ON SOME (: 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 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 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, , 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 Class GASTROPODA Subclass EUTHYNEURA Order CEPHALASPIDEA Superfamily Acteonacea Family Acteonidae A. Acteonidae with established generic position. tt (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 , elevation of the nucleus, and extent of shell covered by the punctate lines. . . . There may be one or several subsuturallines, the middle of the 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. 32]) eats Cirratulidae. I found parts of poly- chaetes in Mysauffa cumingii and Rictaxis punctocaelatus and punctostriatus (1972a: ] 74, ]78). So Rudman's statement that the Acteonidae are spe- cialized vermivores (1972: 311) is once more supported. My key (1972a: 169) must be increased to cover the genera lapo- nactaeon and Maxacteon.

KEY TO THE GENERA OF ACTEONIDAE

1. Radula 00 .0. 00 equal teeth .._. .._A ctean Montfort, ] 810 (Fig. ]) 1. Radula with less numerous teeth __. . .. 2 2. Radula with 5.0.5 teeth 3 2. Radula with more than 5.0.5 teeth 4 3. Teeth of almost equal size and shape Pupa Roeding, 1798 3. One tooth much larger than the rest Rictaxis Dall, 187] (Fig. 2) 4. Radula with 6.0.6 teeth lapanactaeon Taki, 1956 (Fig. 5)l 4. More than 6.0.6 teeth . ..__5 5. Teeth with several cusps of equal length Mysouffa, gen nov. (Fig. 6) 5. Teeth with long main cusp, with or without accessory denticles 6 6. Teeth with main cusp and accessory denticles ______Maxacteon Rudman, ]971 (Fig. 4) 6. Teeth with long cusp without denticles . ______Neactaeonina Thiele, 19]2 (Fig. 3)

A. ACTEONIDS WITH ESTABLISHED GENERIC POSITION Genus Acteon Montfort, 1810 + Acteon tornatilis (Linne, 1758) Rejerences.-G. O. Sars, 1878: 260, pl. 17, fig. 11, pl. XI, fig. 1, a-c, pl. XVIII, fig. 57.-Pilsbry, 1893-95: 152, pl. ]9, figs. 7-11, 15.-Pelseneer, 1894: 5, figs. 1-15.-Lemche, 1948: 70.-Fretter & Graham, 1954: 565- 585, figs. 1-9.-Johansson, 1954: 223-232, pl. I.-Hurst, 1965: 326 ff., figs. 25-27.-Marcus, 1972a: 167. Distribution.-From Norway to Morocco and the Mediterranean. Remark.-Though it is improbable that the species will be found in the western Atlantic, I have included it, because it is the most studied species of the genus.

1Rudman suggests that Taki's interpretation of the anatomy of Japonactaeoll might be erroneous. 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 305

FIGURES1-9.-1-6, Radulae of Acteonidae: 1, Acteon pelecais d. B.-R. Marcus; 2, Rictaxis punctostriatus (C. B. Adams); 3, Neactaeonina cingulata (Strebel) (from Thiele, 1912); 4, Maxacteon hancocki Rudman (from Rudman, 1971); . 5, Japonactaeon nipponensis (Yamakawa) (from Taki, 1956); 6, MysoufJa, gen. nov., cumingii (A. Adams).-7-9, Features of acteonid shells: 7, "Acteon" splendidllius Morch, 1875; 8, MysoufJa cumingii, periostracum (the deep black lines were bright orange); 9, "Acteon" exiguus March.

1. Acteon candens Rehder, 1939 Figs. 10-15 Reterences.-Rehder, 1939: 21, pI. 6, fig. 7.-Abbott, 1955: 275.-Mar- cus, 1972a: 182. Material.-Gulf of Mexico, Louisiana, Chandeleur Sound, four specimens, thanks to the kindness of Dr. Renate Schlenz True.-Brazil, off Espirito Santo, 19° 23' S, 39° 40' W, 11 m, Institute of Oceanography, University of Sao Paulo, one specimen.

Further Distribution.-From Cape Hatteras to Dry Tortugas? Brazil, off Espirito Santo. 306 Bulletin of Marine Science [24(2)

FIGURES 10-15. A cteon candens Rehder: 10, A, shell from Gulf of Mexico; 10, B, shell from Brazil; 11, tips of two jaw platelets; 12, preserved taken out of shell; 13, operculum; 14, penis; 15, radular tooth. (f, foot; 0, operculum; p, penis.)

