmium crypticum Rudman, 1981 Size: 40 mm Diagnostic characteristics: The body is covered in densely packed irregular cerata: these are flattened, covered in small pustules, and curled at the tips. The smooth tapering rhinophores are long and project far above the cerata. Usually the long oral tentacles also project anteriorly. The colour is generally pale, opaque, a fairly nondescript beige. The digestive gland branches into the cerata.

OL

Phyllodesmium crypticum

JK

Species accounts 261

Family Glaucidae

Suborder Aeolidina

Phyllodesmium crypticum

JK

JK

Occurrence:

Several photographic records from Eilat, Israel, usually in spring (B. & S. Koretz, pers. comm.) and Jeddah, Saudi Arabia. Geographical distribution: First records for the Red Sea; otherwise only known from the western Pacific. It is recorded here from Réunion for the first time (photographs P. Bidgrain). Phyllodesmium hyalinum Ehrenberg, 1831 Size: 35 mm Diagnostic characteristics:

The body is beige brown with long cerata which are pustulate and curved at the tips. Cerata are few in number, and the larger ones are pustulose on both surfaces for the distal third. The darker speckling present on some of the cerata is due to zooxanthellae.

Phyllodesmium hyalinum

G. Reinicke

262 Species accounts

Family Glaucidae

Suborder Aeolidina

OL

Phyllodesmium hyalinum On Heteroxenia

G. Reinicke

JK

Biology and ecology: Always in association with Heteroxenia fuscescens (Ehrenberg) and Xenia umbellata Lamarck on which it feeds, and where it is very cryptic. Numerous aeolids can be found in a single colony of these alcyonarians. In addition to the usual method of feeding, Phyllodesmium acquire zooxanthellae from their prey, which continue to photosynthesise in the nudibranchs, providing it with additional nutrients. Breeding reaches a peak during May to October in the northern Red Sea (Gohar & Aboul-Ela 1957b). Occurrence: Common where Heteroxenia and Xenia occur, usually in shallow water and on reef platforms, but currently recorded only from the northern Red Sea. Geographical distribution: Indo-West Pacific, if both are the same species. Notes: Red Sea specimens clearly differ from Indo-West Pacific records of the same name. The Indo-West Pacific, if species have many more cerata with no gap between clusters 1 and 2. The radulae of these Xeniafeeding species are similar, and so external morphology takes on more importance in identification.

Phyllodesmium magnum Rudman, 1991 Size: 100 mm Diagnostic characteristics:

This large species is long and ‘leggy’ with a pale blue or lilac body. The clustered somewhat flattened cerata are also blue or lilac, and banded with creamy yellow and tipped with brighter yellow; the tips curl up over the body. The juvenile is 15 mm long, from Sharm el Sheikh, Egypt, and shows the characters of the adults (over). Species accounts 263

Phyllodesmium magnum JK

Suborder Aeolidina

Family Glaucidae Occurrence:

Found subtidally in coral rubble with secondary colonizers; many photographic records throughout the Red Sea. Geographical distribution: Widespread in the tropical Indo-West Pacific. Note: This is probably the same species as Eolida bella Rüppell & Leuckart (Tab. 10 fig. 4; see p. 44), which should have taken precedence.

15 mm

JH

Phyllodesmium magnum

264 Species accounts

JK

Suborder Aeolidina

Family Glaucidae

Pteraeolidia ianthina (Angas, 1864) Size: 70 mm Diagnostic characteristics: Dragon-like aeolids which have a very long and sinuous body, with clumps of short cerata held close to the body. Colour is extremely variable, from brownish to violet, but always with distinctive banding on the long and mobile oral tentacles. The lamellated rhinophores are bulbous, and each tip is purple. The juvenile adequately demonstrates the colour variation in the species, but the cerata are not as numerous nor as large as in the adults (below left). Biology and ecology: As well as using zooxanthellae, these aeolids can store nematocysts from their hydroid prey and can give an unpleasant sting if roughly handled. Occurrence: Very common, usually associated with live coral, sublittoral; more common further south in the Red Sea. Geographical distribution: Common in the Indo-Pacific oceans.

NY

Pteraeolidia ianthina Juvenile

JK

DPS

JH

JK

Species accounts 265

Suborder Aeolidina

Family Embletoniidae

Family Embletoniidae Debate continues on whether Embletonia is a dendronotacean or an aeolid. Here it is classified with the aeolids due to a number of characteristics, including the presence of nematocysts in the peculiarly shaped cerata, always a single row on each side, and the simple rhinophores without the sheath typical of dendronotaceans. They feed on cnidarians and hydroids, but possibly also bryozoans and detritus. This family contains one genus with three or four species. Embletonia gracile Risbec, 1928 Size: 10 mm Diagnostic characteristics: The tubular body has a single row of cupshaped cerata on each side. It is translucent to white in colour, including the long simple rhinophores, the bilobed oral veil, and the unusual cerata. Biology and ecology: Found in the shallow intertidal, in rubble and reef flats, and under rocks and dead coral slabs. Occurrence: Single published record (Yonow 2000); second specimen record from Sudan (Suakin Expedition). Geographical distribution: Tropical and temperate Indo-West Pacific.

8 mm

JH

Embletonia gracile 8 mm

JH

266 Species accounts

Suborder Aeolidina

Family Tergipedidae

Family Tergipedidae Medium to large aeolids with fusiform cerata arranged in rows or on elevated brackets or cushions. Propodial tentacles are short to medium in length, the oral tentacles are long, and the rhinophores are long and smooth but can appear crinkled. The majority of species feeds on hydroids but species belonging to the genus Phestilla feed on corals. Cuthona sibogae (Bergh, 1905) Size: 25 mm Diagnostic characteristics: The body and cerata are pale pink or lilac deepening to magenta towards the ends of the cerata, tentacles, and tip of the

JH

Cuthona sibogae On Sertularella

JH

Species accounts 267

Suborder Aeolidina

Family Tergipedidae

Cuthona sibogae

JH

tail. The long fusiform cerata additionally are capped in dense yellow orange pigment. The smooth rhinophores, long oral tentacles, and recurved propodial tentacles are deep magenta with no yellow. Biology and ecology: Feeds on the thecate hydroids belonging to the genus Sertularella. Occurrence: Several photographic records from southern Egypt. Geographical distribution: First Red Sea records; Cuthona sibogae is found in South Africa and the West Pacific, up to 35 mm (Miller & Willan 2005). Phestilla lugubris (Bergh, 1870) Size: 50 mm Diagnostic characteristics: The translucent body and cerata have opaque white speckling. The cerata are long and knobbly, and appear to surround the head completely. They are capped in white and have small white patches along their lengths. The long rhinophores are a little crumpled but straighter than the cerata, with white pigment concentrated towards the distal ends. Biology and ecology: Feeds on the coral Porites; found in shallow waters. Occurrence: Single specimen record from the northern Red Sea (Yonow 2000); no further records. Geographical distribution: First Red Sea record, otherwise recorded in the tropical Indo-Pacific. 268 Species accounts

Suborder Aeolidina

Phestilla lugubris

Family Tergipedidae

JH

Phestilla melanobrachia Bergh, 1874 Size: 35 mm Diagnostic characteristics: The translucent pale orange body has many short plump cerata which vary in colour depending upon the diet, but always with an orange or white tip. Colour varies from bright to dull orange, brown, and black. A central band along the body is clear of cerata. The rhinophores and oral tentacles are both long and translucent orange deepening in colour towards the tips.

Phestilla melanobrachia

JK

NY

Species accounts 269

Suborder Aeolidina

Family Tergipedidae Biology and ecology: A number of differently coloured Phestilla melanobrachia can be seen to be feeding on the coral Tubastrea, which have retracted their polyps in response (upper left of photograph). Another specimen is crawling towards the prey colony, probably sensing both the presence of the coral as well as that of feeding conspecifics. The trailing mechanism is clearly seen in the lower photograph, where a number of aeolids are all following one another; three were head to tail. Although the coral colony was dead, they all filed past, presumably following the chemical trail left by the first animal. Occurrence: Probably common, wherever Tubastrea is found; there are a few photographic records from the Red Sea. Geographical distribution: Widespread throughout the Indo-Pacific.

JK

Phestilla melanobrachia

270 Species accounts

NY

NY

“Incertae Sedis” Cephalaspidea

“Incertae Sedis” This brief section contains photographs of single individuals taken in the Red Sea and Gulf of Eilat. These names are not definitive, and we must await collections of material before they can be identified further. However, this is a useful exercise in demonstrating the variety of unknown and undescribed species from the Red Sea, of which there are sure to be many more. I hope this book will encourage scientists, divers, and photographers to contribute towards filling the gaps in our of knowledge of these wonderful marine invertebrates.

Aglajid 12 mm

Chelidonura sp. Eilat,

Chelidonura sp. 10 mm

Eilat, B&SK

Chelidonura sp. 13 mm

Eilat, B&SK

Eilat, B&SK

OL

Chelidonura sp. Eilat,

“Incertae sedis” 271

B&SK

“Incertae Sedis”

Cephalaspidea / Sacoglossa

Philinopsis sp.

Philinopsis sp. 15 mm

Eilat, B&SK

Noalda sp. 5 mm

Philinopsis sp. 20 mm

Eilat, B&SK

Eilat, OL

Eilat, B&SK

Elysiella sp. Eilat,

OL

272 “Incertae sedis”

Elysia cf. tomentosa Jeddah,

GS

“Incertae Sedis”

Doridina

Goniodorid 15 mm

Eilat, B&SK

Gymnodoris sp. 15 mm

Eilat, B&SK

Analogium sp. 13 mm

Hallaxa sp. Eilat,

Hallaxa sp. 10 mm

Eilat, B&SK

Discodoris sp.

“Incertae sedis” 273

Eilat, B&SK

OL

Jeddah, JK

“Incertae Sedis”

Doridina

Discodoris ? mauritiana 35 mm

Egypt, JH

Discodoris ? mauritiana 35 mm, ventral

Egypt, JH

Jorunna sp. 20 mm

Eilat, B&SK

Atagema sp. 22 mm

Eilat, B&SK

Jorunna sp.

Jeddah, JK

Atagema sp.

Jeddah, JK

Sclerodoris cf. rugosa

Jeddah, JK

Hoplodoris cf. armata Jeddah,

274 “Incertae sedis”

JK

“Incertae Sedis”

Doridina / Arminina

Chromodoris sp.

Suakin, CT

Chromodoris sp.

Eilat, OL

Phyllidiopsis sp.

Jeddah, H. Schmid

Chromodoris sp., ventral

Noumea sp. 7 mm

Janolus sp. Jeddah,

“Incertae sedis” 275

Suakin, CT

Eilat, B&SK

JK

“Incertae Sedis”

Armina sp. 25 mm

Arminina / Aeolidina

Eilat, B&SK

Armina cf. major 100 mm

Eilat, B&SK

Eilat, B&SK

Cuthona sp.

Eilat, B&SK

Cerberilla sp. 45 mm

Eubranchus sp.

