Philinopsis cyanea (Martens, 1879)

Valdés, Ángel, 2008, Deep-sea “ cephalaspidean ” heterobranchs (Gastropoda) from the tropical southwest Pacific, Mémoires du Muséum national d'Histoire naturelle 196, pp. 587-792 : 783-792

Philinopsis cyanea (Martens, 1879)

Figs 97B, 101, 102

Doridium cyaneum von Martens, 1879: 738 .

TYPE MATERIAL. — Not examined .

TYPE LOCALITY. — Mozambique .

MATERIAL EXAMINED. — New Caledonia proper. BATHUS 1 : Fiji. BORDAU 1 : stn CP 1445, 17°10’S, 178°42’W, 350-365 m, 1 lv. stn CP 668, 20°57’S, 165°35’E, 205-219 m, 1 lv. Tonga. BORDAU 2 : stn CP 1526, Eua, 21°16’S, 174°59’W, 463- Vanuatu. MUSORSTOM 8: stn CP 963, 20°20’S, 169°49’E, 464 m, 1 lv (Fig. 102); stn CP 1641, NW of Tongatapu , 21°09’S, 400-440 m, 1 lv (Fig. 97B). 175°22’W, 395 m, 1 lv .

DISTRIBUTION. — Widespread in the Indo-Pacific reported from numerous localities. Material herein collected from Fiji, New Caledonia, Tonga and Vanuatu (Fig. 101), in 209-463 m.

DESCRIPTION. — External morphology. The body is oval, 21 mm long (largest specimen examined). The cephalic shield is oval and large, comprising about 2/3 of the body length (Fig. 102A). At the posterior end of the cephalic shield there is a short and triangular dorsal crest. The 2 Hancock’s organs are composed of about 15 simple folds. The parapodia are narrow and do not reach the midline of the body. The posterior shield is rounded posteriorly, with 2 lobes. The gill is simple, with 16 simple lamellae. The colour of the living animals is pale grey, with a number of opaque white spots all over the body surface, more densely arranged near the lateral edges (Fig. 97B). There is dark brown pigment on the edges of the parapodia and the anterior and posterior ends of the body and opaque white elongate spots at the end of the posterior lobes and head.

Shell morphology. The shell is internal, wide and curved, with only the posterior end calcified. It is on the posterior end of the body. Sculpture of a number of simple growth lines (Fig. 102D).

Anatomy. The pharynx is large and muscular, and connects posteriorly with the wide oesophagus (Fig. 102C). The oesophagus opens into a large and lobate digestive gland.

The reproductive system is monoaulic (Fig. 102B). The ampulla is long and convoluted. It connects directly to the mucous gland, which has 2 curved lobes. The small and pyriform seminal receptacle also connects to the mucous gland near the connection of the ampulla. From the common atrium a long duct connects to the elongate bursa copulatrix. From the gonopore an open seminal groove runs anteriorly to the protrusible cephalic penis and the prostate. The prostate is thin and very long. The penis is thin and long, with a lobe and a retractor muscle attached.

REMARKS. — The external colouration of the specimens here examined from the southwest Pacific matches the redescription of Philinopsis cyanea (Martens, 1879) by Rudman (1974), and therefore they are included in this species.

Philinopsis cyanea is included in the genus Philinopsis based on the presence of the following combination of characters: cephalic shield simply rounded, posterior shield with 2 short lobes and lacking a flagellum, head with a single pair of sensory processes, dorsal and ventral labial glands present, gill simple, buccal mass large and muscular, reproductive system monaulic, with the seminal receptacle entering in the genital atrium, mucous gland bilobed, hermaphrodite duct without a branch connecting to the albumen and mucous glands, penis unarmed, simple.

DISCUSSION

SHELL MORPHOLOGY VS. ANATOMY

Most of the deep-water material studied herein consists of empty shells. Therefore, numerous species have been identified and in some cases described here based solely on their shell morphology. Rudman (1971a), Marcus (1977) and Gosliner (1979) stressed the importance of the soft-body anatomy in “cephalaspidean” systematics. These authors used the external morphology, the radula, and the reproductive anatomy for the description and diagnosis of genera and species. In most cases the shell morphology contains information that allows differentiation of species, but the anatomy is fundamental for a correct generic placement in numerous groups of “cephalaspidean” heterobranchs. In the present study several species of Acteonidae and Retusidae have not been assigned to a genus because of the lack of anatomical data. Other species have been provisionally assigned to genera until complete specimens become available for study and anatomical data can be used to verify those placements.

Only in the cases in which the shell morphology is clearly distinct and diagnostic, have new species been named based on empty shells. This approach, far from being ideal, serves the purpose of describing a poorly known fauna that otherwise would remain undocumented until a new series of expeditions, including the use of expensive collecting techniques, allows collection of undamaged specimens. It will probably be several decades before such a series of expeditions is conducted in the southwest Pacific.

THE SOUTHWEST PACIFIC DEEP SEA FAUNA OF “CEPHALASPIDEAN” HETEROBRANCHS

The present collecting effort in the southwest Pacific has produced a large number of previously undocumented species. The largest number of species was found in the area comprising the Coral Sea, New Caledonia, Vanuatu, Fiji, Tonga and Wallis and Futuna, with 97 species. This is a reflection of the more intense collecting effort conducted in this area, which is also evident in the greater abundance of material. The area comprising the Philippines and Indonesia has produced less material and a smaller number of species, total 66, but the number of expeditions conducted in the area was considerably smaller.

Comparison among lists of species collected in different regions shows that numerous species have been found exclusively in the southwest Pacific, and are probably endemic to this area. In total, 116 species appear to occur only in the southwest Pacific, whereas 5 species are also found in Japan, and 3 species are widespread in the tropical Indo- Pacific. It is possible that this high number of endemics is an artefact caused by the absence of a comparable collecting effort in other regions of the tropical Indo-Pacific. However, a more detailed look at the different areas sampled, reveals that 67 species are found exclusively in the area including the Coral Sea, New Caledonia, Vanuatu, Fiji, Tonga and Wallis and Futuna, and only 10 species are found exclusively in the Philippines and Indonesia region. The number of species shared between the Coral Sea, New Caledonia, Vanuatu, Fiji, Tonga and Wallis and Futuna region and the Philippines and Indonesia region is only 18, and only 2 species are shared between all these areas and Japan (a relatively well studied area). Six widespread species are found exclusively in deep waters, but 3 of them are dubious records, based on a single record outside the New Caledonia region.