Description.- The shells of the present (Fig. 10) are 4-5 mm long, hence smaller than in the original description (Rehder, 1939: 7.5 mm). Their ratio of width to length is 50-56.2 per cent and lies between the extremes of the previous shells with 43.5-61.2 per cent. The length of the amounts to 52.9-76.4 per cent. The rather solid shells are whitish transparent. They have 4-4.5 whorls. Their sculpture consists of 9-12 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 307 spiral striae and brown, punctate grooves between them in the anterior part of the body whorl, leaving the posterior half smooth. The head shield (Fig. 12) has two anterior lobes, the labial tentacles, and two posterior lobes, the cephalic tentacles, folded forward as in Fretter & Graham's figure 2 (1954: 567). The anterior border of the foot is slightly notched and has two lateral extensions similar to those of the head shield mentioned by Rudman for Maxacteon (1971: 208, fig. 1). The thin operculum (Fig. 13) is 2 mm long, 0.96 mm wide, in the 5- mm animal. The pigment in the roof of the mantle cavity forms a ramified pattern (Fig. 12) in contrast to the pigment-free mantle border of A. pelecais (No.2). The pallial caecum is reduced to a short cone (Fig. 12); but it may be contracted due to the preservation in alcohol (Pelseneer, 1894: 5-6). The jaw platelets and the radular teeth (Figs. 11, 15) are typical of Acteon (Gabe & Prenant, 1953: fig. 3; Marcus, 1958: 33, figs. 5-10; Marcus, 1972a: 170,173, figs. 6-9,12-13). The narrow mouth-tube widens where it passes between the small jaw plates, composed of spiny rodlets. The radula lies in a long and narrow buccal mass which receives the short, club-shaped salivary glands at its hind end. The slender oesoph- agus is short and leads into the wide stomach. The prostatic gland has a high, smooth epithelium. The penial papilla is slender and smooth with a widened tip and a strongly coiled efferent duct inside (Fig. 14). One specimen from Espirito Santo, Brazil, corresponds, with its broad operculum, pigmented mantle roof, short pallial caecum, and long male organ with a widened tip, to the organization of Acteon candens. Its shell (Fig. 10, B) is different from the ones from the Gulf of Mexico. It has a deep suture and a pronounced shoulder, and its apex is more slender. Remarks.-The radula of the present material is decisive for its generic position. It differs from the other known semistriate West Atlantic species, Acteon pelecais Marcus (No.2), by the pigment pattern of the mantle roof, the short pallial caecum, and the shape of the penial papilla. The species is new for Brazil.

+ 2. Acteon pelecais d. B.-R. Marcus, 1972 References.-Marcus, 1958: 32, figs. 1-10 (A. punctostriatus); 1970: 924, fig. 5 (? A. punctostriatus); 1972a: 170, figs. 1, 6-10 (A. pelecais); 1972b: 301.-non Acteon punctostriatus (C. B. Adams, 1840). Distribution.-Brazil, Sao Paulo, in shallow water. Remarks.-A. pelecais belongs to the group with smooth posterior part of the body whorl. 308 Bulletin of Marine Science [24(2) The subsutural line mentioned in the description (Marcus, 1958: 32) was seen in the periostracum of one shell, not in the shell proper of the others whose periostracum was lost. The shape of the shell of "Acteon" exiguus (No.5) with a single line below the suture is sufficiently different to maintain the species separate, even if the line in pelecais should be more frequent. Genus Rictaxis Dall, 1871 + 3. Rictaxis punctostriatus (c. B. Adams, 1840) Fig. 2 Reterences.-Pilsbry, 1893-95: 157, pI. 18, figs. 98, 99, pI. 19, figs. 22, 23 (Actaeon punctostriatus).-Marcus, 1972a: 178, figs. 2, 30-37 (Rictaxis punctostriatus).-non Acteon punctostriatus Marcus, 1958: 32, figs. 1-10. Distribution.-From Buzzards Bay, Massachusetts, to Cuba and San Do- mingo. The radula is known only from Chesapeake Bay and Florida, in shallow water. Remarks.-Due to the radular formula 5.0.5, "Acteon" punctostriatus is allotted to Rictaxis Dall, 1871 (Marcus, 1972a). The limits of its dis- tribution must be determined by the study of the radulae of future catches, as the shells are only minutely different from those of Acteon pelecais (see 1972a, figs. 1 and 2) with the formula 00 .0. 00 from Brazil.