Suakin, CT

Eubranchus sp. 5 mm

276 “Incertae sedis”

Egypt, JH

GLOSSAR

Y

alcyonarian

colonial soft coral which is solid and fleshy, or octocoral; class Anthozoa, phylum Cnidaria

anemone

solitary or gregarious polyps (q.v.) with a column and a series of tentacles, or hexacorals; class Anthozoa, phylum Cnidaria

anterior

front end of an animal

aperture

opening of the shell

apex

peak or top of a shell

arborescent

repeatedly branched, tree-like

ascidian

also known as sea squirts and tunicates; sessile filter feeding animals belonging to the class Ascidiacea, phylum Chordata; can be solitary or colonial

autotomize

deliberate loss of a body part by an animal, usually the mantle, often when disturbed as a defense mechanism

benthic

an organism living on the sea bed

bilateral symmetry

a condition in which the animal can be divided into two equal halves by a line drawn down the midline

bipinnate

secondary branches bearing leaflets, paired or alternate from a main stem; referring to gills

branchia

respiratory organs or gills

bryozoan

also known as moss animals, lace corals, or sea mats; sessile filter feeding animals with colonies forming encrusting sheets or erect growths; phylum Bryozoa

carnivorous

animals which feed on other animals

cerata

finger-like extensions of the dorsum, containing ramifications of the digestive gland and functioning in digestion, respiration, and defense (see papillae) (sing. ceras)

chloroplast

green photosynthesizing organelles in algae and seaweeds

Glossary 277

columella

the main axis of a gastropod shell, which is usually visible on the left side of the aperture; may be sculptured or have teeth

cnidarian

belonging to anemones, zooanthids, corals and other species of the phylum Cnidaria

cnidosac

small cavity at the end of the cerata (q.v.) in aeolids used to store intact nematocysts (q.v.) (fig. 39)

cryptic

body camouflaged to resemble the habitat

cryptobranch

dorid nudibranchs which have pockets into which the gills and rhinophores retract

ctenidia

gills (sing. ctenidium)

digitiform

finger-like (usually with reference to cerata, rhinophores, or papillae)

distal

area furthest away from the attachment point

dorsum

upper surface or back (= notum)

endemic

species which live in a restricted zoogeographical area, in this case the Red Sea

epifauna

animals (fauna) which live on something else such as rocks, sea weeds, harbour pilings, ship hulls, etc.

epiflora

plants (flora) which live on something else such as rocks, sea weeds, harbour pilings, ship hulls, etc.

epiphyte

plant or animal growing on plants such as sea weeds and sea grasses

gonad

reproductive organs of most invertebrate animals, comprising the production of eggs and sperm

gorgonian

also known as sea fans; arborescent colony with a horny skeleton and filter-feeding polyps (q.v.); class Anthozoa, phylum Cnidaria

herbivorous

animals which feed on algae and seaweeds

hermaphrodite

both male and female reproductive organs functioning in a single individual, producing both eggs and sperm

hydroid

solitary or colonial filter feeders with sessile polyps (q.v.) and free-swimming medusa generations in the life cycle; class Hydrozoa, phylum Cnidaria 278 Glossary

hyponotum

the underside of the mantle extending beyond the foot

invertebrate

in the marine habitat, all animals without a spine, such as snails, starfish, sea urchins, ascidians, hydroids, anemones, jellyfish, corals, etc.

lamellae

often with regards to gills: secondary leaflets located ventrally in the Phyllidiidae and Armina, or the smallest units of the dorsal gills

lamellate

referring to the rings on the rhinophores (= perfoliate)

mantle

layer on the back of most gastropods, primarily responsible for shell secretion; in opisthobranchs refers to the upper surface

metapodium (al)

posterior or tail end of the foot

nematocysts

stinging cells found in animals such as jellyfish, corals, sea anemones, and hydroids; stored and used by some nudibranchs

notum

the back or upper surface of opisthobranchs (= dorsum)

ocelli

eye-like spots of one colour with a ring of another colour around them (sing. ocellus)

oophagous

animals which feed on the eggs of other animals

operculum

horny plate which shuts the opening of the shell, and is attached to the foot; present in very primitive opisthobranchs and all prosobranchs

oral veil

frontal membrane extending from the head above the mouth

oral tentacles

paired elongated projections on the head near the mouth

ovotestis

reproductive organs (= gonad)

papilla

extensions of the body, lacking digestive gland (see cerata) (pl. papillae)

parapodia

fleshy lobes either side of the body of some opisthobranchs such as Hydatina, Aplysia, and Elysia (sing. parapodium)

pelagic

oceanic, floating or swimming in wide open seas, passive movement with sea currents and winds

pennatulacean

known as sea pens; elongate, rod- or feather-like octocorals living in soft substrata; class Anthozoa, phylum Cnidaria

perfoliate

referring to lamellae on the rhinophores (= lamellate)

Glossary 279

periostracum

external chitinous covering of the shell of some molluscs

phanerobranch

condition of dorid nudibranchs in which the dorsal gills cannot be retracted into a pocket or sheath

pinnate

single leaflets branching from a single main stem of a gill, like a feather

polychaete

bristle worms, mostly marine and free-living; class Polychaeta, phylum Annelida

polyclad

marine flatworms; class Turbellaria, phylum Platyhelminthes

polyp

basic structure of the phylum Cnidaria; sessile, more or less cylindrical, with a mouth surrounded by tentacles at the free end

posterior

tail end of an animal

predator

an animal which feeds on another

prey

an animal upon which another animal feeds

propodium (al)

front end of the foot

proximal

area nearest the point of attachment

pseudobranch

from the Greek for “false gill”, referring to a small structure present at the base of the cerata in some species of Doto

pustule

small rounded pimples or swellings which are present on the mantle and foot of some opisthobranchs (= tubercle)

radula

cuticularized structure used for feeding in most molluscs; bears one or many rows of tiny teeth which are of use in identifying species

reticulate

lines which join or intersect to form a lattice-like pattern

rhinophores

paired chemosensory tentacles on the head of most opisthobranchs

sessile

living attached to the substrate

sheath

sleeve into which the rhinophores and/or gills retract

siphon

tube-like extension of the mantle for the passage of water currents

siphonophore

floating or swimming hydroid with many different units functioning as one colonial organism; individual units are specialized for floating, feeding, reproduction, etc.; class Hydrozoa, phylum Cnidaria 280 Glossary

spatulate

expanded head region of some species of Hypselodoris and Ceratosoma, for example

spawn

mass or strings of eggs imbedded in a transparent mucous jelly

spicule

calcareous structure in many shapes, usually from sponges, incorporated in the mantle tissue of some nudibranchs and used for mechanical protection

sponge

colonial invertebrates belonging to the phylum Porifera; filter feeders growing on many substrates and the food of many nudibranchs

tentacles

paired sensory projections near the head and mouth

trapezoidal

referring to the oral veil of pleurobranchs, which is narrower at the attached side than the free distal side

tripinnate

main stem with primary and secondary branches (gills)

tubercle

short rounded projections, usually on the mantle (= pustule)

ventral

lower part, underneath; in opisthobranchs usually the foot, head, and underside of the mantle

zooxanthellae

single-celled plants living in the tissue of other organisms but continuing to photosynthesize

Glossary 281

ACKNOWLEDGEMENTS

As every author worries, it is impossible to thank everyone, and the fear of omission is great. This book has been ‘in prep’ for nearly 15 years, and my first thanks must go to Doreen and Issam Sharabati for inviting me on their wonderful expedition which circumnavigated the Red Sea during the summer months of 1983. My addiction to opisthobranchs began on an undergraduate field course in 1981, where the only species I could successfully identify were the opisthobranchs; for this I sincerely thank the author of the key, Tom Thompson, who later became my mentor and friend. He died tragically in 1990, and I hope he would be pleased with my efforts: I dedicate this book to him. Many people have helped in different spheres: for enjoyable and productive discussions on many topics, I am grateful to Ingo Burghardt, Dan Forman, Binyamin Koretz, and Richard Willan. A big thank you to Anna Ratcliffe and Alan Cutliffe for cheerfully scanning my never-ending supply of slides, and to my ‘search’ assistants Matt and Imogen Elwick and Jenni Aazem for finding some of the illustrations used in the introduction. Michael Miller is gratefully acknowledged for a copy of Vayssiére (1912) as are my friends for patiently submitting to endless conversations about sea slugs. Most of all, I thank my father for teaching me to swim before I could walk, for my first mask and snorkel at the age of eight when in the Arabian Gulf, and for all the snorkelling holidays, not only in the Red Sea. I would like to thank my publisher for his enthusiasm from the very start of this project, and the designer for producing a book beyond my wildest dreams. Any mistakes are mine. Finally, this book would have been very dull indeed without the generosity of the photographers, and I cannot thank them enough: I hope they are pleased with the results. The majority of photographs were taken by the following, and their photos are designated by their initials: B&SK JH JK OL PK

Binyamin & Shulamit Koretz Johann Hinterkircher Jürgen Kuchinke Oren Lederman Pam Kemp

282 Acknowledgements

All other photos have the photographers’ names or initials underneath, and I thank them most sincerely for making a tedious scientific tome so much more interesting and attractive. Arik Diamant Alex Double Amir Stern Angel Valdés Bernard Picton Chris Todd (CT) Doreen Sharabati (DPS) Ewald Lieske Gunnar Bemert Götz Reinicke Gordon Smith (GS) Helmut Göthel Hagen Schmid Helmut Schuhmacher (HS) Hans Sjöholm Jacob Dafni Johnny Walker Kevin Metcalfe Mohamed Habib Moshira Hassan Michel Jozon Michael Schrödl Nechama Ben-Eliahu Nicole Kenward Philibert Bidgrain Razi Cohen Schmuel Schrenkel Tuula Metcalfe Thomas Paulus (TP) Thilo Wanner Viviana Farstey Volker Neumann

Acknowledgements 283

BIBLIOGRAPHY

Aboul-Ela, I. A. 1959. On the food of nudibranchs. Biological Bulletin 117 (3): 439–442. Abraham, P. S. 1877. Revision of the anthobranchiate nudibranchiate Mollusca, with descriptions or notices of forty-one hitherto undescribed species. Proceedings of the Zoological Society, London: 196–269. Audouin, J. V. 1826. Explications sommaire des planches de Mollusques de l’Egypte et de la Syrie publiées par J. C. Savigny... Histoire Naturelle, Animaux Invertébrés 1 (4): 7–56. Avila, C., Kelman, D., Kashman, Y. & Benayahu, Y. 1999. An association between a dendronotid nudibranch (Mollusca, Opisthobranchia) and a soft coral (Octocorallia, Alcyonaria) from the Red Sea. Journal of Natural History 33 (10): 1433–1449. Baba, K. 1949. Opisthobranchia of Sagami Bay. Iwanami Shoten, Tokyo, Japan. 194 pp. Baba, K. 1959. The genus Petalifera and a new species, P. ramosa, from Japan. Publications of the Seto marine Biological Laboratory 7(3): 43–44. Barash, A. & Danin, Z. 1972 . The Indo-Pacific species of Mollusca in the Mediterranean and notes on a collection from the Suez Canal. Israel Journal of Zoology 21: 301–376. Barash, A. & Danin, Z. 1980 . Notes on the Aplysiidae from the Red Sea (Mollusca, Gastropoda, Opisthobranchia). Levantina 28: 325–329. Barash, A. & Zeniper, Z. 1993. Notes on Opisthobranchia from the Red Sea. Conchiglia 25 (269): 18–23. Barash, A. & Zeniper, Z. 1994. Notes on Opisthobranchia from the Red Sea. Conchiglia 26: 7–9. Behrens, D. W. & Hermosillo, A. 2005. Eastern Pacific Nudibranchs. A Guide to the Opisthobranchs fr om Alaska to Central America. Sea Challengers, California, U.S.A. 137 pp. Bemert, G. & Ormond, R. 1971. Red Sea Coral Reefs. Routledge & Keegan Paul, London, U.K. 192 pp. Bergh, L. S. R. 1905. Die Opisthobranchiata der Siboga Expedition. Monographie 50: 1–248. Bergh, L. S. R. 1908. Malacologische Untersuchungen. In: Reisen im Archipel der Philippinen (ed. C. G. Semper) 9: 148–163. Bouchet, P. & Danrigal, F. 1982. Napoleon’s Egyptian campaign (1798–1801) and the Savigny collection of shells. The Nautilus 96 (1): 9–24. Brodie, G. D., Willan, R. C. & Collins, J. D. 1997. Taxonomy and occurrence of Dendrodoris nigra and Dendrodoris fumata (Nudibranchia: Dendrodorididae) in the Indo-West Pacific region. Journal of Molluscan Studies 63: 407–423.

