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On the Anaspidea (Gastropoda: Opisthobranchia) of the Warm Waters of the Western Atlantic

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On the Anaspidea (Gastropoda: Opisthobranchia) of the Warm Waters of the Western Atlantic ON THE ANASPIDEA (GASTROPODA: OPISTHOBRANCHIA) OF THE WARM WATERS OF THE WESTERN ATLANTIC EVELINE D. B.-R. MARCUS Caixa Postal 6994, 01000 sao Paulo, Brazil ABSTRACT A key is given for the 16 western Atlantic species of Anaspidea and two Middle Atlantic ones. These are eight (and two) species of Aplysia, and Notarchus punctatus armatus, Stylocheilus longicauda, Stylocheilus citrinus, Bursatella leachii pleii, Dolabrifera dolabrifera, Petalifera petalifera bon- airei, Petalifera ramosa, and Phyllaplysia engeli. Descriptions are given for Petalifera p. bonairei and Petalifera ramosa. A key to the subgenera of the genus Aplysia is also furnished. Comparative notes are made on anatomical and systematic characters, sole, pallial cavity, shell, and nervous system. My subdivision of the Notarchidae leaves only Notarchus in the Notar- chinae; the remainder of the genera are allotted to the Dolabriferinae. Three species of Aplysia were dredged in 348-357 m depth. INTRODUCTION The present paper deals with seven western Atlantic genera and 16 species of the order Anaspidea. Taylor & Sohl (1962: 12) considered the order Anaspidea as consisting of the superfamily Aplysiacea, including Aplysiidae and Akeridae. However, the Akeridae are often allotted to the Cephalaspidea (Hoffmann, 1933: 308-309; Wirz, 1952: 167; Pruvot-Fol, 1954: 92; Marcus, 1967b: 613; 1970c: 939; and others). The strepto- neurous Akera (Guiart, 1901: 116; Marcus, 1970c: 938, fig. 32) really occupies an intermediate position between Cephalaspidea with cephalic shield and pallial caecum and the euthyneurous Anaspidea with numerous gizzard teeth. However, the argument that Akera has a head shield like the Cephalaspidea, while the Anaspidea have tentacles and rhinophores, favors the placing of Akera in the Cephalaspidea in any key. The Anaspidea can be separated by the length of the visceral loop of the nervous system into the suborder Longicommissurata (Pruvot-Fol, 1954: 95) with the family Aplysiidae, including the Aplysiinae and Dolabel- linae; and the Brevicommissurata (1954: 101) with the family Notarchidae, divided into Notarchinae and Dolabriferinae. In all six genera of the Brevicommissurata, the body may be beset with knobs or warts or branched papillae, but these protuberances can be so completely contracted that the animals appear quite smooth. The genus Notarchus, with a pallial cavity all around the visceral sac and with a quite narrow foot, differs widely from the rest of the Brevicommissurata, in which the pallial cavity does not extend under the visceral sac and which have a broader foot. 842 Bulletin of Marine Science [22(4) Martin (1966: 72) distinguished only three subfamilies of the Aplysiidae: Aplysiinae, Dolabellinae, and Dolabriferinae, and alloted Notarchus, of which he gave beautiful photographs, to the Dolabriferinae. ACKNOWLEDGMENTS Prof. Dr. Frederick M. Bayer, Rosenstiel School of Marine and Atmo- spheric Sciences of the University of Miami, entrusted me with hundreds of specimens of opisthobranchs collected by himself and other members of the school and on cruises of the R/V JOHN ELLIOTT PILLSBURY and R/V GERDA. Drs. Marc Kempf, Recife, and Pierre Montouchet, Campinas, gave me their opisthobranchs from the Brazilian coast. From Lt. Col. Corinne E. Edwards, Miss Kaoru Hiroki, Mrs. Helen Hughes, Dr. John Lewis, Dr. Donald Moore, Messrs. Edgardo Ortiz Corp's, Bruce Ott, Robert Austin Smith, Robert C. Work and Warren Zeiller I received specimens. To all of them I am grateful for the interesting material and much help. Mr. Robert Austin Smith kindly revised my manuscript and the bibliography. Dr. Eurico de Oliveira classified the algal contents of the stomachs of three aplysiids dredged in 348-357 m depth. The "FundaC;ao de Amparo a Pesquisa do Estado de Sao Paulo" (FAPESP) granted me the trip to Recife, Curac;ao, Puerto Rico, and Miami for collecting and comparing material (Proc. 71/961), for which I express my sincere thanks. Dr. and Mrs. Ingvar Kristensen invited me to their house during my stay at Curac;ao, and I am grateful for the very pleasant time at their home. LIST OF SPECIES Order ANASPIDEA Suborder LONGICOMMISSURATA Family Aplysiidae Subfamily Aplysiinae Genus Aplysia Linne, 1767 1. Aplysia (Pruvotaplysia) parvula March, 1863 2. Aplysia (Varria) brasiliana Rang, 1828 3. Aplysia (Varria) cervina Dall & Stimpson, 1901 4. Aplysia (Varria) dactylomela Rang, 1828 5. Aplysia (Varria) donca Marcus, 1960 6. A plysia (V arria) morio Verrill, 1901 7. Aplysia (Aplysia) juliana Quoy & Gaimard, 1832 Aplysia (Varria) willcoxi Heilprin, 1886-of this species, I had no specimens. Suborder BREVICOMMISSURATA Family Notarchidae Subfamily