From these numbers it appears that there is a high degree of endemism in the deep sea fauna in the southwest Pacific. It also appears that there is some sort of isolation between the Coral Sea, New Caledonia, Vanuatu, Fiji, Tonga and Wallis and Futuna region and the Philippines and Indonesia region, which is reflected in the small number of species shared between these two areas.

BATHYMETRIC RANGES Valdés (2001a, b) found a clearly distinct fauna of deep sea nudibranch dorids in the southwest Pacific, with 30 new species and only 4 (13% of the total) previously described. These 30 new species are apparently only found in the deep sea (below 100 m) and some of them have adaptations to the deep sea environment, such as white colour and absence of eyes. The 4 previously described species also occur in shallow waters (above 100 m). The “cephalaspidean”

heterobranch fauna here studied is very different. Of the 121 species found, 26 (21%) have previously been described or reported from shallow waters above 100 m and the rest are either unidentified or have only been found in deep waters.

Valdés (2001a, b) also found restricted bathymetric ranges for most species of dorids, which are normally found within a 200 m range or less. The “cephalaspidean” heterobranch species here studied have much larger bathymetric ranges, with many species having ranges of 500 m or more, up to 1000 m (including records for empty shells). Only 40 species have ranges narrower than 200 m, but many of them are uncommon and known only from one or a few lots. Further sampling in this area may result in larger known ranges for some of these species.

There are two hypotheses that could explain the large bathymetric ranges of “cephalaspidean” heterobranchs compared to dorid nudibranchs:

— “Cephalaspidean” heterobranchs could have more adaptability than nudibranchs to survive at different depths.

The fact that most dorid nudibranchs have strict feeding habits, on one or a few species of sponges or other inver-

tebrates, may restrict their real bathymetric ranges to the ranges of their prey. Most “cephalaspidean” heterobranchs

found in deep waters feed on a variety of prey belonging to different groups of invetebrates or protozooans, or are

detritivorous. Therefore these species are potentially able to survive at a wider range of depths, where sources of

food are available.

— Because most of the material collected consists of empty shells, it is not possible to conclude that the animals

actually live at the depths sampled. Transport of dead shells by currents or gravity would produce artefacts in the

known bathymetric range of shell-bearing species, especially in areas with a complex orography. When only the

bathymetric ranges of specimens collected alive are considered, there is a much clearer picture, with 35 out of 41

species having ranges narrower than 200 m.

Figure 103 is a graphic representation of the bathymetric ranges of the species here studied. Valdés (2001b) found evidence for a considerable species turnover in the dorid nudibranch fauna: of the 5 species found at more than one station in the 200-400 m depth interval, just one was recorded from deeper than 500 m and another from shallower than 100 m. Conversely, of the 5 species found at more than one station in the 500-1000 m interval, 2 were recorded from shallower than 500 m. For the species here studied there appears to be a turnover at about 1000-1200 m, but the 200-400 m barrier is much more blurred than in the case of dorid nudibranchs. There are 21 species of “cephalaspideans” here examined whose range stops between 1000-1200 m. There are also 12 species that cross this boundary, but most of them (with one exception) have extremely wide bathymetric ranges (800 m or more), and only one of them is based on specimens collected alive, so transport of empty shells could be involved in producing this pattern. There are also 14 species found exclusively at depths greater than 1000-1200 m. The 200-400 m turnover could be masked by shell transport. There are numerous species that cross this boundary, but when only specimens collected alive are considered there is a sharp boundary at 200 m. There is only one species that is found above 200 m and below 400 m, Scaphander japonicus , the range of which stops at 452 m. All other species collected alive above 200 m have not been found alive below this depth. Conversely, all other species found alive below 200 m have not been collected above this depth. It appears that the “cephalaspidean” heterobranch fauna below 200 m is clearly distinct from the shallow water fauna. Some of these shallow water species have been included in the study because living specimens were found below 100 m, but none of them seem to occur below 200 m. Also, several other species collected below 100 m, but only as empty and damaged shells, were excluded from the study. These are: Pupa solidula (Linné, 1758) , Pupa strigosa (Gould, 1859) , Maxacteon fabreanus (Crosse, 1874) , Bullina lineata (Gray, 1825) , Bullina melior Iredale, 1926 , Bullina nobilis Kuroda in Habe, 1950, Bullina callizona Sakurai & Habe in Habe, 1961, Haminoea cf. nigropunctata Pease, 1868, Haminoea sp. , Atys naucum (Linné, 1758) , Atys cylindrica (Helbling, 1779) , Atys semistriata Pease, 1860 , Atys sp. , Mnestia bizona (A. Adams, 1850) , Liloa curta (A. Adams in Sowerby, 1850), Limulatys tortuosa (A. Adams in Sowerby, 1850), Limulatys debilis (Pease, 1860) , Limulatys perforata (Thiele, 1925) , Limulatys sp. , Diniatys cf. monodonta (A. Adams in Sowerby 1850), Diniatys sp. 1 , Diniatys sp. 2 , Bulla cf. vernicosa Gould, 1859 and Bulla sp.

FIG. 103. Bathymetric range of the species studied in this paper.Solid black bar: data from the present study based on specimens collected alive; solid gray bar: data from the present study based on empty shells; dotted bar: data from the literature.

HERBIVOROUS VS. CARNIVOROUS

The deep sea fauna contains a large number of detritivorous species, a smaller number of carnivores and no herbivores below depths where algae are not able to survive. In the present study, some species of Haminoeidae were collected at greater depths than expected. For instance, several common shallow water species of Haminoea and Atys were collected below 500 m, but all as empty shells only and therefore these records are considered to be caused by transportation by currents or other factors.