284 Bibliography

Campbell, A. C. 1987. Echinoderms of the Red Sea. In: Red Sea (eds. A. J. Edwards & S. M. Head). Pergamon Press, Oxford, U.K. pp 215–232. Cervera, J. L., Garcia Gomez, J. C. & Megina, C . 1996. Redescription of Nembrotha megalocera Yonow, 1990 (Gastropoda: Nudibranchia: Polyceratidae) from the Red Sea. The Veliger 39 (1): 55–59. Collingwood, C. 1881. On some new species of nudibranchiate Mollusca from the Eastern seas. Transactions of the Linnean Society, London: Zoology (series 2) 2 (2): 128–140. Crossland, C. 1937. Four molluscs rarely recorded from the Red Sea. Nature 139: 31. Dayrat, B. 2006. A taxonomic revision of Paradoris sea slugs (Mollusca, Gastropoda, Nudibranchia, Doridina). Zoological Journal of the Linnean Society 147: 125–238. De Ventier, L., Turck, E., Al-Sheikh, K. & De’ath, G . 2000. Coral communities of the central and northern Saudi Arabian Red Sea. Fauna of Arabia 18: 23–66. Dekker, H. & Or lin, Z. 2000. Check-list of Red Sea Mollusca. Spirula 47 (supplement): 1–46. Drouin, J.-M. 2001. Analogies et contrastes entre l’éxpedition d’Egypte et le voyage d’Humboldt et Bonpland. História, Ciêcias, Saúde–Manguinlos 8 (supplement): 839–861. Eales, N. B. 1938. A systematic and anatomical account of the Opisthobranchia. Scientific Reports of the John Murray Expedition 5: 77–122. Eales, N. B. 1960. Revision of the world species of Aplysia (Gastropods, Opisthobranchia). Bulletin of the British Museum (Natural Histor y) Zoology 5: 267–404. Eales, N. B. 1979. The Aplysiidae from the Red Sea. Argamon, Israel Journal of Malacology 7 (1): 1–19. Eales, N. B. & Engel, H. 1935 . The genus Bursatella de Blainville. Proceedings of the Malacological Society 21 (5): 279–303. Edmunds, M. 1968. On the swimming and defensive response of Hexabranchus marginatus (Mollusca, Nudibranchia). Journal of the Linnean Society, Zoology 47 (313): 425–429. Edmunds, M. 1969. Opisthobranchiate Mollusca from Tanzania. I. Eolidacea (Eubranchidae and Aeolidiidae). Proceeding of the Malacological Society of London 38: 451–469. Ehrenberg, C. G. 1828, 1831. Decas 1. Mollusca. Symbolae physicae, seu icones et descriptiones animalium e vertebraterum sepositis insectis quae ex itiner e per Africam Borealum et Asiam Occidentalem–novae ant illustratae redierunt (pages unnumbered) (plates 1828, text 1831). Eisenberg, J. M. 1981. A collector’s guide to Seashells of the World. McGraw Hill Book Co., New York, U.S.A. 239 pp. Eliot, C. N. E. 1908. Reports on the marine biology of the Sudanese Red Sea. XI. Notes on a collection of nudibranchs from the Red Sea. Journal Linnean Society, Zoology 31: 86–122.

Bibliography 285

Eliot, C. N. E. 1911. Chromodorids from the Red Sea, collected and figured by Mr. Cyril Crossland. Proceedings of the Zoological Society, London 1068–1072. Engel, E. & van Eeken, C. J. 1962. Red Sea Opisthobranchia from the coast of Israel and Sinai. Bulletin of Sea Fisheries Research Station, Israel 30: 15–34. Fahrner, A. & Schrödl, M. 2000a. Description of Phyllidia schupporum, a new nudibranch species from the northern Red Sea. Spixiana 23 (1): 55–60. Fahrner, A. & Schrödl, M. 2000b. Taxonomic revision of the common Indo-West Pacific nudibranch Phyllidia varicosa Lamarck, 1801. The Veliger 43 (2): 164–171. Fahrner, A. & Schrödl, M. 2000c. Redescription of Phyllidiopsis sinaiensis (Yonow, 1988) (Nudibranchia: Doridoidea: Phyllidiidae), with a review of the Red Sea Phyllidiidae. Journal of Molluscan Studies 66: 467–476. Farran, G. P. 1905. Report on the opisthobranchiate Mollusca collected by Prof. Herdman at Ceylon, in 1902. In: Report to the Gover nment of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar (ed. W. A. Herdman) 3, supplementary report 21: 329–364. Fishelsen, L. 1971. Ecology and distribution of the benthic fauna in shallow waters of the Red Sea. Marine Biology 10: 113–133. Foin, T. C. & Ruebush, L. P. 1969. Cypraeidae of the Red Sea at Massawa, Ethiopia, with a zoogeographical analysis based on the Schilders’ regional lists. The Veliger 12 (1): 201–206. Forskål, P. 1775, 1776. Vermes. I. Mollusca. In: Descriptiones animalium, avium ...vernum quae in itiner e orientali observavit Petrus Forsskål (ed. C. Niebuhr). Hauniae pp 99–140 (1775) and pp 1–13 (1776). Gat, G. & Fainzilber, M. 1984. A flock of Spanish dancers. Levantina 53: 164. Gohar, H. A. F. 1948. A description and some biological studies of a new alcyonarian species Clavularia hamra Gohar. Publications of the marine biological Station Ghardaqa 6: 3–33. Gohar, H. A. F. & Aboul-Ela, I. A. 1957a . The development of Berthellina citrina. Publications of the marine biological Station Ghardaqa 9: 69–84. Gohar, H. A. F. & Aboul-Ela, I. A. 1957b . On a new nudibranch Phylodesmium xenia. Publications of the marine biological Station Ghardaqa 9: 131–144. Gohar, H. A. F. & Aboul-Ela, I. A. 1957c . The development of three chromodorids. Publications of the marine biological Station Ghardaqa 9: 203–228. Gohar, H. A. F. & Aboul-Ela, I. A. 1959 . On the biology and development of three nudibranchs. Publications of the marine biological Station Ghardaqa 10: 41–62. Gohar, H. A. F. & Soliman, G. N. 1963. The biology and development of Hexabranchus sanguineus (Rüppell & Leuckart) (Gastropoda, Nudibranchiata). Publications of the marine biological Station Ghardaqa 12: 219–247.

286 Bibliography

Gohar, H. A. F. & Soliman, G. N. 1967a. The biology and development of Dendrodoris (Doriopsis) fumata (Rüppell & Leuckart) (Gastropoda, Nudibranchiata). Publications of the marine biological Station Ghardaqa 14: 31–54. Gohar, H. A. F. & Soliman, G. N. 1967b. The biology and development of Chromodoris inornata Pease (Gastropoda, Nudibranchiata). Publications of the marine biological Station Ghardaqa 14: 79–94. Gohar, H. A. F. & Soliman, G. N. 1967c. The biology and development of the nudibranch Chromodoris tinctoria (Rüppell & Leuckart) (Gastropoda, Nudibranchiata). Publications of the marine biological Station Ghardaqa 14: 95–108. Gohar, H. A. F. & Soliman, G. N. 1967d. The direct development of the nudibranch Casella obsoleta (Rüppell & Leuckart) (Gastropoda, Nudibranchiata). Publications of the marine biological Station Ghardaqa 14: 149–166. Gohar, H. A. F. & Soliman, G. N. 1967e. The biology and development of Asteronotus cespitosa (van Hasselt) (Gastropoda, Nudibranchiata). Publications of the marine biological Station Ghardaqa 14: 177–195. Gohar, H. A. F. & Soliman, G. N. 1967f. The biology and development of Discodoris concinna (Alder & Hancock) (Gastropoda, Nudibranchiata). Publications of the marine biological Station Ghardaqa 14: 197–214. Gohar, H. A. F. & Soliman, G. N. 1967g. On two rare nudibranchs of the genus Trippa Bergh (Gastropoda, Nudibranchiata). Publications of the marine biological Station Ghardaqa 14: 269–293. Gosliner, T. M. 1987. Nudibranchs of Southern Africa. E.J. Brill, Leiden, Netherlands. 136 pp. Gosliner, T. M. & Behrens, D. W. 2000. Two new species of Chromodorididae (Mollusca: Nudibranchia) from the tropical Indo-Pacific, with a redescription of Hypselodoris dollfusi (Pruvot-Fol, 1933). Proceedings of the California Academy of Sciences 52 (10): 111–124. Gosliner, T. M., Behrens, D. W. & Willams, G. C. 1996. Coral Reef Animals of the Indo-Pacific. Sea Challengers, California, U.S.A. pp 150–178. Gunthorpe, L. & Cameron, A. M. 1987. Bioactive properties of extracts from Australian dorid nudibranchs. Marine Biology 94: 39–43. Haas, F. 1920. Opisthobranchier aus verschiedenen warmen Meeren. Archiv für Mollusken 52 (3): 138–142. Hadfield, M. & Switzer-Dunlap, M. 1984. Opisthobranchs. In: The Mollusca (eds. A. S. Tompa et al.). Academic Press, New York, U.S.A. Vol. 7, pp 209–350. Hägg, R. 1904. Two new opisthobranchiate Mollusca from the Red Sea. Results of the Swedish Zoological Expedition to Egypt and the W hite Nile, 1901, under L.A. Jagerskiold. Upsala, Sweden. 16 pp. Head, S. M. 1987. Introduction. In: Red Sea (eds. A. J. Edwards & S. M. Head). Pergamon Press, Oxford, U.K. pp 1–21. Heller, J. & Thompson T. E. 1983. Opisthobranch molluscs of the Sudanese Red Sea. Zoological Journal of the Linnean Society 78: 317–348.