Notarchinae Genus N otarch us Cuvier, 1817 8. Notarchus punctatus armatus Baba, 1938 1972] Marcus: Anaspidea of the Western Atlantic 843 Subfamily Dolabriferinae Genus Stylocheilus Gould, 1852 9. Stylocheilus longicauda Quoy & Gaimard, 1824 10. Stylocheilus citrin us (Rang, 1828) Genus Bursatella Blainville, 1817 11. Bursatella leachii pleii (Rang, 1828) Genus Dolabrifera Gray, 1847 12. Dolabrifera dolabrifera (Cuvier, 1817) Genus Petalifera Gray, 1847 13. Petalifera petalifera bonairei Engel, 1936 14. Petalifera ramosa Baba, 1959 Genus Phyllaplysia P. Fischer, 1872 15. Phyllaplysia engeli Marcus, 1955 Of the well-described worldwide species, I do not give new descriptions nor all the localities. KEY TO THE ANASPIDEA Figs. 1-8 1. Pleurovisceral nerve cords long, visceral ganglia far behind pedal ganglia . Longicommissurata, Aplysiidae, 2 Pleurovisceral nerve cords quite short, visceral ganglia over pedal ganglia . Brevicommissurata, Notarchidae, 3 2. Animals with dorsocaudal, oblique disc _ _______________________________________Dolabellinae (Dolabella, Indopacific, Fig. I) No dorsocaudal disc Aplysiinae, Aplysia (Syphonota, Indopacific) 3. Dorsal slit in anterior half of body, sole quite narrow, pallial cavity surrounds entire visceral mass Notarchinae, Notarchus Dorsal slit in middle or posterior half, pallial cavity dorsal and lateral to visceral mass . Dolabriferinae, 4 4. Foot narrower than greatest diameter of body, no shell .___ 5 Underside set off by a ridge along widest part of body 7 5. A row of small papillae around the broad sole . B ursatella No such papillae, long and slender tail Stylocheilus, 6 6. Color yellow to orange with white spots . S. citrinus Color greyish with dark stripes and eyespots S. longicauda 7. Normal, primary shell present . 8 Shell absent or a small, secondary disc . Phyllaplysia 8. Shell foramen closed _.. Dolabrifera Shell foramen wide . Petalifera, 9 9. Only two denticles on the outer side of the lateral teeth of the radula ______. .. ._. Petalifer a pe tali fera bo nair ei More than two dentic1es Petalifera ramosa 844 Bulletin of Marine Science [22(4 ) KEY TO THE SUBGENERA OF Aplysia 1. Parapodia joined high up posteriorly 2 Parapodia joined low down posteriorly _ 3 2. Sole narrow, no sucker Aplysia (Pruvotaplysia) Engel, 1936) 1. A. parvula Sole broad, its hind end producing a temporary sucker _ Aplysia (Aplysia) Linne, 1767 7. A. juliana 3. Long and slender tail, shell with platelike, flattened apex __ ________________________________________Aplysia (Neaplysia) Cooper, 1863 (Pacific) No long tail, apex a reduced spiral A plysia (V arria) Eales, 1960 KEY TO WESTERN AND MIDDLE ATLANTIC SPECIES OF Aplysia (Varria) 1. Black rings on outer side of para podia 4. dactylomela No black rings 2 2. Mantle aperture closed 6. morio Man tie aperture open 3 3. Mantle aperture a flat hole 4 Mantle aperture tubular, on tip of papilla 5 4. Opaline gland multiporous 5. donca Opaline gland uniporous . fasciata Poiret, 1789* 5. Scattered, round, dark spots, diameter 2-3 mm, on parapodia, caecum curved 3. cervina No round spots, caecum straight .______6 6. Only tip of caecum visible on surface of digestive gland . _ ________________________________________________________________________wilcoxi, Heilprin, 1886t Caecum straight on surface of digestive gland 7 7. Para podia with smooth edges 2. brasiliana Parapodia with fimbriated edges . winneba Eales, 1960* 1. Aplysia (Pruvotaplysia) parvula Morch, 1863 References.-Eales, 1960a: 287, figs. 10, 11 (synonyms and bibliography); Marcus, 1967a: 154; 1970a: 26; Barash & Danin, 1971: 167. Material.-Brazil: Bahia, Olivem;a, 17. I. 1969; Bahia, Salvador, Ondina, 18. X. 1968; Espirito Santo, Guarapari, 6. XII. 1968, a total of 25 speci- mens in the lower tidal zone, Pierre Montouchet leg. Further Distribution.-From 400N to 40·S in all warm and temperate seas. Remark.- The Israeli locality of Barash & Danin (1971) seems to be the first indication for the Mediterranean. • For the two Middle Atlantic species of Aplysia. see Eales (1960a: 315, 352). t As I had no specimens of Aplysia willcoxi, it is not discussed in this paper; for a description, see Eales (1960; 350). 1972] Marcus: Anaspidea of the Western Atlantic 845 1 8 FIGURES 1-8. Aspect of living animals: 1, Dolabella; 2, Aplysia; 3, Notarchus punctatus armatus (from Baba, 1938); 4, Stylocheilus longicauda; 5, Bursatella leachii pleii; 6, Dolabrifera dolabrifera; 7, Petalifera ramasa (from Baba, 1959); 8, Phyllaplysia engeli. (Shell stippled.) 2. Aplysia (Varria) brasiliana Rang, 1828 Fig. 9 References.-Marcus, 1957b: 4, figs. 1-23; 1959: 8, figs. 11, 15-18; Eales, 1960a: 297, figs. 14, 15 (bibliography). Material.-PILLSBURY Sta. 580: 21°0S'N, 86°23'W, 348-357 m, one speci- men, 23 mm long, preserved, 22. V. 1967, together with A. donca and A. moria. 846 Bulletin of Marine Science [22(4 ) Further Distribution.-From
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