Only one species of Haminoeidae , Micratys wareni n. sp., was found alive at great depths. The anatomy of Micratys has been described in the present study for the first time and there are questions regarding the systematic placement of this taxon. The placement of Micratys in Haminoeidae has been exclusively based on shell characters. Considering the range of this species it is most likely that species of Micratys are either detritivorous or carnivorous. A phylogenetic analysis of this group is necessary to determine its evolutionary relationships and correct systematic placement.

“CEPHALASPIDEAN” HETEROBRANCH DEEP SEA DIVERSITY

The MUSORSTOM expeditions produced a total of 34 species of dorid nudibranchs from deep waters in the southwest Pacific (Valdés 2001a, b), which is a small number compared to the 121 species of “cephalaspideans” (excluding Ringiculidae ) collected during the same expeditions and here studied. This proportion of almost 4: 1 contrasts with the faunal composition of shallow waters in this region. From the Koumac expedition in northwest New Caledonia, Bouchet et al. (2002) documented 98 species of shallow water “cephalaspideans” (including Ringiculidae ) and a similar number of species of dorid nudibranchs (proportion 1: 1). For the Great Barrier Reef, Marshall & Willan (1999) listed 75 species of shallow water “cephalaspideans” (including Ringiculidae ) and 107 species of dorid nudibranchs (proportion 1: 1.5) and Gosliner (1987) listed 60 species of shallow water “cephalaspideans” (including Ringiculidae ) and 151 dorid nudibranchs in South Africa (proportion 1: 2.5). These data suggest that “cephalaspideans” are comparatively more successful in the deep sea than dorid nudibranchs, which are the most diverse group of opisthobranchs in shallow waters. However, because most species of “cephalaspideans” have shells, it is likely that by using dredging as the main collecting method more species of “cephalaspideans” would be found, whereas in shallow-water sampling more soft-bodied and brightly coloured species would be collected, most of which would be destroyed using dredging. If the high proportion of “cephalaspideans” in the deep sea is an artefact, we would expect the diversity of dorid nudibranchs in the deep sea to be much larger than is currently known. Bouchet et al. (2002) used a variety of indirect collecting techniques as well, including underwater aspirators, breaking and brushing hard substrates, and trawling, which also produce empty shells, and that may account for the higher proportion of “cephalaspideans” versus dorids found, compared to the Great Barrier Reef and South Africa.

ACKNOWLEDGEMENTS

Philippe Bouchet (MNHN) made available the southwest Pacific material examined in this study, which was collected with the financial support of the MNHN and the IRD (formerly ORSTOM) during the MUSORSTOM expeditions. Specimens were collected by Bertrand Richer de Forges, Philippe Bouchet, Anders Warén, Pierre Lozouet and Bernard Métivier.

Virginie Héros (MNHN) and Lindsey Groves (LACM) provided assistance with the curation and processing of the material. Matthias Glaubrecht (MNHB), Robert Moolenbeek (ZMA) and Amelia Campbell (BMNH) made available the type material from the Valdivia , Siboga and Challenger expeditions.

Robert H. Cowie edited the manuscript and made numerous constructive comments.

Nancy Smith (LACM volunteer) provided invaluable assistance during the preparation of the manuscript. Paul Valentich-Scott, Patricia Sadeghian and the staff of the Santa Barbara Museum of Natural History provided assistance with the digital photography of the specimens studied.

The geographic range maps were prepared using the freeware software OMC 4.1 (Online Map Creation) by Martin Weinelt.

The Scanning Electron Microscopy work was conducted at the LACM facility, with financial support from the US National Science Foundation (DBI - 0216506).

REFERENCES

ADAMS A. 1862. — On some new species of Cylichnidae, Bullidae and on the Mollusca dredged by the Blake in 1880, including descriptions of

Philinidae , from the Seas of China and Japan. Annals and Magazine of several new species. Bulletin of the Museum of Comparative Zoology 23:

Natural History (3) 9: 150-161. 199-244, 2 pls.

ADAMS H. & ADAMS A. 1854-1858. — The genera of recent Mollusca; CHABAN E. M. 1996. — Opisthobranch molluscs of the family Diaphanidae

arranged according to their organization. Volume 2. Van Voorst, London, ( Gastropoda, Opisthobranchia ) of the Russian Seas. Ruthenica 6: 127-

661 p. [Dates of publication: p. 1- 92 (1854), p. 93-284 (1855), p. 285- 148. [In Russian]

412 (1856), p. 413-540 (1857), p. 541-661 (1858)]. COSSMANN M. 1889. — Catalogue illustré des coquilles fossiles de l’Éocène

ASCANIUS P. 1772. — Philine quadripartita, ein sonst unbekanntes Seethier des environs de Paris. Annales de la Société Royale Malacologique de

abgezeichnet und beschrieben. Kongliga Svenska Vetenskaps Akademiens Belgique 24: 3-385, pls 1-12.

Handlingar 34: 325-327. COSSMANN M. 1895. — Essais de Paléoconchologie comparée. Livraison 1

BABA K. & TOKIOKA T. 1965. — Two more new species of Gastropteron from [Tectibranchia, Nucleobranchiata, Pulmonata (Thalassophila) et Annexe].

Japan,with further notes on G.flavum T.& B.( Gastropoda: Opisthobranchia ). Chez l’auteur, Paris. 159 p, 7 pls.

Publications of the Seto Marine Biology Laboratory 12: 363-378, 1 pl. CROSSE J. C. H. 1885 [review of]. — Nomenclatura generica e specifica di

BELLARDI L. 1876. — Descrizione di un nuovo genera della famiglia delle alcune Conchiglie Mediterranee,pel marchese di (Nomenclature générique

Bullide fossile del terreno pliocenico inferiore del Piemonte e della Liguria. et spécifique de quelques coquilles méditerranéennes, par le marquis de

Bullettino della Società Malacologica Italiana 2: 207-210. Monterosato. Palerme, 1884. Journal de Conchyliologie 33: 139-141.

BERGH R. 1905. — Die Opisthobranchiata der Siboga-Expedition, in WEBER CUVIER G. L. 1810.— Mémoire sur les Acères,ou Gastéropodes sans tentacules

M. (ed.), Uitkomsten op Zoologisch, Botanisch, Oceanographisch en apparens. Annales du Muséum National d’Histoire Naturelle 16: 1-18.