Bibliography 287

Hughes, R. N. 1977. The biota of reef flats and limestone cliffs near Jeddah, Saudi Arabia. Journal of Natural History 11: 77–96. I.C.Z.N. 1999. International Code of Zoological Nomenclature. Fourth Edition. The Natural History Museum, London, U.K. 306 pp. Issel, A. 1869. Malacologia del Mar Rosso. Memoria letta al cogresso dei naturalisti Italiani in Vicenza nel 1868. Pisa, Italy. 387 pp. Johnson, R. & Valdés, A. 2001. The Hypselodoris infucata, H. obscura and H. saintvincentius species complex (Mollusca, Nudibranchia, Chromodorididae), with remarks on the genus Brachyclanis Ehrenberg, 1831. Journal of Natural History 35: 1371–1398. Jousseaume, F. 1888. Descriptions des mollusques receuillis par M. le Dr. Faurot dans la Mer Rouge et le Golfe d’Aden. Memoires de la Societé Zoologique de France 1: 165–223. Klausewitz, W. 2002. Frankfurt versus Berlin: the Red Sea explorers Wilhelm Hemprich, Christian Ehrenberg, and Eduard Rüppell. Zoology in the Middle East 27: 7–12. Lamy, E. 1938. Mollusca Testacea (Molluscs collected by Dollfus) from Gulfs of Suez and Aqaba. Memoires de l’Institut d’Egypte 37: 1–89. Lüttmann, K., Anthes, N., D’Souza, T. G., Riss, S. & Michiels, N. K. 2006. Population size estimate of a reef flat aggregation of Chromodoris annulata (Opisthobranchia, Chromodorididae). Journal of Molluscan Studies 72: 214–216. Maempel, G. Z. 1992. The Arabian Voyage of 1761–67 and Malta: Forsskål and his contribution to the study of local natural history. Proceedings of History Week 1992: 35–76. Marbach, A. & Tsurnamal, M. 1973. On the biology of Berthellina citrina (Gastropoda, Opisthobranchia) and its defensive acid secretion. Marine Biology 21: 331–339. Marcus, E. 1980. Review of western Atlantic Elysiidae (Opisthobranchia Ascoglossa) with a description of a new Elysia species. Bulletin of Marine Science 30: 54–79. Marcus, E. & Marcus, E. 1960 . Opisthobranchia aus dem Roten Meer und von den Maldivien. Abh.Math. Nat. Kl. Akad. W iss. Mainz 12: 1–64 (873–939). Marshall, J. & Willan, R. C. 1999. Nudibranchs of Heron Island, Great Barrier Reef. A Sur vey of the Opisthobranchia (Sea Slugs) of Heron and Wistari Reefs. Backhuys Publishers, Leiden, The Netherlands. 257 pp. Mastaller, M. 1978. The marine molluscan assemblages of Port Sudan, Red Sea. Zoologische Mededelingen 53 (13): 117–144. Mastaller, M. 1979. Beitrage zur faunistik und ökologie der mollusken und echinodermen in den korallenriffen bei Aqaba, Rotes Meer. Ph.D. dissertation, Abt. Biologie and der Ruhr-Univ. Bochum. 344 pp. Mienis, H. K. 1973. A bivalved gastropod from Elat. Argamon, Israel Journal of Malacology 4 (1): 18.

288 Bibliography

Mienis, H. K. 1984. Chromodoris vicina Eliot, 1904 from the Red Sea. Levantina 50: 581. Mienis, H. K. & Gat, G . 1986. A note concerning Flabellina rubrolineata (O’Donoghue, 1929). Levantina 64: 683–684. Miller, M. C. & Willan, R. C. 2005. A redescription and taxonomic reappraisal of the Indo-Pacific aeolid nudibranch Trinchesia sibogae (Bergh, 1905) (Gastropoda, Opisthobranchia). Vita Malacologica 3: 69–75. Nebelsick, J. H. 1996. Biodiversity of shallow-water Red Sea echinoids: implications for the fossil record. Journal of the Marine Biological Association, U.K. 76 (1): 185–194. Odhner, N. H. 1936. Nudibranchia Dendronotacea: a revision of the system. Memoires du Museum Royale d’Histoire Naturelle de Belgique (2) 3: 1057–1128. O’Donoghue, C. H. 1929. Report on the Opisthobranchiata, with a comparison of the nudibranch faunas of the Red Sea and of the Mediterranean. Cambridge Expedition to the Suez Canal 1924. Transactions of the Zoological Society, London 22 (38): 713–841. Ormond, R. F. G. & Edwards, A. J. 1987. Red Sea Fishes. In: Red Sea (eds. A. J. Edwards & S. M. Head). Pergamon Press, Oxford, U.K. pp 251–287. Pagenstecher, H. A. 1877 . Mollusca. In: Zoologische Ergeb. einer im Auftrage der Kon. Adad. W iss. Berlin Reise Kust. Rothen Meeres (ed. R. Kossman) 1 (2): 1–666. Pallary, P. 1926. Explication des planches de J.C. Savigny. Memoires de l’Institut d’Eg ypte 11: 139 pp. Pallary, P. 1931. Marie Jules-César Savigny. Sa vie et son oeuvre. Première partie. Memoires de l’Institut d’Egypte 17: 106 pp. Pawlik, J. R., Kernan, M. R., Molinski, T. F., Harper, M. K. & Faulkner, D. J. 1988. Defensive chemicals of the Spanish dancer nudibranch Hexabranchus sanguineus and its egg ribbons: macrolides derived from a sponge diet. Journal of Experimental Marine Biology and Ecology 119: 99–109. Perrone, A. S. & Doneddu, M. 2001 . Pattern convergence in Nudibranchia of the genus Chromodoris Alder & Hancock, 1855 (Opisthobranchia) from the Red Sea and Polycladida (Platyhelminthes) from the Mediterranean Sea. Vita Marina 47 (4): 121–130. Plaut, I., Borut, A. & Spira, M. E. 1995 . Growth and metamorphosis of Aplysia oculifera larvae in laboratory culture. Marine Biology 122: 425–430. Plaut, I., Borut, A. & Spira, M. E. 1998 . Seasonal cycle and population dynamics of the sea hare Aplysia oculifera in the northern Gulf of Eilat (Aqaba), Red Sea. Journal of Molluscan Studies 64: 239–247. Pruvot-Fol, A. 1933. Mission Robert Ph. Dollfus en Egypte. Opisthobranchiata. Memoires de l’Institut d’Egypte 21: 89–159. Pruvot-Fol, A. 1951. Révision du genre Glossodoris Ehrenberg. Journal de Conchyliologie 91 (3): 76–132.

Bibliography 289

Randall, J. E. 2003. A review of cardinalfishes (Perciformes: Apogonidae) of the Red Sea. The South African Institute for Aquatic Biodi versity 1 (1): 1–48. Reinicke, G. B. 1997. Xeniidae (Colenterata: Octocorallia) of the Red Sea, with descriptions of six new species of Xenia. Fauna of Saudi Arabia 16: 5–62. Righton, D., Kemp, J. & Ormond, R. 1996. Biogeography, community structure, and diversity of Red Sea and western Indian Ocean butterfly fishes. Journal of the Marine Biological Association, U.K. 76 (1): 223– 228. Rochebrune, A. T. 1894. Diagnoses de formes nouvelles appartenant au genre Ceratosoma. Le Naturaliste, Paris (2) 16: 168. Rudman, W. B. 1978. The dorid opisthobranch genera Halgerda Bergh and Sclerodoris Eliot from the IndoWest Pacific. Zoological Journal Linnean Society 62: 59-88. Rudman, W. B. 1983. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: Chromodoris splendida, C. aspersa, and Hypselodoris placida colour groups. Zoological Journal Linnean Society 78 (2): 105–173. Rudman, W. B. 1984. The Chromodorididae (Opisthobranchia, Mollusca) of the Indo-West Pacific: a review of the genera. Zoological Journal Linnean Society 81: 115–273. Rudman, W. B. 1985. The Chromodorididae (Opisthobranchia, Mollusca) of the Indo-West Pacific: Chromodoris aureomarginata, C. verrieri and C. fidelis colour groups. Zoological Journal Linnean Society 83 (3): 241–299. Rudman, W. B. 1987a. The Chromodorididae (Opisthobranchia, Mollusca) of the Indo-West Pacific: Chromodoris epicuria, C. aureopurpurea, C. annulata, C. coi and Risbecia tryoni colour groups. Zoological Journal Linnean Society 90 (4): 305–407. Rudman, W. B. 1987b. The genus Trapania (Nudibranchia: Goniodorididae) in the Indo-West Pacific. Journal of Molluscan Studies 53: 189–212. Rudman, W. B. 1988. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: the genus Ceratosoma J.E. Gray. Zoological Journal Linnean Society 93 (2): 133–185. Rudman, W. B. & Dar vell, B. W. 1990. Opisthobranchs of Hong Kong, part I: Goniodorididae, Onchidorididae, Triophidae, Gymnodorididae, Chromodorididae (Nudibranchia). Asian Marine Biology 7: 31–79. Rüppell, W. P. E. S. & Leuckart, F. S. 1828, 1830. Mollusca. In: Atlas zu der Reise im nordlichen Afrika von Eduard Rüppell. Zool. Neue wirbellose Thier e des Rothen Meer es, pp 15–22 and Tab. 4, 5 (1828), pp 23–47 and Tab. 7–11 (1830). Savigny, J. C. 1817. Description de l’Egypte, ou receuil des observations et des recherches qui ont été faites en Egypte pendant l’Expédition de l’armée française, publiée par ordre du gouvernement. Histoire Naturelle. Imprimerie Royale, Paris. Planches, vol. II.

290 Bibliography

Schmid, H. 1996. Desert Sea. Fauna of the Saudi Arabian Red Sea Coast . Paramount Printing Co. Ltd., Hong Kong. 201 pp. Schmid, H. 2006. Out of the Blue. Underwater realm of the Red Sea. Heidelberger Reprographic, Andreas Grosch GmbH, Germany. 165 pp. Schrödl, M. 1999. Glossodoris charlottae, a new chromodorid nudibranch from the Red Sea (Gastropoda, Opisthobranchia). Vita Marina 46 (3–4): 89–94. Schuhmacher, H. 1973a. Das Kommensalische Verhaltnis zwischen Periclimenes imperator und Hexabranchus sanguineus. Marine Biology 22: 355–360. Schuhmacher, H. 1973b. Notes on occurrence, feeding, and swimming behaviour of Notarchus indicus and Melibe bucephala at Elat, Red Sea (Mollusca: Opisthobranchia). Israel Journal of Zoology 22: 13–25. Schupp, P., Eder, C., Proksch, P., Wray, V., Schneider, B., Herderich, M. & Paul, V. 1999. Staurosporine derivatives from the ascidian Eudistoma toealensis and its predatory flatworm Pseudoceros sp. Journal of Natural Products 62: 959–962. Seager, J. R. 1982. Population dynamics of the Antarctic opisthobranch Philine gibba Strebel. Journal of Experimental Marine Biology and Ecology 60: 163–175. Sharabati, D. P. 1981. Saudi Arabian Seashells. Royal Smeets Offset, B.V. Weert, Netherlands. 119 pp. Sharabati, D. P. 1984. Red Sea Shells. K.P.I. Ltd., London, U.K. 128 pp. Soliman, G. N. 1978. The redescription, reproduction and development of the dorid nudibranch Platydoris scabra (Cuvier) from the north-western Red Sea. Journal of Molluscan Studies 44: 151–165. Soliman, G. N. 1980. On the nudibranch Sebadoris crosslandi (Eliot) from the north-western Red Sea. Journal of Molluscan Studies 46: 227–238. Soliman, G. N. 1986. The reproduction and development of the dorid nudibranch Phyllidia varicosa Lamarck, 1801 from the northern Red Sea. Proceedings of the 8th Inter national Malacological Congress, Budapest 1983: 255–260. Sommerfeldt, N. & Schrödl, M. 2005. Microanatomy of Hedylopsis ballantinei, a new interstitial acochlidian gastropod from the Red Sea, and its significance for phylogeny. Journal of Molluscan Studies 71: 153–165. Strack, H. L. 1993. The Polyplacophora of the Red Sea. Journal of the Malacological Society, Australia 14: 1–40. Sturany, R. 1903. Gastropoden des Rothen Meeres. Denkschriften der Kaiser. Akad. W iss. Math.-Natur. Wien, Kl. Zoologische Ergebnisse 23: 210–283. Taylor, J. D. & Reid, D. G. 1984. The abundance and trophic classification of molluscs upon coral reefs in the Sudanese Red Sea. Journal of Natural History 18: 175–209.

