On Some Cephalaspidea (Gastropoda: Opisthobranchia) from the Western and Middle Atlantic Warm Waters
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ON SOME CEPHALASPIDEA (GASTROPODA: OPISTHOBRANCHIA) FROM THE WESTERN AND MIDDLE ATLANTIC WARM WATERS EVELINE D. B.-R. MARCUS Caixa Postal 6994, 01000 Siio Paulo, Brazil ABSTRACT Seventy-two species of the Acteonidae and of the genera Scaphander and Philine, known from the western Atlantic from Cape Cod to the Falklands, east to the Azores and Canaries, are listed; the 25 species represented in the present material are described and figured. Only for three, Aeteon eandem', Seaphander nobilis, and S. (Sabatina) bathymophilus, are the soft parts described for the first time. Several species are redescribed. Of the known species, nine have not been figured till now; 48 are descriptions of empty shells, so that their generic position remains uncertain, and the soft parts are known only for 26 species. Acteon cumingii A. Adams, 1854, is the type-species of Mysoufja, gen. nov. Aeteon eandens, Seaphander nobilis, and Philine pr. jalklandiea are new for Brazil. INTRODUCTION The present paper is the third part of my purpose to catalogue the western warm water Opisthobranchia. The first dealt with the Anaspidea (Marcus, 1972c), the second with the genus Bosellia (Marcus, 1973). A great number of specimens was entrusted to me by Prof. Frederick M. Bayer, Rosenstiel School of Marine and Atmospheric Science, University of Miami; they were collected during the oceanographic cruises of the R/V JOHN ELLIOTT PILLSBURY and the R/V GERDA during the years 1964-70. Furthermore, I received one sample from Chandeleur Sound, Gulf of Mexico, from Dr. Renate Schlenz True (New Orleans); several samples from Drs. Plinio Moreira Soares and Roberto Tommasi (Sao Paulo); and one sample from Dr. Eliezer de Carvalho Rios (Rio Grande do Sui). Two collections of dry shells I got from Dr. J. Meyer (Amsterdam), and Frere Fridericus (Aruba, N.A.). One species (Philine aperta guineensis) not found in the area (marked -1+) is an addendum of the PILLSBURY Deep-Sea Expedition to the Gulf of Guinea, 1964-65. Of the 61 species with figured shells from the mentioned area, the soft parts have been described for only 25 species, four of which are in the present paper. The generic position of empty shells is often difficult to decide. Pilsbry (1893-95: 18],242) and Zilch (1959-60: 6) expressed the need for anatomical studies. There are several examples for erroneous allocations. Manuscript accepted July 1973. 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 301 For a reliable classification the operculum, jaw elements, radula, gizzard plates, hermaphroditism or separate sexes, and the male organ are desirable. The comparison of shells with previous descriptions often meets with subjective methods of counting whorls and sculptural striae. In the former the protoconch is often not considered, due to the fact that it is frequently worn. The striae are of unequal breadth, and there may be new grooves in different degrees of development; hence, generally large shells have more numerous striae than smaller ones of the same species. Dall evidently counted only the main striae and left aside beginning and short rows around the umbilical region. The ratio of width to length or length to width as percentage is also a doubtful criterion. The small and delicate shells are not easy to measure, and generally the relative breadth diminishes with the growth in length. ACKNOWLEDGMENTS My sincere thanks are due to those who furnished me with opisthobranch specimens: Dr. Renate Schlenz True, New Orleans; Dr. Meyer, Amster- dam; Frere Fridericus, Aruba; Drs. Plinio Moreira Soares and Roberto Tommasi, Sao Paulo; Dr. Eliezer de Carvalho Rios, Rio Grande; and last, not least, Prof. Frederick M. Bayer of Miami, who gave not only specimens, but encouragement and friendship. Furthermore, I thank Miss Roberta J. Imrie, Capetown; Dr. P. N. Kilburn, Pietermaritzburg; Prof. Tadashige Habe, Tokyo; and Dr. Robert Robertson, Philadelphia, for advice and bibliographic help, and especially Dr. Tor G. Kading, Stockholm, for lend- ing me the valuable old volume of G. O. Sars, 1878. Mr. Eduardo do Patrocinio Fernandes made the fine photographs of the shells of Scaphander. Once again Robert Austin Smith and Mrs. Gisela Marmol had the trouble to revise my manuscript for language and correct citations, and I am very grateful to both. LIST OF SPECIES (Those not represented in the present collection are marked with +, and those not in the treated area are marked with tt) Phylum MOLLUSCA Class GASTROPODA Subclass EUTHYNEURA Order CEPHALASPIDEA Superfamily Acteonacea Family Acteonidae A. Acteonidae with established generic position. tt Acteon tornatilis (Linne, 1758) 1. Acteon candens Rehder, 1939 (Figs. 10-15) + 2. Acteon pelecais d. B.-R. Marcus, 1972 (Fig. 1) 302 Bulletin of Marine Science [24(2) + 3. Rictaxis punctostriatus (C. B. Adams, 1840) (Fig. 2) 4. MysoufJa, gen. nov., cumingii (c. B. Adams, i854) (Figs. 6, 8) B. Acteonidae with unknown radula in the present collections. 