Geologisch Gebied verzameld in Nederlandsch Oost-Indië 1899-1900 DALL W. H. 1871. — Descriptions of sixty new forms of mollusks from the

aan boord H.M. Siboga onder commando van Luitenant ter zee Iekl. G.F. west coast of North America and the north Pacific Ocean, with notes on

Tydeman. Volume 50. Brill, Leiden: 1-248, 20 pls. others already described. American Journal of Conchology 7: 93-160,

BLAINVILLE H. M. DE. 1825. — Manuel de Malacologie et de Conchyliologie. pls 13-16.

Levrault, Paris, 664 p., 87 pls. DALL W.H. 1881.— Reports on the results of dredging,under the supervision

BOUCHET P. 1975.— Opisthobranches de profondeur de l’Océan Atlantique. of Alexander Agassiz, in the Gulf of Mexico, and in the Caribbean Sea,

I.- Cephaslaspidea. Cahiers de Biologie Marine 16: 317-365, pls 1-4. 1877 -79, by the United States Coast Survey Steamer Blake, Lieutenant-

BOUCHET P., LOZOUET P., MAESTRATI P. & HÉROS V. 2002. — Assessing the Commander C. D. Sigsbee, U. S. N., and Commander J. R. Bartlett, U. S.

magnitude of species richness in tropical marine enviroments:Exceptionally N., commanding. XV. Preliminary report on the Mollusca. Bulletin of the

high numbers of molluscs at a New Caledonia site. Biological Journal of Museum of Comparative Zoology 9: 32-144.

the Linnean Society 75: 421-436. DALL W. H. 1882. — American work on recent Mollusca in 1881. American

BOUCHET P., HÉROS V., LOZOUET P. & MAESTRATI P. 2008. — A quater- Naturalist 16: 874-887.

century of deep-sea malacological exploration in the South and West DALL W. H. 1889. — Reports on the results of dredging, under the supervision

Pacific:Where do we stand? How far to go?, in HÉROS V., COWIE R. H. & of Alexander Agassiz,in the Gulf of Mexico (1877 -78) and in the Caribbean Sea (1879-80),by the United States Coast Survey Steamer Blake, Lieutenant-

BOUCHET P. (eds), Tropical Deep-Sea Benthos 25. Mémoires du Muséum Commander C. D.Sigsbee, U.S. N., and Commander J.R. Bartlett, U. S. N.,

national d’Histoire naturelle 196: 9-40. commanding.XXIX Report on the Mollusca,Pt.2: Gastropoda and Scaphopoda.

BRAZIER J. 1878. — Continuation of the Mollusca collected during the Bulletin of the Museum of Comparative Zoology 18: 1-492, pls 10-40.

Chevert Expedition. Proceedings of the Linnean Society of New South DALL W.H. 1890.— Contributions to the Tertiary fauna of Florida,with especial

Wales 2: 74-89. reference to the Miocene silex-beds of Tampa and the Pliocene beds of the

BROWN T. 1827.— Illustrations of the conchology of Great Britain and Ireland. Caloosahatchie River. Part I. Pulmonate, opisthobranchiate and orthodant

Drawn from nature. Lizards, Edinburgh, 144 p, 59 pls. gastropods. Transactions of the Wagner Free Institute of Science 3: 1-200.

BRUGUIÈRE J. G. 1789. — Encyclopédie Méthodique. Histoire naturelle des DALL W.H. 1895.— No.XXXIV. Scientific results of explorations by the U.S.Fish

vers. Volume 1. Panckoucke, Paris, 757 p. Commission Steamer Albatross. Report on the Mollusca and Brachiopoda

BULLIS H. R. 1956. — The genus Scaphander in the Gulf of Mexico and notes dredged in deep water, chiefly near the Hawaiian Islands,with illustrations

on the western Atlantic species. Bulletin of Marine Science of the Gulf

of hitherto unfigured species from Northwest America. Proceedings of the

and Caribbean 6: 1-17. United States National Museum 17: 675-733, pls 23-32.

BUSH K.J. 1893.— Reports on the results of dredging,under the supervision DALL W.H. 1896.— Diagnoses of new Tertiary fossils from the southern United

of Alexander Agassiz,in the Gulf of Mexico (1877-78),and in the Caribbean States. Proceedings of the United States National Museum 18: 21-46.

Sea (1879-80),and along the Atlantic coast of the United States (1880),by DALL W.H.1902.— Illustrations and descriptions of new,unfigured,or imperfectly

the U.S. coast survey steamer Blake, Lieutenant-Commander C.D.Sigsbee, known shells,chiefly American, in the U.S. National Museum. Proceedings

U. S. N., and Commander J. R. Bartlett, U. S. N., commanding. Report

of the United States National Museum 24: 499-566, pls 27-40. DALL W.H. 1908.— Reports on the dredging operations off the west coast of HABE T. 1952b. — Atydae in Japan, in KURODA T. (ed.), Illustrated catalogue Central America to the Galapagos,to the west coast of Mexico,and in the of Japanese shells. Volume 20. Malacological Society of Japan, Tokyo: Gulf of California, in charge of Alexander Agassiz, carried on by the U.S. 137-151.

Fish Commission Steamer Albatross, during 1891,Lieut.-Commander Z.L. HABE T. 1954. — Report on the Mollusca chiefly collected by the S.S. Soyo- Tanner, U.S. N.,commanding,XXXVII.Reports on the scientific results of the Maru of the Imperial Fisheries Experimental Station on the continental expedition to the eastern tropical Pacific,in charge of Alexander Agassiz, shelf bordering Japan during the years 1922-1930.Part 1. Cephalaspidea . by the U.S. Fish Commission Steamer Albatross, from October, 1904, to Publications of the Seto Marine Biological Laboratory 3: 301-318, pl 38. March,1905,Lieut.-Commander L.M. Garrett, U.S. N., commanding,XIV. HABE T. 1955. — A list of the cephalaspid Opisthobranchia of Japan. Bulletin The Mollusca and the Brachiopoda. Bulletin of the Museum of Coparative of the Biogeographical Society of Japan 16-19: 54-79, pl 4.