Bibliography 291

Thompson, T. E. & McFarlane, I. D. 1967. Observations on a collection of Glaucus from the Gulf of Aden with a critical review of published records of Glaucidae (Gastropoda, Opisthobranchia). Proceedings of the Linnean Society, London 178 (2): 107–123. Tomlin, J. R. 1927. Report on the Mollusca (Amphineura, Gastropoda, Scaphopoda, Pelecypoda). Cambridge Expedition to the Suez Canal 1924. Transactions of the Zoological Society, London 22: 291–320. Valdés, A. 2002. Review of the genus Actinocyclus Ehrenberg, 1831 (Opisthobranchia: Doridoidea). The Veliger 45 (3): 193–202. Vayssière, A. 1898. Monographie de la famille des Pleurobranchidés. Annals des sciences natur elles (Zoologie) 8: 209–402. Vayssière, A. 1906. Recherches zoologiques et anatomiques sur les Opisthobranches de la Mer Rouge et du Golfe d’Aden. Parte 1. Annals de la Faculté des Sciences, Univ. Marseille 16: 1–72. Vayssière, A. 1911a. Sur les opisthobranches et sur les Marséniadés du Golfe de Tadjoura. Compte Rendu à l’Academie des Sciences, Paris 153: 1164–1166. Vayssière. A. 1911b. Note sur les Mollusques Nudibranches, Marséniadés, et Onchididés receuillis dans le Golfe d’Aden, à Djibouti, par M. Ch. Gravier, en 1904. Bulletin du Museum National d’Histoir e Naturelle, Paris 17: 442–443. Vayssière, A. 1912. Recherches zoologiques et anatomiques sur les Opisthobranches de la Mer Rouge et du Golfe d’Aden. Parte 2. Annals de la Faculté des Sciences, Univ. Marseille 20: 5–157. Wehe, T. & Fiege, D. 2002. Annotated checklist of the polychaete species of the seas surrounding the Arabian Peninsula: Red Sea, Gulf of Aden, Arabian Sea, Gulf of Oman, Arabian Gulf. Fauna of Arabia 19: 7–238. Wells, F. E. and Bryce, C. W. 1993. Sea Slugs of Western Australia. Western Australian Museum, Perth, Australia. 184 pp. White, K. M. 1951. On a collection of molluscs, mainly nudibranchs, from the Red Sea. Proceedings of the Malacological Society, London 28: 241–253. Willan, R. C. & Morton, J. 1984. Marine Molluscs Part 2: Opisthobranchia. University of Auckland, New Zealand. 106 pp. Winckworth, R. 1942. A note on Rang, Leuckart and Ehrenberg. Proceedings of the Malacological Society 29 (5–6): 2. Yaron, I., Schiotte, T. & Wium-Andersen, G. 1986. A review of molluscan taxa described by P. Forsskål and C. Niebuhr with citation of original descriptions, discussion of type material available, and selection of some lectotypes. Steenstrupia 12: 157–203. Yonow, N. 1986. Red Sea Phyllidiidae (Mollusca, Nudibranchia) with descriptions of new species. Journal of Natural History 20 (6): 1401–1428.

292 Bibliography

Yonow, N. 1988. Red Sea Opisthobranchia. 1: The family Phyllidiidae (Mollusca, Nudibranchia). Fauna of Saudi Arabia 9: 138–151. Yonow, N. 1989. Red Sea Opisthobranchia. 2. The family Chromodorididae (Mollusca, Nudibranchia). Fauna of Saudi Arabia 10: 290–309. Yonow, N. 1990. Red Sea Opisthobranchia. 3. The orders Sacoglossa, Cephalaspidea, and Nudibranchia: Doridacea (Mollusca, Opisthobranchia). Fauna of Saudi Arabia 11: 286–299. Yonow, N. 1992. Observations on the diet of Philinopsis cyanea (Martens) (Cephalaspidea, Aglajidae). Journal of Conchology 34 (4): 199–204. Yonow, N. 1994a. A new species and new record of Chelidonura from the Red Sea (Cephalaspidea: Aglajidae). Journal of Conchology 35 (2): 141–147. Yonow, N. 1994b. Opisthobranchs from the Maldive Islands, including descriptions of seven new species (Mollusca: Gastropoda). Revue française d’aquariologie herpétologie 20 (4): 97–130. Yonow, N. 2000. Red Sea Opisthobranchia. 4. The orders Cephalaspidea, Anaspidea, Notaspidea, and Nudibranchia: Dendronotacea and Aeolidacea. Fauna of Arabia 18: 87–131. Yonow, N. 2001. Results of the Rumphius Biohistorical Expedition to Ambon (1990). Part 11. Doridacea of the families Chromodorididae and Hexabranchidae (Mollusca, Gastropoda, Opisthobranchia, Nudibranchia), including additional Moluccan material. Zoologische Mededelingen, Leiden 75 (1–15): 1–50. Yonow, N. in press. Opisthobranchs of the Gulf of Eilat and the Red Sea: an account of similarities and differences. The Improbable Gulf – Oceanography and Marine Biology of the Gulf of Aqaba (Eilat). Magnes Press, Jerusalem, Israel. Yonow, N. & Hayward, P. J. 1991. Opisthobranches de l’lle Maurice, avec la description de deux espèces nouvelles (Mollusca: Opisthobranchia). Revue française d’aquariologie herpétologie 18 (1): 1–30. Yonow, N., Anderson, R. C. & Buttress, S. G. 2002. Opisthobranch molluscs from the Chagos Archipelago, Central Indian Ocean. Journal of Natural History 1: 831–882.

Bibliography 293

INDEX

All species, genera, and families recorded from the Red Sea are listed in this index. If they are in CAPITALS they denote families and higher taxa; italic typeface denotes valid species while synonyms are listed in nonitalics. SMALL CAPITALS are misidentifications; however, when a name is both a valid Red Sea species and a misidentification, it is not listed in small capitals. Page numbers in bold numerals refer to main entries in “Species Accounts” and are illustrated. Page numbers in (bold) in parentheses refer to photographs of tentatively identified animals with no formal entry in the “Species Accounts”. A acicula, Creseis 8, 20 Aclesia erythraea 55 Acteocina fusiformis 52 Acteocina inconspicua 53 Acteocina involuta 53 Acteocina simplex 53 ACTEON FlAMMEUS 52 Acteon sp. 52 ACTEON SULCATA 52 ACTEONIDAE 52 ACTINOCYCLIDAE 58, 149 Actinocyclus fragilis 58 Actinocyclus sordida 58 Actinocyclus velutinus 59 Actinocyclus verrucosus 58, 149 Actinodoris sponsa 65 acuta, Ringicula 52, 82 Aegires citrinus 57, 143 Aegires villosus 57, 143 AEGIRIDAE 57, 142 Aeolidia habessinica 64 Aeolidiella alba 64, 251 Aeolidiella indica 64 AEOLIDIELLA ORIENTALIS 64 AEOLIDIIDAE 64, 251 AEOLIDINA 63, 244 affinis, Pupa 52 affinis, Tambja 57, 137 africana, Chromodoris 59, 65, 174 africana, Thorunna 61, 171 Aglaja cyanea 53 Aglaja phaeoreticulata 53, 82 AGLAJIDAE 53, 76

294 Index

Akera soluta 54 alba, Aeolidiella 64, 251 alba, Aplysiella gravieri var. 55 alba, Gymnodoris 57, 130, 131 albiguttatus, Pleurobranchus 34, 56, 110 alboannulata, Noumea 61, 198 albocollaris, Phanerophthalmus 54, 93, 104 albocrossata, Berthella stellata 56 albo-limbata, Doris 38, 61 albomaculata var. erythraea, Glossodoris 50, 59 albus, Dermatobranchus 63, 240 Aldisa sp. 1 58, 155 Aldisa sp. 2 58, 156 amboinei, Euselenops ? 56 ampulla, Bulla 54, 82, 88, 95 Analogium striatum 48, 57, 129 Analogium sp. 57, 130, 132 ANASPIDEA 54, 96 annnuligera, Kentrodoris 59 annulata, Cerberilla 64 annulata, Chromodoris 59, 178 APICULATA, HALGERDA 58 apiculata, Sclerodoris 58, 160 Aplysia argus 38, 54, 97 Aplysia cornigera 54 Aplysia dactylomela 38, 54, 96 Aplysia fasciata 55 APLYSIA GRACILIS 55 Aplysia oculifera 55, 97, 106, 238 Aplysia parvula 55, 98, 101 Aplysia sp. 55, 98, 100 Aplysiella gravieri 55 Aplysiella gravieri var. alba 55 APLYSIIDAE 54, 96

arabica, Phyllidia 62 Archidoris o’donoghuei 59 Archidoris staminea 58 Archidoris sp. 58 Archidoris vayssierea 58 Ardeadoris egretta 59, 169 areolata, Atagema 58 areolata, Trippa 58, 157 argus, Aplysia 38, 54, 97 argus, Deridobranchus 65 argus, Tethys 54 armata, Hoplodoris ? (275) Armina cf. major (276) Armina semperi 50, 63 Armina semperi var. erythraea 50, 63 ARMINIDAE 63, 240 ARMININA 63, 240 Artachaea clavata 59 Artachaea intermedia 59 Artachaea verrucosa 59 aspersa, Chromodoris 59, 179 Asteronotus cespitosus 50, 59, 162 Asteronotus denticulata 59 Asteronotus hemprichi 50, 59 Atagema areolata 58 Atagema ornata 58, 156 Atagema spongiosa 58, 157 atlanticus, Glaucus 64 atromarginata, Casella 46, 60 atromarginata, Glossodoris 46, 60 Atys cylindricus 53 Atys ehrenbergi 54 Atys naucum 54, 88 Atys semistriatus 54, 89 auricularia, Dolabella 28, 55, 101 australis, Spurilla 64, 254 B babai, Chelidonura 81 Baeolidia fusiformis 64, 251 Baeolidia cf. fusiformis (252) Baeolidia moebii 64 ballantinei, Hydelopsis 19 bella, Eolida 46, 64, 263 Berthelinia sp. 56 Berthella granulata 55 Berthella pellucida 56 Berthella stellata 55 Berthella stellata albocrossata 55