5. "Acteon" exiguus March, 1875 (Fig. 9) 6. "Acteon" splendidulus March, 1875 (Fig. 7) 7. "Acteon" melampoides DaH, 1881 (Figs. 16, 17) 8. "Acteon" perforatus DaH, 1881 (Figs. 18-20) 9. "Ac/eon" incisus Dall, 1881 (Fig. 23) 10. "Acteon" turri/us Watson, 1883 (Figs. 21, 22) 11. "Acteon" vagabundus Mabille, 1885 (Figs. 24, 25) 12. "Acteon" spec. 904 (Figs. 26, 27) 13. "Acteon" spec. 1261 (Figs. 28,29) 14. "Acteon" (Lissacteon) exilis (Jeffreys, 1870) (Fig. 30) C. Acteonidae with unknown radula, not in the present collections. + 15. "Ac/eon" senegalensis Petit, 1852 + 16. "Acteon" danaida DaH, 1881 + 17. "Ac/eon" amabilis Watson, 1883 + 18. "Acteon" hebes Verrill, 1885 + 19. "Acteon" delicatus Dall, 1889 + 20. "Acteon" monterosatoi Dautzenberg, 1889 + 21. "Acteon" semisculptus E. A. Smith, 1890 + 22. "Acteon" grimaldii Dautzenberg & Fischer, 1896 + 23. "Acteon" azoricus Locard, 1897 + 24. "Acteon" torrei Aguayo & Rehder, 1935 + 25. "Acteon" finlayi McGinty, 1955 + 26. "Acteon" (Lissacteon) nitidus (Verrill, 1882) + 27. Leucotina minuta E. A. Smith, 1890 + 28. Neactaeonina chariis (Watson, 1883) + 29. Bullina exquisita McGinty, 1955 D. List of unfigured Acteonidae with unknown radula (see p. 319). Superfamily Philinacea Family Scaphandridae A. Species treated. + 30. Scaphander lignarius (Linne, 1758) (Fig. 31) 31. Scaphander punctostriatus (Mighels, 1841) (Fig. 33) 32. Scaphander watsoni Dall, 1881 (Figs. 35-41) 33. ScaphandermundusWatson, 1883 (Figs. 81, 84) 34. Scaphander? gracilis Watson, 1883 (Fig. 32) 35. Scaphander nobilis Verrill, 1884 (Figs. 42-50, 81, 83) 36. Scaphander clavus Dall, 1889 (Figs. 34, 81, 85) 37. Scaphander stigmaticus Dall, 1927 (Figs. 51-56) 38. Scaphander darius Marcus, 1967 (Figs. 57-66, 81, 86) 39. Scaphander (Sabatina) bathymophilus (DaIl, 1881) (Figs. 67-82) Family Philinidae 40. Philine finmarchica M. Sars, 1858 (Figs. 88-92) 41. Philine infundibulum DaIl, 1889 (Figs. 93-97) 42. Philine pI. falklandica Powell, 1951 (Figs. 98-101) 43. Philine alba Mattox, 1958 (Figs. 102, 103) 44. Philine mera Marcus, 1969 (Figs. 105-111) tt Philine aperta guineensis Marcus, 1969 (Fig. 104) 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 303 B. Speciesof Philine not in the present collections. + 45. Philine aperta aperta (Linne, 1767) + 46. Philine scabra (0. F. MUller, 1776) + 47. Philine catena (Montagu, ]803 ) + 48. Philine lima (Brown, 1825) + 49. Philine quadrata (S. Wood, 1839) + 50. Philine sinllata (Stimpson, ]850) + 51. Philine trachyostraca Watson, 1897 + 52. Philine gibba Strebel, ]908 + 53. Philine thurmanni thurmanni Marcus, 1969 + 54. Philine amabilis Verrill, 1880 C. List of species of Philine known only for their shells (see p. 364). Family Acteonidae Summarizing my previous paper (1972a): the generic position of "Acteon" punctostriatus from North America was changed to Rictaxis Dall, due to its radula, while the South American "punctostriatus" received the new name Aeteon peleeais (1972b: 30 I). The positions of "Aeteon" danaida and vagabundus (1970: 924, figs. 3, 4, 6, 7) remain uncertain as long as their soft parts are not known. In his description of Aeteon punetostriatus, Dall (1889: 40) mentioned the variation of "color, height of the spire, elevation of the nucleus, and extent of shell covered by the punctate lines. There may be one or several subsuturallines, the middle of the whorl is generally smooth .... " This description suggests that Da\l's material included several species besides punetostriatus (No.3), perhaps Aeteon eandens (No.2), "Aeteon" exiguus (No.5), "Aeteon" torrei (No. 24), or others, even completely striated ones. When I published my paper (1 972a), I had not seen several papers concerning the family, e.g., Taki's paper on Japonaetaeon (1956) and that of Rudman (1971) on Maxaeteon. Moreover, I had followed Strand's error in giving the name Tomlinula as a second name to Tomlin's Alexandria ( 1926), to which mistake Prof. Habe kindly called my attention. I had not found the paper of Robertson & Habe (1965) on Alexania, for a copy of which I am grateful to Miss Roberta Imrie (Capetown) and Dr. R. N. Kilburn (Pietermaritzburg). The genus Alexania was removed to the Proso- branchia Ptenoglossa Epitoniidae. The species of Alexania which Robert- son studied from India (1965: 141) is a true prosobranch with "small to dwarf males." As in the case of Alexania, sometimes even the knowledge of the radula is not sufficient. Thiele (1912: 220) compared the radula of Neaetaeonina eingulata (Strebel, 1908) with that of the ptenoglossan Janthina. Also the absence of a free pallial caecum in that species (Odhner, 1926: 4) is not acteonid. Strebel's prosobranch Ohlinia (1905: 597, pI. 22, fig. 32), family Cerithiopsidae, was synonymized with the opisthobranch Toledonia Dall, 1902 (Thiele, 1931: 382). 304 Bulletin of Marine Science [24(2) The uncertain generic position of species known only by their shells is also evident in Lemche's synonymic lists (1948: 70-99). While Fretter & Graham (1954: 572) could not determine the food of Acteon tornatiUs, Hurst (1965: 326) showed that it feeds upon worms. Rudman (] 972: 317) observed Pupa eating sand-dwelling polychaetes, and Maxacteon cratericulus (p.