Zoology 43: 205-487, pls 1-22. HABE T. 1958. — On the shell-bearing opisthobranchiate molluscan fauna DAUTZENBERG P.1929.— Mollusques testacés marins de Madagascar. Faune from off Choshi, Chiba Pref., Japan. Annotationes Zoologicae Japonenses des Colonies FranÇaises 3: 1-636. 31: 117-120.

DE GEER G.1910.— Quaternary sea-bottoms in Western Sweden. Geologiska HABE T. 1961 . — Coloured illustrations of the shells of Japan. Hoikusha, FÖreningens i Stockholm FÖrhandlingar 32: 1339. Osaka, 183 p. [In Japanese] .

DRAPARNAUD J.P.R.1800.— Observations sur la Gioenia . Journal de Physique, HABE T. 1976. — Two new striped Punctacteon (Mollusca) from Japan and de Chimie, d’Histoire Naturelle et des Arts 50: 146. the Philippines. Bulletin of the National Science Museum (A - Zoology) FÉRUSSAC A. E. J. DE. 1821-22. — Tableaux Systématiques des Animaux 2: 7-9.

Mollusques, classés en familles naturelles, dans lesquels où est établi la HARRY H. W. 1967. — A review of the living tectibranch snails of the genus concordance de tous les systèmes. Bertrand, Paris, xlvii, i + 27 p + 110 p Volvulella , with descriptions of a new subgenus and species from Texas. [Dates of publication: i-xlvii, i (1822), 1-27 + 1-110 (1821)]. The Veliger 10: 133-147.

FÉRUSSAC A.E.J.DE.1822.— Bulline, in AUDOUIN J.V.et al.(eds.), Dictionnaire HEDLEY C. 1899. — The Mollusca of Funafuti.Part I.- Gasteropoda. Memoirs classique d’Histoire Naturelle. Volume 2. Rey et Gravier, Paris: 578-579. of the Australian Museum 3: 395-488.

FISCHER P. 1880-87. — Manuel de Conchyliologie et de Paléontologie HERRMANNSEN A. N. 1846-52.— Indicis generum Malacozoorum primordia. Conchyliologique, ou histoire naturelle des mollusques vivants et fossiles. Nomina subgenerum, generum, familiarum, tribuum, ordinum, classium; Savy,Paris,1369 p., 23 pls [Dates of publication:pp 1-112 (1880),pp 113- adjectis auctoribus, temporibus, locis systematicis atque literariis, etymis, 304 (1881), pp 305-416 (1882), pp 417-608 (1883), pp 609-688 (1884), synonymis. Praetermittuntur Cirripeda, Tunicata et Rhizopoda. Fischeri, pp 689-896 (1885), pp 897-1008 (1886), pp 1009-1369 (1887)]. Kasel,xxvii, 637 p. [Vol. 1] + 717 p. [Vol. 2] + v, 140 p. [suppl.]. [Dates of GABB W.M.1872.— Description of some new genera of Mollusca. Proceedings publication: Vol. 1 (1846), Vol. 2 (1847-49), suppl. (1852)].

of the Academy of Natural Sciences of Philadelphia 24: 270-274. HIGO S., CALLOMON P. & GOTO Y. 2001. — Catalogue and bibliography GIOENI G.1783.— Descrizione di una nuova famiglia e di un nuovo generre di of the marine shell-bearing mollusca of Japan. Gastropoda, Bivalvia, testacei trovati nel littorale di Catania. Author’s edition, Naples, 34 p., 1 pl. Polyplacophora, Scaphopoda. Type Figures. Elle Scientific Publications, GOSLINER T. M. 1979. — A review of the systematics of Cylichnella Gabb Osaka-fu, 208 p.

( Opisthobranchia : Scaphandridae ). The Nautilus 93: 85-92. HORI S. 2000a. — Family Acteonidae , in OKUTANI T. (ed.), Marine Mollusks GOSLINER T. M. 1980. — Systematics and phylogeny of the Aglajidae in Japan. Tokai University Press, Tokyo: 732-735.

( Opisthobranchia : Mollusca). Zoological Journal of the Linnean Society HORI S. 2000b. — Family Cylichnidae , in OKUTANI T. (ed.), Marine Mollusks 68: 325-360. in Japan. Tokai University Press, Tokyo: 740, 741-745, 746.

GOSLINER T.M.1987.— Nudibranchs of Southern Africa. A guide to opsithobranch HORI S. 2000c . — Family Retusidae , in OKUTANI T. (ed.), Marine Mollusks in molluscs of Southern Africa. Sea Challengers, Monterey , 136 p. Japan. Tokai University Press , Tokyo: 746-748 .

GOSLINER T.M.1988.— The Philinacea (Mollusca: Gastropoda: Opisthobranchia ) HORI S. 2000d. — Family Bullinidae / B. ovulina , in OKUTANI T. (ed.), Marine of Aldabra Atoll, with descriptions of five new species and a new genus. Mollusks in Japan. Tokai University Press, Tokyo: 754-757, 759.

Bulletin of the Biological Society of Washington 8: 79-100. ICZN [INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE]1954. GOSLINER T. M. 1989. — Revision of the Gastropteridae ( Opisthobranchia : — Opinion 287.Validation,under the plenary powers,of the generic name Cephalaspidea ) with descrptions of a new genus and six new species. The “ Scaphander ” Montfort, 1810 (class Gastropoda, order Tectibranchiata ). Veliger 32: 333-381. Opinions and Declarations rendered by the International Commission on GOSLINER T. M. 1995. — Introduction and spread of Philine auriformis Zoological Nomenclature 8: 4962.