Berthellina citrina 38, 55, 109 Berthellina saidensis 56 bicolor, Flabellina 63, 233, 244 bicolor, Gymnodoris 134 bifurcata, Hoplodoris 58, 158 bilas, Flabellina 63, 246 bizonata, Ventomnestia 53 Bornella digitata 48, 63 Bornella marmorata 63 Bornella stellifer 48, 63, 233 BORNELLIDAE 63, 233 Bornellopsis kabretiana 63 bourbonica, Julia 56 Brachyclanis pantherina 60 brachyphylla, Pterodoris 65 brevipes, Notarchus 55 brevitentaculus, Favorinus horridus 65 brunneomarginatum, Siphopteron 87 bucephala, Melibe 63, 239 Bulla ampulla 54, 82, 88, 95 Bulla smaragdina 46, 54, 94 BULLA STRIATA 54 BULLIDAE 54, 95 Bursatella leachii savignana 34, 38, 55 Bursatella savignana 55 BURSATELLA SP. 56 C Cadlinella ornatissima 59, 172 Cadlinella sp. 59, 173 CALIPHYLLA SP. 57 Caloria indica 64, 257 calyculata, Smaragdinella 54 carbunculosa, Dendrodoris 214 cardinalis, Phyllidiopsis 62, 221 Casella atromarginata 46, 60 Casella foxi 60 Casella obsoleta 60 castanea, Goniodoris 57 Catriona susa 64 CEPHALASPIDEA 52, 74 Ceratosoma cornigerum 50, 61 Ceratosoma jousseaumei 61 Ceratosoma rhopalicum 61 Ceratosoma tenue 50, 61, 203 Ceratosoma trilobatum 61, 203, 204 Cerberilla annulata 64 cespitosus, Asteronotus 50, 59, 162 ceylonica, Gymnodoris 57, 132, 136

Index 295

ceylonicus, Plocamopherus 141 charlottae, Chromodoris 59, 180 charlottae, Glossodoris 59 CHELIDONURA AFRICANA 53 Chelidonura babai 81 Chelidonura flavolobata 53, 76 Chelidonura fulvipunctata 53, 78 Chelidonura livida 53, 78 Chelidonura punctata 53, 79 Chelidonura sandrana 29, 53, 80, 82 chlorophaeus, Plakobranchus 56 CHROMODORIDIDAE 59, 169 Chromodoris africana 59, 65, 174 Chromodoris annulata 59, 178 Chromodoris aspersa 59, 179 Chromodoris charlottae 59, 180 Chromodoris cf. decora 60, 186 Chromodoris diardii 60 Chromodoris fidelis 59, 182 Chromodoris geminus 59, 183 CHROMODORIS ? INOPINATA 59, 185 Chromodoris inornata 59 Chromodoris marindica 185 Chromodoris obsoleta 46, 50, 59, 184 Chromodoris pulchella 61 Chromodoris quadricolor 46, 50, 60, 65, 174, 176, 177 Chromodoris reticulata 185 CHROMODORIS RETICULATA VAR. 59 Chromodoris runcinata 48, 60 Chromodoris strigata 60, 177 Chromodoris sp. 59, 60, 61, 186 Chromodoris tennentana 60 Chromodoris tenui-linearis 48, 61, 197 Chromodoris tinctoria 46, 60, 180, 184 Chromodoris verrieri 60, 185 Chromodoris cf. verrieri (185) Chromodoris vicina 60 cincta, Glossodoris 27, 60, 188 citrina, Berthellina 38, 55, 109 citrina, Gymnodoris 57, 134 citrinus, Aegires 57, 143 citrinus, Pleurobranchus 38, 55 citrinus, Stylocheilus 106 clavata, Artachaea 59 Clio convexa 20 colemani, Phyllodesmium 64, 260 collaris, Phanerophthalmus 54 Colpodaspis thompsoni 52, 76 concentrica, Doris 34, 58

296 Index

convexa, Clio 20 cornigera, Aplysia 55 coronata, Dendrodoris 61, 208 Coryphellina rubrolineata 63 Creseis acicula 8, 20 crosslandi, Sebadoris 58 crosslandi, Thordisa 58 Crosslandia fusca 63 Crosslandia viridis 63 cruenta, Platydoris 168 crypticum, Phyllodesmium 64, 261 Cryptophthalmus olivaceus 36, 46, 54, 94 Cryptophthalmus smaragdinus 94 cuprea, Dendrodoris 61 cuprea, Doris 61 curta, Liloa 54, 92 Cuthona sibogae 64, 267 Cuthona sp. 64 cuvieri, Dobabrifera 55 cyanea, Aglaja 53 cyanea, Philinopsis 53, 81 cyanobranchiata, Marioniopsis 38, 62, 226 cyanobranchiata, Tritonia 38, 62 cyanocaudata, Haminoea 54 cyanomarginata, Haminoea 54, 91 Cyerce elegans 56, 126 CYLICHNA MICA 53 Cylichna mongii 53 CYLICHNA PULVISCULUS 53 CYLICHNA SEMISULCATA 54 Cylichna villersii 53 CYLICHNIDAE 53 cylindricus, Atys 53 D dactylomela, Aplysia 38, 54, 96 dautzenbergi, Phyllidia 48, 62 dautzenbergi, Phyllidiopsis 48, 62, 222 decora, Chromodoris cf. 60, 186 decorata, Elysia 56 decorata, Hallaxa 58 decorata, Thuridilla 56, 123 DENDRODORIDIDAE 61, 208 Dendrodoris carbunculosa 214 Dendrodoris coronata 61, 208 Dendrodoris cuprea 61 Dendrodoris elongata 61, 210 Dendrodoris fumata 38, 48, 61, 211 Dendrodoris immaculata 61

Dendrodoris jousseaumei 48, 61 Dendrodoris leptopus 61 Dendrodoris lugubris 61 Dendrodoris nigra 34, 38, 48, 61, 212 Dendrodoris nigropunctata 61 DENDRODORIS PUSTULOSA 62 Dendrodoris rubra 61 Dendrodoris sp. 214 Dendrodoris tigrina 34, 46, 61 Dendrodoris tuberculosa 62, 213 DENDRONOTINA 62, 225 denticulata, Asteronotus 59 denticulata, Peronodoris 59 dentifer, Diniatys 54. 90, 91 Deridobranchus argus 65 Dermatobranchus albus 63, 240 Dermatobranchus glabrus 63 Dermatobranchus gonatophorus 63, 240 Dermatobranchus ornatus 63, 242 Dermatobranchus striatus 63 Dermatobranchus sp. 63, 242 desgenettii, Retusa 53 DIAPHANIDAE 52, 76 diardii, Chromodoris 60 digitata, Bornella 48, 63 Diniatys dentifer 54. 90, 91 Diniatys dubius 54, 91 DISCODORIDIDAE 58, 150 DISCODORIS AMBOINENSIS 58 DISCODORIS CONCINNA 58 Discodoris erythraeensis 58 Discodoris fragilis 58, 150, 153 Discodoris granulata 58 Discodoris ? mauritiana (274) Discodoris schmeltziana 58, 151 Discodoris sp. 58 Dolabella auricularia 28, 55, 101 DOLABELLA GIGAS 55, 102 Dolabella hemprichii 55 Dolabella rumphii 55 Dobabrifera cuvieri 55 Dolabrifera dolabrifera 55, 102 dolabrifera, Dolabrifera 55, 102 dollfusi, Glossodoris 50, 60 dollfusi, Hypselodoris 50, 60, 192 DORIDIDAE 58, 155 DORIDINA 57, 128 Doridopsis sp. 61 Doriopsilla sp. 62

Doriopsis granulosa 34, 58 Doriopsis jousseaumei 48, 61 Doriopsis nigra 48, 61 Doriopsis nigropunctata 61 Doriopsis rosea 48, 61 Doris albo-limbata 38, 61 Doris concentrica 34, 58 Doris cuprea 61 Doris erythraea 65 Doris exanthemata 58 Doris fumata 38, 61 Doris granulata 58 Doris immaculata 34, 61 Doris impudica 46 Doris infucata 46, 60 Doris marmorata 34, 60 Doris obsoleta 46, 60 Doris pallida 46, 60 Doris plumbea 60 Doris pulchella 46, 61 Doris punctata 46, 61 Doris quadricolor 46, 60 Doris sanguineus 38, 57 Doris sordida 58 Doris sp. 58 Doris tigrina 34, 61 Doris tinctoria 46, 60 Doris xantholeuca 60 Doto orcha 63, 234 DOTOIDAE 63, 234 dubius, Diniatys 54, 91 Dunga ocellata 64, 255 Durvilledoris lemniscata 60, 187 Durvilledoris pusilla 60, 188 E egretta, Ardeadoris 59, 169 ehrenbergi, Atys 54 electra, Glossodoris 60, 170 elegans, Cyerce 56, 126 elegans, Tritonia 36, 62 elegans, Tritoniopsis 36, 62, 232 elongata, Dendrodoris 61, 210 Elysia decorata 56 ELYSIA GRANDIFOLIA 56, 122 ELYSIA ORNATA 122 Elysia sp. 1 56, 120 Elysia sp. 2 56, 121 Elysia tomentosa 56, 119, (272)

Index 297

Elysia trilobata 56, 120 Elysiella halimedae 122 Elysiella pusilla 56, 122 ELYSIIDAE 56, 119 Embletonia gracile 64, 266 EMBLETONIIDAE 64, 266 engeli, Melibe 237, 239 Eolida bella 46, 64, 263 erythraea, Aclesia 55 erythraea, Armina semperi var. 50, 63 erythraea, Doris 65 erythraea, Glossodoris albomaculata var. 50, 60 erythraeensis, Discodoris 58 erythraeensis, Paradoris 27, 58, 153 EUBRANCHIDAE 64, 255 Eubranchus rubropunctatus 64, 256 Euselenops ? amboinei 56 Euselenops luniceps 56 exanthemata, Doris 58 exquisita, Julia 22, 56 F Facelina rhodopos 64, 248 “Facelina” sp. 64, 250 FACELINIDAE 64, 248 fasciata, Aplysia 55 fastuosum, Phyllodesmium 65 Favorinus horridus brevitentaculus 65 Favorinus japonicus 64, 258 Favorinus tsuruganus 64, 259 ferruginosa, Madrella 25, 63, 243 fidelis, Chromodoris 59, 182 Flabellina bicolor 63, 233, 244 Flabellina bilas 63, 246 Flabellina rubrolineata 63, 247 “Flabellina” sp. 63, 245 FLABELLINIDAE 63, 244 flava, Noumea 61, 199 flavolobata, Chelidonura 53, 76 forskalii, Pleurobranchus 38, 56, 112 fourierrii, Pyrunculus 53 foxi, Casella 60 fragilis, Actinocyclus 58 fragilis, Discodoris 58, 150, 153 Fryeria marindica 216 Fryeria rueppelii 46, 62, 215 fulvicola, Marioniopsis 62, 228 fulvipunctata, Chelidonura 53, 78 fumata, Dendrodoris 38, 48, 61, 211