Genus MysoufIa, gen. nov. Acteonidae with many, almost uniform, strong teeth per half-row, with a restricted number of pointed denticles. Jaw elements with many spines on one border. Type-species.-Mysoufja cumingii (A. Adams, 1854) Marcus, 1972a: 173. The generic name Tomlinula, which I had applied to Acteon cumingii (1972a: 173), is a synonym of Alexania Strand, 1928. This genus was removed to the prosobranch Ptenoglossa Epitoniidae, so the true acteonid cumingii must receive a new generic name. I call it Mysoufja. Mysoufj was Alexandre Dumas's cat.

4. Mysoufja cumingii (A. Adams, 1854) Figs. 6, 8 References.-Pilsbry, 1893-95: 162, pI. 19, figs. 16, 17 (Acteoncumingii). -Marcus, 1972a: 173, figs. 4, 15-18 (Tomlinula cumingii). Material.-Brazil, off Ubatuba, 23° 05'-39' S, 45° 06'-40' W, 22-28 m, 6. V. and 8. V. 1970, Plinio Moreira Soares leg., one empty shell and four animals, 4, 3, 2.5 and 2 mm long. ]974] Marcus: Cephalaspidea from Atlantic Warm Waters 309 Further Distribution.-From Florida, 15 m, to Brazil, Sao Paulo. Remarks.-Rios (1970: 130, pI. 49) listed and figured "Acteon" finlayi (No. 25) from the Brazilian coast. His photograph might just as well be the present species, and the soft parts are not known. Figure 8 illustrates the periostracum with the borders of the spiral grooves, which in the stretches drawn deep black were rusty orange, and in the parts between, colorless. The radula of the 4-mm specimen has 16 teeth per half-row, that of the 2-mm animal has 13 teeth per half-row.

B. ACTEONIDAE WITH UNKNOWN RADULA IN THE PRESENT COLLECTION 5. "Acteon" exiguus March, ] 875 References.-Pilsbry, 1893-95: 161.-Thiele, 1926: 95, fig. 104. Material.-Suriname, coastal waters, ] 968, J. Meyer leg., two empty shells, 4.5 and 5.0 mm high.-PILLSBURY Sta. P-758: ] 1042.2'-]] 044' N, 690 40' W, 14-18 m, 27. VII. 1968, one 4-mm shell. Further Distribution.-Antilles (March); West Indies (Thiele).

Description.- The transparent white shells measure from 2.2 X 1.2 to 6.2 X 3.1 mm (Thiele). They are moderately thick and regular in shape; their greatest width corresponds to 50 per cent, the height of the aperture to 60 per cent, of the total height. The largest shells have five whorls. The spire is rather elevated. The sculpture consists of spiral grooves in the anterior half of the body whorl, and one more single fine groove just below the very distinct suture. The spiral grooves at the level of the insertion of the outer are far apart; farther in front they lie close together. The aperture is rounded in front. The columellar fold is distinct. Discussion.-The unfigured description of March, copied by Pilsbry, com- prises five lines. However, the groove which margins the suture is a distinc- tive character sufficient to recognize the species, of which March evidently had two specimens, different in their ratios, the one 6.5 X 3 mm, the other 6.0 X 3.1 mm. Thiele (1926) gave an original figure of a shell from the West Indies. Remarks.-The allocation of the present material to March's species is beyond doubt; however, its generic position remains uncertain, as long as neither radula nor operculum are found. 6. "Acteon" splendidulus March, 1875 Fig. 7 Reference.-Pilsbry, 1893-95: 161. 310 Bulletin of Marine Science [24(2) Material.-Suriname, coastal waters, 1966, J. Meyer leg., six empty shells. -Aruba, from sand pumped up in the harbour, Frere Fridericus leg., seven empty shells. Further Distribution.-St. Thomas, one specimen. Description.-The largest shells from Aruba measure 3.1 and 3.15 mm in height, 1.6 mm in greatest width, and have five whorls. The greatest width lies in the middle. The shells from Suriname are smaller, up to 2.0 mm high. The shells are rather solid, transparent white. The ratio of width to length is 50.1-64.1 per cent. The aperture occupies 57-63 per cent of the length; it is rounded in front. The protoconch is smooth and quite trans- parent. The suture is well marked by a round shoulder. The single colu- mellar fold is sharp. The upper part of the whorls is smooth; in the anterior part there are spiral grooves consisting of rows of longish pits. Below the suture there are about 20 rows on the body whorl; their interspaces diminish forward, and there are two or more above the insertion of the outer lip, and these two or three rows appear above the suture in the third and fourth whorl (counted from the protoconch) (Fig. 7). Discussion.- There are only six western Atlantic species of acteonids with a smooth upper part of the body whorl, of which now only "Acteon" semi- cingulatus Dail (1927: 19) is unfigured. The present species corresponds to March's description, cited from Pilsbry (1893-95: 161-162), that the spire bears two punctate lines. As March's only specimen came from St. Thomas, there is no doubt that my material is conspecific with his. As the soft parts and the radula are not known, the generic place of the present species cannot be determined.