( Gastropoda: Opisthobranchia ) from New Zealand to San Francisco Bay ICZN [INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE] and Bodega Harbor. Marine Biology 122: 249-255. 1974.— Opinion 1030. Cylindrella Swainson,1840 (Mollusca, Gastropoda): GOULD A. A. 1859. — Descriptions of shells collected in the North Pacific suppressed under the Plenary Powers. Bulletin of Zoological Nomenclature Exploring Expedition under Captains Ringgold and Rodgers. Proceedings 31: 190-191.

of the Boston Society of Natural History 7: 138-142. ICZN [INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE]1977. GRAY J.E.1847.— A list of the genera of recent Mollusca,their synonyma and — Opinion 1079. Aglaja Renier, [1807] , A. depicta Renier, [1807] and A. types. Proceedings of the Zoological Society of London 15: 129-219. tricolorata Renier,[1807] (Mollusca: Gastropoda) rendered available under HABE T. 1950a. — Ringiculidae and Retusidae (1) in Japan, in KURODA T. the plenary powers. Bulletin of Zoological Nomenclature 34: 16-20.

(ed.), Illustrated catalogue of Japanese shells. Volume 2. Malacological ICZN [INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE]1999. Society of Japan, Tokyo: 17-24 . — International code of zoological nomenclature, 4th edition.International HABE T.1950b.— Philinidae in Japan, in KURODA T.(ed.), Illustrated catalogue of Trust for Zoological Nomenclature , London , xxix + 306 p.

Japanese shells. Volume 8. Malacological Society of Japan, Tokyo: 48-52. IREDALE T.1918.— Molluscan nomenclatural problems and solutions.- No.1. HABE T.1952a.— Descriptions of new genera and species of the shell-bearing Proceedings of the Malacological Society of London 13: 28-40.

opisthobranchiate molluscs from Japan ( Cephalaspidea, Tectibranchia ). IREDALE T.1925.— Mollusca from the continental shelf of eastern Australia. Venus 17: 69-77 [In Japanese]. Records of the Australian Museum 14: 243-270, pls 41-43, map.

IREDALE T.1926.— Strange molluscs in Sydney Harbour. Australian Zoologist MENKE K. T. 1854. — Zur Familie Bullacea und deren Gattungen und Arten. 5: 337-352, pls 37-38. Malakozoologische Blätter 1: 33-48.

IREDALE T.1930. — Some notable name changes. Australian Zoologist 6: 175. MIKKELSEN P. M. 1996. — The evolutionary relationships of Cephalaspidea IREDALE T.1936 .— Australian Molluscan Notes.No.2. Records of the Australian s. l. ( Gastropoda: Opisthobranchia ): a phylogenetic analysis. Malacologia Museum 19: 267-340, pls 20-24. 37: 375-442.

JEFFREYS J.G.1883.— Mediterranean Mollusca (No.3) and other Invertebrata. MIKKELSEN P. S. & MIKKELSEN P. M. 1984. — Comparison of Acteocina Annals and Magazine of Natural History (5) 11: 393-401. canaliculata (Say, 1826) , A. candei (d’Orbigny, 1841) , and A. atra spec. KOSSE J.F.J.1813.— De Pteropodum ordine et nova ipsius genera. Hendelianis, nov. ( Gastropoda: Cephalaspidea ). The Veliger 27: 164-192.

Hallae, 16 p., 1 pl. MONTEROSATO T.A. DI.1884.— Nomenclatura generica e specifica di alcune KURODA T. & HABE T. 1952. — Check list and bibliography of the recent conchiglie mediterranee. Virzì, Palermo, 152 p.

marine Mollusca of Japan. Stach , Tokyo, 210 p. MONTEROSATO T. A. DI. 1890 . — Conchiglie delle profondità del mare di KURODA T. & HABE T. 1954 . — On some Japanese Mollusca described by Palermo (Part 3). Il Naturalista Siciliano 9: 181-191.

A. Adams, whose specimens are deposited in the Redpath Museum of MONTFORT P. D. de. 1810. — Conchyliologie systématique, et classification Canada. Venus 18: 1-16, pls 1-2. méthodique des coquilles; offrant leurs figures,leur arrangement générique, KURODA T. & HABE T. 1955. — A list of the cephalaspid opisthobranchiate leurs descriptions caractéristiques, leurs noms; ainsi que leur synonymie molluscs from Japan. Bulletin of the Biogeographical Society of Japan en plusieurs langues. Volume 2. Schoell, Paris, 676 p.

16-19: 54-79, pl. 4. MÜLLER O. F. 1776. — Zoologiae Danicae prodromus, seu Animalium Daniae KURODA T., HABE T. & OYAMA K. 1971. — The sea shells of Sagami Bay et Norvegiae indigenarum characteres, nomina, et synonyma imprimis collected by his Majesty the Emperor of Japan. Maruzen , Tokyo, 489 + popularium. Hallageriis , Havnioe , xxxii + 282 p.

51 p., 121 pls. NEWTON R.B. 1891.— S ystematic list of the Frederick E.Edwards collection of LAMARCK J. B. 1801. — Système des animaux sans vertèbres, ou tableau British Oligocene and Eocene Mollusca in the British Museum (Natural History), général des classes, des ordres et des genres de ces animaux. Deterville, with references to the type-specimens from similar horizons contained in Paris, 232 p. other collections belonging to the Geological department of the Museum. LAMARCK J. B. 1816. — Mollusques et Polypes divers, in BRUGUIÈRE J. G. British Museum (Natural History), London, xxviii + 365 p., 1 tab.

(ed.), Tableau encyclopédique et méthodique des Trois Règnes de la Nature. NOMURA S. 1935. — Catalogue of the Tertiary and Quaternary Mollusca Contenant l’helminthologie, ou les vers infusoires, les vers intestins,les vers from the Island of Taiwan (Formosa) in the Institute of Geology and mollusques, & c. Volume 4, Part 23. Agasse, Paris: 1-16, pls 391-488. Paleontology,Tôhoku Imperial University,Sendai, Japan.Part 2, Scaphopoda LEACH W. E. 1847. — The classification of the British Mollusca. Annals and and Gastropoda. Science Reports of the Tôhoku Imperial University 18: Magazine of Natural History 20: 267-273. 53-228, pls 6-10.

LEMCHE H. E. 1951. — Proposed use of the Plenary Powers to validate the NORDSIECK F.1972.— Die europäischen Meeresschnecken ( Opisthobranchia generic name “ Scaphander ” Montfort, 1810 (Class Gastropoda, Order mit Pyramidellidae ; Rissoacea). Vom Eismeer bis Kapverden, Mittelmeer Tectibranchiata ). Bulletin of Zoological Nomenclature 2: 35-36. und Schwarzes Meer. Fischer, Stuttgart, 327 p.