298 Index

fumata, Doris 38, 61 funebris, Jorunna 59, 164 fusca, Crosslandia 63 fusiformis, Acteocina 52 fusiformis, Baeolidia 64, 251 fusiformis, Baeolidia cf. (252) G GASTROPTERIDAE 53, 85 geminus, Chromodoris 59, 183 geographica, Syphonota 55 ghardaqana, Risbecia 61, (203) ghomfodensis, Scyllaea 63 girardi, Ventomnestia 53 glabrus, Dermatobranchus 63 glama, Marionia 38, 63, 225 glama, Tritonia 38, 62 GLAUCIDAE 64, 257 Glaucilla marginata 64, 260 Glaucus atlanticus 64 Glossodoris albomaculata var. erythraea 50, 59 Glossodoris atromarginata 46, 60 Glossodoris charlottae 59 Glossodoris cincta 27, 60, 188 Glossodoris dollfusi 50, 60 Glossodoris electra 60, 170 Glossodoris hikuerensis 60, 190 GLOSSODORIS LUTEOROSEA 60 Glossodoris pallida 46, 60, 191 Glossodoris plumbea 60 Glossodoris quadricolor 60 GLOSSODORIS RUFOMARGINATA 60 Glossodoris symmetricus 60, 192 Glossodoris sp. 60 gonatophorus, Dermatobranchus 63, 240 Goniodoridella savignyi 57 GONIODORIDIDAE 57, 128 Goniodoris castanea 57 Goniodoris joubini 57, 128 Goniodoris savignyi 57 gracile, Embletonia 64, 266 grandiflora, Hoplodoris 58, 158 grandis, Pleurobranchus 56, 112, 114, 116 grandis, Pleurobranchus cf. 56, 114, 115 granulata, Berthella 55 granulata, Discodoris 58 granulata, Doris 58 granulosa, Doriopsis 34, 58 Graviera rugosa 58, 59

gravieri, Aplysiella 55 gravieri, Tritoniopsis 48, 62 gravieri var. alba, Aplysiella 55 guamensis, Odontaglaja 53, 83 GYMNODORIDIDAE 57, 129 Gymnodoris alba 57, 130, 131 Gymnodoris bicolor 134 Gymnodoris ceylonica 57, 132, 136 Gymnodoris citrina 57, 134 Gymnodoris impudica 46, 57, 135 Gymnodoris inornata 57, 135 Gymnodoris cf. nigricolor 57 Gymnodoris rubropapulosa 57, 135 Gymnodoris striata 57 Gymnodoris sp. 57, 136 H habessinica, Aeolidia 64 HALGERDA APICULATA 58 Halgerda willeyi 48, 59, 163 HALGERDIDAE 59, 162 halimedae, Elysiella 122 Hallaxa decorata 58 Hallaxa indecora 48, 58, 150 Haminoea cyanocaudata 54 Haminoea cyanomarginata 54, 91 Haminoea pemphis 54 HAMINOEA TENERA 54 HAMINOEIDAE 54, 88 hemprichi, Asteronotus 50, 59 hemprichii, Dolabella 55 HEXABRANCHIDAE 57, 144 Hexabranchus marginatus 30, 146 Hexabranchus plicatus 57 Hexabranchus praetextus 36, 57 Hexabranchus sanguineus 29, 36, 38, 57, 144, 147 Hexabranchus sp. 58, 114, 147 Hexabranchus suezensis 57 hikuerensis, Glossodoris 60, 190 Hoplodoris ? armata (275) Hoplodoris bifurcata 58, 158 Hoplodoris grandiflora 58, 158 Hoplodoris nodulosa 159 Hoplodoris pustulata 58, 158 horridus brevitentaculus, Favorinus 64 hyalinum, Phyllodesmium 64, 262 HYDATINA AMPLUSTRE 52 Hydatina physis 52, 74, 75 Hydatina stromfelti 75 Hydatina velum 52, 75

Hydatina vesicaria 75 Hydatina zonata 52, 75 HYDATINIDAE 52, 74 Hydelopsis ballantinei 19 Hypselodoris dollfusi 50, 60, 192 Hypselodoris infucata 34, 46, 48, 60, 193 Hypselodoris maculosa 60, 195 Hypselodoris maridadilus 60, 196 Hypselodoris nigrostriata 60, 197 I ianthina, Pteraeolidia 64, 265 immaculata, Dendrodoris 61 immaculata, Doris 34, 61 imperator, Periclimenes 114, 145, 147, 203 imperialis, Plocamopherus 141 impudica, Doris 46 impudica, Gymnodoris 46, 57, 135 incertae, Platydoris 59 inconspicua, Acteocina 53 indecora, Hallaxa 48, 58, 150 indica, Aeolidiella 64 indica, Caloria 64, 257 indicus, Notarchus 29, 30, 55, 97, 105, 238 indicus, Plocamopherus 29, 48, 57, 140 indopacifica, Thuridilla ? 56, 124 infucata, Doris 46, 60 infucata, Hypselodoris 34, 46, 48, 60, 193 inornata, Chromodoris 59 inornata, Gymnodoris 57, 135 intecta, Trippa 156 intermedia, Artachaea 59 involuta, Acteocina 53 J japonicus, Favorinus 64, 258 Jorunna funebris 59, 164 Jorunna pantherina 59, 165 Jorunna sp. 59, 165 joubini, Goniodoris 57, 128 jousseaumei, Ceratosoma 61 jousseaumei, Dendrodoris 48, 61 jousseaumei, Doriopsis 48, 61 Julia bourbonica 56 Julia exquisita 22, 56 JULIIDAE 56 K kabretiana, Bornellopsis 63 katalexis, Mexichromis 61, 197 Index 299

KENTRODORIDIDAE 59, 164 Kentrodoris annnuligera 59 Kentrodoris labialis 59 Kentrodoris rubescens 164 L labialis, Kentrodoris 59 laciniatus, Notarchus 38, 55 leachii savignana, Bursatella 34, 38, 55 lemniscata, Durvilledoris 60, 187 leptopus, Dendrodoris 61 levis, Marioniopsis 62, 228 Liloa curta 54, 92 Limacia trochiformis 20 limaciformis, Nembrotha 57 limaciformis, Tambja 57, 139 Limenandra nodosa 64 Limenandra sp. 64, 253 lithensis, Roxania 53 livida, Chelidonura 53, 78 LOMANOTIDAE 63, 235 Lomanotus vermiformis 63, 235 longicauda, Stylocheilus 55, 106 lugubris, Phestilla 64, 268 lugubris, Dendrodoris 61 luniceps, Euselenops 56 M maculosa, Hypselodoris 60, 195 Madrella ferruginosa 25, 63, 243 MADRELLIDAE 63, 243 magnifica, Miamira 61, 206 magnum, Phyllodesmium 46, 65, 263 major, Armina cf. (276) major, Spurilla 64, 255 mamillatus, Pleurobranchus 56, 117 marginata, Glaucilla 64, 260 marginatus, Hexabranchus 30, 146 maridadilus, Hypselodoris 60, 196 marindica, Chromodoris 185 marindica, Fryeria 216 Marionia glama 38, 62, 225 Marioniopsis cyanobranchiata 38, 62, 226 Marioniopsis fulvicola 62, 228 Marioniopsis levis 62, 228 Marioniopsis rubra 38, 62, 229 Marioniopsis viridescens 62, 230 marmorata, Bornella 63 marmorata, Doris 34, 60 mauritiana, Discodoris ? (274) 300 Index

megalocera, Nembrotha 29, 57, 137 melanobrachia, Phestilla 64, 269 melanocera, Phyllidia 62 Melibe bucephala 63, 239 Melibe engeli 237, 239 Melibe rangii 29, 63, 97, 106, 237 Mexichromis katalexis 61, 197 MEXICHROMIS MARIEI 61, 198 Mexichromis multituberculata 198 Miamira magnifica 61, 206 miamirana, Orodoris 48, 61, 207 moebii, Baeolidia 64 monacha, Phyllidia 62 monacha, Phyllidiopsis 62, 223 mongii, Cylichna 53 mucronata, Utriculastra 53 multifaria, Phyllidia 62, 216, 217 multituberculata, Mexichromis 198 N naucum, Atys 54, 88 Nembrotha limaciformis 57 Nembrotha megalocera 29, 57, 137 nigra, Dendrodoris 34, 38, 48, 61, 212 nigra, Doriopsis 48, 61 nigricolor, Gymnodoris cf. 57 nigromarginatum, Siphopteron 87 nigropunctata, Dendrodoris 61 nigropunctata, Doriopsis 61 nigrostriata, Hypselodoris 60, 197 nodosa, Limenandra 64 nodulosa, Hoplodoris 159 norba, Noumea 61, 199 NOTARCHIDAE 55, 105 Notarchus brevipes 55 Notarchus indicus 29, 30, 55, 97, 105, 238 Notarchus laciniatus 38, 55 Noumea alboannulata 61, 198 Noumea flava 61, 199 Noumea norba 61, 199 NOUMEA PURPUREA 61, 199 Noumea simplex 61, 200 Noumea sudanica 61, 200 nubilosa, Sebadoris 29, 58, 153 NUDIBRANCHIA 57, 128 O o’donoghuei, Archidoris 59 oblongus, Pleurobranchus 34, 56, 110 obsoleta, Casella 60

obsoleta, Chromodoris 46, 50, 59, 184 obsoleta, Doris 46, 59 ocellata, Dunga 64, 255 ocellatus, Plocamopherus 38, 58, 142 ocellatus, Plakobranchus 48, 57, 124, 130 oculifera, Aplysia 55, 97, 106, 238 Odontaglaja guamensis 53, 83 Ohola pacifica 57 olivaceus, Cryptophthalmus 36, 46, 54, 94 olivaceus, Phanerophthalmus 36, 54, 94, 104, 122 orcha, Doto 63, 234 orientalis, Polybranchia 56, 127 ornata, Atagema 58, 156 ornatissima, Cadlinella 59, 172 ornatum, Sagaminopteron cf. 53, 85 ornatus, Dermatobranchus 63, 242 ornatus, Stiliger 36, 56 Orodoris miamirana 48, 61, 207 Oxynoe viridis 56, 118 OXYNOIDAE 56, 118 P pacifica, Ohola 57 pacifica, Thecacera 57, 140 pallida, Doris 46, 60 pallida, Glossodoris 46, 60, 191 pantherina, Brachyclanis 60 pantherina, Jorunna 59, 165 Paradoris erythraeensis 27, 58, 153 parvula, Aplysia 55, 98, 101 pelagica, Scyllaea 63, 236 pellucida, Berthella 56 pemphis, Haminoea 54 Periclimenes imperator 114, 145, 147, 203 Peronodoris denticulata 59 perrieri, Pleurobranchus 56 Petalifera petalifera 55 petalifera, Petalifera 55 Petalifera ramosa 55, 103 phaeoreticulata, Aglaja 53, 82 Phanerophthalmus albocollaris 54, 93, 104 Phanerophthalmus collaris 54 Phanerophthalmus olivaceus 36, 54, 94, 104, 122 Phanerophthalmus smaragdinus 54 Phestilla lugubris 64, 268 Phestilla melanobrachia 64, 269 Philine vaillanti 50, 53 PHILINIDAE 53 Philinopsis cyanea 53, 81