7. "Acteon" melampoides Dall, 1881 Figs. 16, 17 References.-Dall, 1889: 41, pI. 17, fig. 2; 1903: 84, pI. 17, fig. 2, pI. 46, fig. 15.-Pilsbry, 1893-95: 158, pI. 20, fig. 33.

Material.-P-904: 13045.5' N, 61005.7' W, 457 m, 9. VII. 1969, one specimen. Further Distribution.-Off. Cuba, 567 m; Gulf of Mexico, 1472 m. From Virginia to Barbados, 310-2574 m.

Description.-The single delicate and broken empty shell measures 9.6 X 7 mm, the ratio of width to length is 72.9 per cent. The aperture is 7.3 mm high, the spire 1.5 mm. The protoconch is worn. There are six whorls. 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 311

oocco oecOCl ~~ooc ~oooo OOOoC> 19 (Jfl{JlJ lJ{]fJ{j (J{j{j{j 000 20 0000 ()(j(JO 1700o( ':>ODD

00000 00000 ====c::> 000000 22

FIGURES16-23. Features of acteonid shells: 16, "Acteon" melampoides DaIl, shell; 17, same, sculpture of shell; 18, "Acteon" perforatus Dall, shell; 19, same, sculpture near posterior end; 20, same, sculpture near anterior end; 21, "Acteon" turritus Watson, shell; 22, same, sculpture; 23, "Acteon" incisus Dall, shell.

The body whorl bears 32 striae, the second whorl, five. The interspaces are as wide as the roundish pits in the posterior part of the body whorl, narrower towards the anterior end (Fig. 17). There is no callus on the body whorl. The reflexed has a distinct fold and covers a narrow umbilical chink (Fig. 16).

Remarks.-Dall's original single specimen measured 6 X 4 mm. Dall (1889: 41) had little doubt of the identity of Verrill's Acteon hebes (1885; No. 18) from off North Carolina in 4707 m, which had lost its spire and cannot be allotted to anyone of the short-spired species. He synonymized it with A. melampoides. However, in 1927 (p. 19) Dall classified two young specimens of A. melampoides from off Georgia as A. hebes Verrill. Dall would call "Acteon" torrei Aguayo & Rehder (1935: 268, pI. 24, 312 Bulletin of Marine Science [24(2) fig. 8) (No. 24) probably synonymous (1889: 40) with melampoides, because it comes from the same locality, Habana, 16 m. However, the biggest shell of torrei is 8.5 mm high and has only 4.5 whorls and a very distinct nucleus. 8. "Acteon" perforatus Dall, 1881 Figs. 18-20 References.-Dall, 1889: 42, pI. 18, fig. 3; 1903: 84-85, pI. 18, fig. 3.- Pilsbry, 1893-95: 159, pI. 20, fig. 36. Material.-P-607: 18° 30' N, 87° 37' W, 731-860 m, 17.-18. III. 1967. -P-904: 13° 45.5' N, 61 ° 05.7' W, 457 m, 9. VII. 1969. Further Distribution.-Florida Straits; Cuba, 339 m.

Description.-The two present empty shells measure 14 X 9.5 and 13.5 X 9.5 mm, respectively. Their ratio of width to length is 67.8 and 70.4 per cent; the greatest width lies in the middle. The aperture is 8 mm high, and the spire 4.0 mm. They have 6 to 6% whorls. The smaller shell is heavier than the biggest one. There are 34 and 33 spiral striae, a little narrower than the large, distinctly separate pits (Figs. 19, 20). The shape of the latter varies from roundish posteriorly to high and narrow near the anterior end, more pronounced in the heavier shell. The outer lip is broken. The inner lip has a light callus. The columella has a thickening in its middle and is reflexed over a deeply perforated umbilical chink (Fig. 18). The protoconch is worn. The suture is set off by the shoulder of the following whorl.

Remark.-Dall's specimen was 7.75 X 4.62 mm. Its columellar fold could hardly be seen.