LEMCHE H. E. 1956. — The anatomy and histology of Cylichna ( Gastropoda ODHNER N. H. 1913. — Ptisanula limnaeoides , a new fossil and recent Tectibranchia). Spolia Zoologica Musei Hauniensis 16: 1-278, pls 1-46. opisthobranchiate mollusc. Geologiska FÖreningens i Stockholm FÖrhandlingar LIN G.-Y. & QI, Z.-Y. 1983. — A preliminary survey of the Cephalaspidea 35: 329-332.

( Opisthobranchia ) of Hong Kong and adjacent waters, in MORTON B. & ODHNER N.H.1914.— Ptisanula limnaeoides , a new Arctic opisthobranchiate DUDGEON D. (eds), Proceedings of the Second International Workshop mollusc, its anatomy and affinities. Arkiv fÖr Zoologi 7: 1-18, pl 1.

on the Malacofauna of Hong Kong and Southern China. Hong Kong OKEN L. 1815 . — Lehrbuch der Naturgeschichte. Dritter Theil.Zoologie Erste University Press, Hong Kong: 109-124. Abth. Fleischlose Thiere. Schmid und Comp., Leipzig, 45 p.

LOVÉN S. L. 1846. — Index Molluscorum littora Scandinaviae occidentalia OKUTANI T. 1987. — Description of a new gigantic scaphanderid gastropod, habitantium. Öfversigt af Kongl. vetenskaps-akademiens FÖrhandlingar Bucconia centa , from Koshiki Islet, West of Kyushu. Venus 45: 219-221. 3: 1-50. PEASE W.H.1860.— Descriptions of new species of Mollusca from the Sandwich MARCUS EV. 1972. — On some Acteonidae ( Gastropoda, Opisthobranchia ). Islands. Proceedings of the Zoological Society of London 28: 18-36.

Papéis Avulsos de Zoologia 25: 167-188. PHILIPSSON L. M. 1788. — Dissertatio historico naturalis sistens testaceorum MARCUS EV.1974.— On some Cephalaspidea ( Gastropoda: Opisthobranchia ) genera. Lundae, Berlingianis, 23 p.

from the western and middle Atlantic warm waters. Bulletin of Marine PILSBRY H.A. 1893.— Manual of conchology; structural and systematic, with Science 24: 300-371. illustrations of the species. Polyplacophora (Chitons), Acanthochitidae, MARCUS EV. 1977 . — On the genus Tornatina and related forms. Jounal of Cryptoplacidae and Appendix. Tectibranchiata . Volume 15. Academy of Molluscan Studies supplement 2: 1-35. Natural Sciences, Philadelphia, 436 p., 61 pls.

MARCUS EV.& MARCUS ER.1960.— Opisthobranchs from American Atlantic PILSBRY H. A. 1895-96. — Manual of conchology; structural and systematic, warm waters. Bulletin of Marine Science of the Gulf and Caribbean 10: with illustrations of the species. Philinidae , Gastropteridae , Aglajidae , 129-203. Aplysiidae , Oxynoeidae , Runcinidae , Umbraculidae , Pleurobranchidae . MARSHALL J.G.& WILLAN R.C. 1999.— Nudibranchs of Heron Island, Great Volume 16. Academy of Natural Sciences, Philadelphia, 262 p, 74 pls. Barrier Reef. A survey of the opisthobranchia (sea slugs) of Heron and [Dates of publication: p 1-112 (1895), p 113-262, i-vii (1896)].

Wistari Reefs. Backhuys, Leiden, 257 p. POWELL A. W. 1937. — New species of marine Mollusca from New Zealand. MARTENS E. C. VON 1879. — Übersicht der von ihm (W. Peters) von 1843 bis Discovery Reports 15: 153-222, pls 45-56.

1847 in Mossambique gesammelten Mollusca. Monatsberichte der KÖniglichen POWELL A. W. 1979.— New Zealand mollusca: Marine, land and freshwater Preussischen Akademie der Wissenschaften zu Berlin 1879: 727-749. shells. Collins, Auckland, xiv + 500 p.

MECKEL J.F.1809.— Über ein neues Geschlecht der Gasteropoden.Beyträge PRUVOT-FOL A. 1954. — Mollusques Opisthobranches. Faune de France 58: zur vergleichenden Anatomie, Part 1, Volume 2. Reclam, Leipzig, 14 p. 1-460, pl 1.

QUOY J. & GAIMARD J. 1832-33. — Zoologie, in DUMONT D’URVILLE J. S. des vers testacés avec XXII planches. Schultz, Copenhague, 287 p., C. (ed.), Voyage de découvertes de l’Astrolabe exécuté par ordre du Roi, 22 pls.

pendant les années 1826-1827-1828-1829,sous le commandement de M. SOWERBY G. B. 1839.— A conchological manual . Author’s Edition, London, J.Dumont d’Urville. Volume 2 & Atlas.Tastu,Paris:1-686, pls 1-26. [Dates 130 p., 530 figs [in 23 unnumbered pls], 2 tabs.

of publication: p 1-320 (1832), 321-686 (1833), pls 1-26 (1833)]. SOWERBY G. B. 1848-50. — Thesaurus Conchyliorum , or monographs of RAFINESQUE C. S.1814.— Précis des découvertes et travaux somiologiques ou genera of shells. Volume 2. Sowerby, London, 460 p., 94 pls. [Dates of Choix raisonné de ses principales découvertes en zoologie et en botanique, publication:pp.493-505,pls 92-108 (1848), pp.507-552,pls 109-118bis pour servir d’introduction à ses ouvrages futurs. Royale Typographie (1849), pp. 553-899, pls 119-186 (1850)].