Philinopsis reticulata 53, 82 Phyllaplysia sp. 55, 104 Phyllidia arabica 62 Phyllidia dautzenbergi 48, 62 Phyllidia melanocera 62 Phyllidia monacha 62 Phyllidia multifaria 62, 216, 217 Phyllidia pustulosa 46, 62 PHYLLIDIA QUADRILINEATA 62 Phyllidia schupporum 62, 217 Phyllidia sinaiensis 62 Phyllidia sudanensis 62 Phyllidia sp. 62 Phyllidia undula 62, 218 Phyllidia varicosa 62, 219 Phyllidiella pustulosa 62, 156, 220, 224 PHYLLIDIIDAE 62, 215 Phyllidiopsis cardinalis 62, 221 Phyllidiopsis dautzenbergi 48, 62, 222 Phyllidiopsis monacha 62, 223 Phyllidiopsis sinaiensis 62, 224 Phyllidiopsis sudanensis 63 Phyllodesmium colemani 64, 260 Phyllodesmium crypticum 64, 261 Phyllodesmium fastuosum 64 Phyllodesmium hyalinum 64, 262 Phyllodesmium magnum 46, 64, 263 Phyllodesmium xeniae 64 physis, Hydatina 52, 74, 75 picturata, Pterodoris 60 PLAKOBRANCHIDAE 56, 119 Plakobranchus chlorophaeus 56 Plakobranchus ocellatus 48, 57, 124, 130 PLATYDORIDIDAE 59, 167 Platydoris cruenta 168 Platydoris incertae 59 Platydoris scabra 59, 167 Platydoris striata 59, 168 Platydoris sp. 59 PLEUROBRANCHIDA 56, 109 PLEUROBRANCHIDAE 56, 109 Pleurobranchus albiguttatus 34, 56, 110 Pleurobranchus citrinus 38, 55 Pleurobranchus forskalii 38, 56, 112 Pleurobranchus grandis 56, 112, 114, 116 Pleurobranchus cf. grandis 56, 114, 115 Pleurobranchus mamillatus 56, 117 Pleurobranchus oblongus 34, 56, 110 Pleurobranchus perrieri 56

Index 301

Pleurobranchus ruppelii 56 Pleurobranchus semperi 56 Pleurobranchus tesselatus ? (112) plicatus, Hexabranchus 58 Plocamopherus ceylonicus 141 Plocamopherus imperialis 141 Plocamopherus indicus 29, 48, 57, 140 Plocamopherus ocellatus 38, 58, 142 plumbea, Doris 60 plumbea, Glossodoris 60 Polybranchia orientalis 56, 127 POLYBRANCHIIDAE 57, 126 POLYCERA SP. 57 POLYCERIDAE 57, 137 praetextus, Hexabranchus 36, 57 psychedelicum, Sagaminopteron cf. 53, 86 Pteraeolidia ianthina 64, 265 Pteraeolidia semperi 64 Pterodoris brachyphylla 65 Pterodoris picturata 60 pulchella, Chromodoris 61 pulchella, Doris 46, 61 pulchella, Risbecia 46, 61, 201 punctata, Chelidonura 53, 79 punctata, Doris 46, 61 Pupa affinis 52 Pupa cf. solidula 52 PUPA SULCATA 52 Pupa sp. 52 Pupa tessellata 52 pusilla, Durvilledoris 60, 188 pusilla, Elysiella 56, 122 pustulata, Hoplodoris 58, 158 pustulosa, Phyllidia 46, 62 pustulosa, Phyllidiella 62, 156, 220, 224 Pyrunculus fourierrii 53 Q quadricolor, Chromodoris 46, 50, 60, 65, 174, 176, 177 quadricolor, Doris 46, 60 quadricolor, Glossodoris 60 quadrispinosum, Siphopteron 87 R ramosa, Petalifera 55, 103 rangii, Melibe 29, 63, 97, 106, 237 reticulata, Chromodoris 185 reticulata, Philinopsis 53, 82 Retusa desgenettii 53

302 Index

Retusa tarutana 53 RETUSIDAE 53 rhodopos, Facelina 64, 248 rhopalicum, Ceratosoma 61 rigida, Spongiodoris 58 Ringicula acuta 52, 82 RINGICULA SAVIGNY 52 RINGICULIDAE 52 Risbecia ghardaqana 61, (203) Risbecia pulchella 46, 61, 201 rosea, Doriopsis 48, 61 Roxania lithensis 53 rubescens, Kentrodoris 164 rubra, Dendrodoris 61 rubra, Marioniopsis 38, 62, 229 rubra, Tritonia 38, 62 rubrolineata, Coryphellina 63 rubrolineata, Flabellina 63, 247 rubropapulosa, Gymnodoris 57, 135 rubropunctatus, Eubranchus 64, 256 rueppelii, Fryeria 46, 62, 215 rugosa, Graviera 58 rugosa, Sclerodoris 48, 58, (275) rumphii, Dolabella 55 runcinata, Chromodoris 48, 60 ruppelii, Pleurobranchus 56 S SACOGLOSSA 56, 118 Sagaminopteron cf. ornatum 53, 85 Sagaminopteron cf. psychedelicum 53, 86 saidensis, Berthellina 56 sandrana, Chelidonura 29, 53, 80, 82 sanguineus, Doris 38, 57 sanguineus, Hexabranchus 29, 36, 38, 57, 144, 147 savignana, Bursatella 55 savignana, Bursatella leachii 34, 38, 55 savignyi, Goniodoridella 57 savignyi, Goniodoris 57 scabra, Platydoris 59, 167 SCAPHANDRIDAE 53 schmeltziana, Discodoris 58, 151 schupporum, Phyllidia 62, 217 Sclerodoris apiculata 58, 160 Sclerodoris rugosa 48, 58, (275) Sclerodoris tuberculata 58, 161 Scyllaea ghomfodensis 63 Scyllaea pelagica 63, 236 SCYLLAEIDAE 63, 236

Sebadoris crosslandi 58 Sebadoris nubilosa 29, 58, 153 semistriatus, Atys 54, 89 semperi, Armina 50, 63 semperi, Pleurobranchus 56 semperi, Pteraeolidia 64 semperi var. erythraea, Armina 50, 63 sibogae, Cuthona 64, 267 sieboldi, Smaragdinella 54 simplex, Noumea 61, 200 simplex, Acteocina 53 sinaiensis, Phyllidia 62 sinaiensis, Phyllidiopsis 62, 224 Siphopteron brunneomarginatum 87 Siphopteron nigromarginatum 87 Siphopteron quadrispinosum 87 Siphopteron sp. 53, 87 smaragdina, Bulla 46, 54, 94 Smaragdinella calyculata 54 Smaragdinella sieboldi 54 SMARAGDINELLIDAE 54, 93 smaragdinus, Cryptophthalmus 94 smaragdinus, Phanerophthalmus 54 solidula, Pupa cf. 52 soluta, Akera 54 sordida, Doris 58 sordida, Actinocyclus 58 Spongiodoris rigida 58 Spongiodoris sp. 58 spongiosa, Atagema 58, 157 spongiosa, Trippa 58 sponsa, Actinodoris 65 Spurilla australis 64, 254 Spurilla major 64, 255 staminea, Archidoris 58 stellata albocrossata, Berthella 55 stellata, Berthella 55 stellifer, Bornella 48, 63, 233 STILIGERIDAE 56 Stiliger ornatus 36, 56 striata, Gymnodoris 57 striata, Platydoris 59, 168 striata, Trevelyana 48, 57 striatum, Analogium 48, 57, 129 striatus, Dermatobranchus 64 striatus, Stylocheilus 55, 108 strigata, Chromodoris 60, 177 stromfelti, Hydatina 75 Stylocheilus citrinus 106 Stylocheilus longicauda 55, 106

Stylocheilus striatus 55, 108 sudanensis, Phyllidia 62 sudanensis, Phyllidiopsis 62 sudanica, Noumea 61, 200 suezensis, Hexabranchus 57 susa, Catriona 64 symmetricus, Glossodoris 60, 192 Syphonota geographica 55 T Tambja affinis 57, 137 Tambja limaciformis 57, 139 tarutana, Retusa 53 tennentana, Chromodoris 60 tenue, Ceratosoma 50, 61, 203 tenui-linearis, Chromodoris 48, 61, 197 TERGIPEDIDAE 64, 267 tesselatus ?, Pleurobranchus (112) tessellata, Pupa 52 TETHYIDAE 63, 237 Tethys argus 55 Tethys leporina 55 Thecacera pacifica 57, 140 thompsoni, Colpodaspis 52, 76 Thordisa crosslandi 58 Thorunna africana 61, 171 THORUNNA FURTIVA 61 Thuridilla ? indopacifica 56, 124 Thuridilla decorata 56, 123 tigrina, Dendrodoris 34, 46, 61 tigrina, Doris 34, 61 tinctoria, Chromodoris 46, 60, 180, 184 tinctoria, Doris 46, 60 toddi, Trapania 57 tomentosa, Elysia 56, 119, (272) TORNATINA OLIVAEFORMIS 53 Trapania toddi 57 TREVELYANA BICOLOR 57 TREVELYANA CONCINNA 57 Trevelyana striata 48, 57 trilobata, Elysia 56, 120 trilobatum, Ceratosoma 61, 203, 204 TRIOPHIDAE 57, 140 Trippa areolata 58, 157 Trippa intecta 156 Trippa spongiosa 58 Tritonia cyanobranchiata 38, 62 Tritonia elegans 36, 62 Tritonia glama 38, 62 Tritonia rubra 38, 62 Index 303

TRITONIIDAE 62, 225 Tritoniopsis elegans 36, 62, 232 Tritoniopsis gravieri 48, 62 Tritoniopsis sp. 63 trochiformis, Limacia 20 tsuruganus, Favorinus 64, 259 tuberculata, Sclerodoris 58, 161 tuberculosa, Dendrodoris 62, 213 U undula, Phyllidia 62, 218 Utriculastra mucronata 53 V vaillanti, Philine 50, 53 varicosa, Phyllidia 62, 219 vayssierea, Archidoris 58 velum, Hydatina 52, 75 velutinus, Actinocyclus 59 Ventomnestia bizonata 53 Ventomnestia girardi 53 Ventomnestia villosa 53 vermiformis, Lomanotus 63, 235

304 Index

verrieri, Chromodoris 60, 185 verrieri, Chromodoris cf. (185) verrucosa, Artachaea 59 verrucosus, Actinocyclus 58, 149 vesicaria, Hydatina 75 vicina, Chromodoris 60 villersii, Cylichna 53 villosa, Ventomnestia 53 villosus, Aegires 57, 143 viridescens, Marioniopsis 62, 230 viridis, Crosslandia 63 viridis, Oxynoe 56, 118 W willeyi, Halgerda 48, 59, 163 X xantholeuca, Doris 60 xeniae, Phyllodesmium 64 Z zonata, Hydatina 52, 75