Militaire, Palermo, 55 p. STREBEL H.1905.— Beiträge zur Kenntnis der Molluskenfauna der Magalhaen- REEVE L. A. 1842. — Monograph of the genus Tornatella , a small group of Provinz.No.3 Zoologische Jahrbücher,Abteilung für Systematik,Geographie pectinibranchiate mollusks of the family Plicacea , including descriptions und Biologie der Tiere 22: 575-666, pls 21-24.

of seven new species from the collection of H. Cuming, Esq. Proceedings SUTER H. 1913. — Manual of the New Zealand Mollusca with an atlas of of the Zoological Society of London 10: 58-62. quarto plates. Government Printer, Wellington: xxiii + 1120 p. [Atlas, RENIER S. A. 1807. — Tavole per servire alla classificazione e connoscenza published 1915, 72 pls.].

degli Animali. Padova, 8 pls. SWAINSON W.1840.— A treatise on Malacology; or the natural classification RISSO A. 1826. — Histoire Naturelle des principales productions de l’Europe of shells and shell fish. Longman, London, 419 p.

Méridionale et particulièrement de celles des environs de Nice et des Alpes TAKI I. 1956.— Japonacteon , a new genus of Pupidae. Bulletin of the National Maritimes. Volume 4. Levrault, Paris, 439 p., 12 pls. Science Museum, Tokyo 3: 47-51.

RUDMAN W. B. 1971a. — On a new genus for “ Tornatina ” murdochi Suter, THIELE J. 1903 . — Anatomisch-systematische Untersuchungen einiger 1913 ( Retusidae , Opisthobranchia ). Journal of the Malacological Society Gastropoden, in MARTENS E. C. von (Ed.), Wissenschaftliche Ergebnisse of Australia 2: 187-193. der Deutschen Tiefsee-Expedition auf dem Damfper Valdivia 1898 -1899. RUDMAN W. B. 1971b. — The genus Bullina ( Opisthobranchia , Gastropoda) Volume 7, Issue 2. Fischer, Jena, 180, pls 1-9.

in New Zealand. Journal of the Malacological Society of Australia 2: 195- THIELE J. 1925. — Gastropoda der Deutschen Tiefsee-Expedition, II. Part, 203. in CHUN C. (ed.), Wissenschaftliche Ergebnisse der Deutschen Tiefsee- RUDMAN W.B.1971c.— The family Acteonidae ( Opisthobranchia , Gastropoda) Expedition auf dem Damfper Valdivia 1898 -1899. Volume 17, Issue 2. in New Zealand. Journal of the Malacological Society of Australia 2: 205- Fischer, Jena: 35-382, pls 1-34.

214. THOMPSON T.E. 1976. — Biology of opisthobranch molluscs.Volume 1. The RUDMAN W. B. 1972a. — The genus Philine ( Opisthobranchia , Gastropoda). Ray Society, London, 206 p.

Proceedings of the Malacological Society of London 40: 171-187. VALDÉS A. 2001a. — Phyllidiid nudibranchs (Mollusca) from the deep south- RUDMAN W. B. 1972b. — Studies on the primitive opisthobranch genera western Pacific Ocean, in BOUCHET P. & MARSHALL B. A. (eds) Tropical Bullina Férussac and Micromelo Pilsbry. Zoological Journal of the Linnean Deep-Sea Benthos 22. Mémoires du Muséum National d’Histoire Naturelle Society 51: 105-119. 185: 331-369.

RUDMAN W. B. 1974. — A comparison of Chelidonura , Navanax and Aglaja VALDÉS A.2001b.— Deep-sea dorid nudibranchs (Mollusca, Opisthobranchia ) with other genera of the Aglajidae ( Opisthobranchia : Gastropoda). Zoological from the tropical West Pacific, with descriptions of two new genera and Journal of the Linnean Society 54: 185-212. nineteen new species. Malacologia 43: 237-311.

RUDMAN W. B. 1998a. — Comment on the identity of Californian Philine VALDÉS A. & HÉROS V. 1998. — The types of recent and certain fossil auriformis . Available at: http://www.seaslugforum.net/philunk.htm opisthobranch molluscs in the Muséum national d’Histoire naturelle,Paris. RUDMAN W. B. 1998b. — Philine orientalis? A. Adams, 1854. Available at: Zoosystema 20: 695-742.

http://www.seaslugforum.net/philhk.htm WATSON R. B. 1883. — Mollusca of H.M.S. Challenger Expedition.- Part XX. SARS M. 1870. — Bidrag til Kundskab om Christianiafjordens Fauna. Dahl, Journal of the Linnean Society, Zoology 17: 341-346.

Christiania [Oslo], 114 p., 13 pls. WATSON R.B.1886.— II - Report on the Scaphopoda and Gasteropoda collected by SCHEPMAN M.M.1913.— The Prosobranchia,Pulmonata and Opisthobranchia H.M.S. Challenger during the years1873-1876, in THOMSON C.W.& MURRAY tectibranchiata (tribe Bullomorpha ) of the Siboga Expedition,Pt. 6 Pulmonata J.(ed.), Report on the scientific results of the voyage of the H.M.S.Challenger and Opisthobranchia tectibranchiata,tribe Bullomorpha , in WEBER M.(ed.), during the years1873-76 under the command of Captain George S. Nares , R.N., Uitkomsten op Zoologisch, Botanisch, Oceanographisch en Geologisch F.R.S. and the late Captain Frank Tourle Thompson , R. N. prepared under the Gebied verzameld in Nederlandsch Oost-Indië 1899-1900 aan boord superintendence of the late Sir L. Wyville Thompson , Knt., F.R.S.& c. Volume H.M. Siboga onder commando van Luitenant ter zee Ie kl. G. F. Tydeman. 15: Zoology. H.M. Stationery Office, London: i-v, 1-756, pls 1-50, 1-3.

Volume 49. Brill, Leiden: 453-494, 2 pls. WINCKWORTH R.1932.— The British marine Mollusca. Journal of Conchology SCHIØTTE T. 1998. — A taxonomic revision of the genus Diaphana Brown , 19: 211-252.

1827, including a discussion of the phylogeny and zoogeography of the YOKOYAMA M. 1920. — Fossils from the Miura Peninsula and its immediate genus (Mollusca: Opisthobranchia ). Steenstrupia 24: 77-140. north. Journal of the College of Science, Imperial University of Tokyo 39: SCHUMACHER C. F. 1817. — Essai d’un nouveau système des habitations 1-193, pls 1-20.