9. "Acteon" incisus Dall, 1881 Fig. 23 References.-Dall, 1889: 42, pI. 17, figs. 1, 1b; 1927: 19.-Pilsbry, 1893- 95: 160, pI. 20, figs. 31, 34.-Dautzenberg & Fischer, 1897: 142. Material.-P-904: 13° 45.5' N, 61° 05.7' W, 366 m, 9. VII. 1969, two empty shells.-P-1261: 07° 13' N, 77° 50' W, 600-825 m, 15. VII. 1970, one shell. Further Distribution.- Yucatan Strait; Cuba; Florida, Fernandina, 294- 1360 m.

Description.-The shells from P-904 measure 9.5 X 6 mm and 6.5 x 4 mm, respectively, and that from P-1261 is 10 X 6 mm (Fig. 23); thus, the ratio is 60-63.2 per cent. They have five to six whorls. The aperture of 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 313 the biggest is 6 mm high, its spire, 2 mm. The slightly oblique columella has a narrow reflexed border and a very slight fold (Dall, 1889: 42). The 26 striae are limited by scalloped borders against the grooves, which are almost as wide. Remarks.-The striation of the present shells does not agree with Dall's description of 10-11 somewhat zigzag, nonpunctate grooves and other spiral, microscopically fine striae, about 70 in the width of a millimeter. However, DaB's figure 1 shows about 20 striae, which result in an aspect comparable to the sculpture of the present material.

10. "Acteon" turritus Watson, 1883 Figs. 21, 22 References.-Watson, 1886: 628, pI. 47, fig. I.-Dall, 1889: 40.-Pilsbry, 1893-95: 157, pI. 20, figs. 29, 30. Material.-RjV GERDASta. G-964: 23° 46' N, 81° 16' W, 760-773 m, 1. II. 1968, two shells.-P-1255: 17° 18' N, 78° 32' W, 620-820 m, 14. VII. 1970, one shell.-P-1261: 07° 13' N, 77° 50' W, 600-825 m, 15. VII. 1970, one shell. Further Distribution.-Off Culebra Island, 713 m.

Description.-The empty shells measure from 13 X 6.5 to 14.0 X 7.4 mm. In the latter, the aperture is 9 mm high, the spire, 3 mm. The ratio is 50- 52.9 per cent. There are 40-45 flat spiral striae on the body whorl with scalloped borders against the grooves. Their breadth is irregular. The bottom of the grooves is divided into pits a little longer than high (Fig. 22). The outer lip is broken in all four specimens. The inner lip has a callus along the body and the columella (Fig. 21). The columella has a fold covered by the callus near its upper end and is reflexed. There is no chink. Discussion.-Dall (1889: 40) compared turritus with punctostriatu.s: "though the figures are not very similar, the locality Culebra Island is suspicious." However, "Acteon" (now Rictaxis) punctostriatus (No.3) is semisculptured and much smaller, hence distinctly different from turritus, while delicatus Dall (1889: 41, pI. 15, fig. 5) (No. 24), measuring 10 X 5.6 mm, is to my opinion synonymous with turritus, as long as the soft parts are not proved to be different. The aspect of the punctations varies with the angle of the illumination.

11. "Acteon" vagabundus Mabille, 1885 Figs. 24, 25 Reference.s.-Mabille, 1885: 208 (Tornatella vagabunda) (ref. not seen). -Rochebrune & Mabille, 1891: 12, pI. 6, fig. 6 (Tornatella vagabunda). 314 Bulletin of Marine Science [24(2)

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FIGURES24-30. Features of acteonid shells: 24, "Acteon" vagabundus Mabille, shell; 25, same, sculpture of shell; 26, "Acteon" spec. 904, shell; 27, same, sculp- ture; 28, "Acteon" spec. 1261, shell; 29, same, sculpture; 30, "Acteon" (Lis- sacteon) exi/is (Jeffreys), shell, dorsal (left) and ventral (right) views.

-Pilsbry, 1893-95: 164, pi. 18, figs. 95, 96 (Actaeon vagabunda).- Marcus, 1970: 924, figs. 6, 7 (Acteon vagabundus). Material.-P-607: 18° 30' N, 87° 37' W, 750-786 m, 17.-18. III. 1967, one empty shell and a fragment. Further Distribution.-Cape Horn; Brazil, Alagoas, 09° 01' S, 34° 51' W, 370 m. Description.-The length of the present shell is 12 mm, its diameter, 6 mm, the ratio 50 per cent. The aperture occupies 67 per cent of the height, the spire is 2 mm high. There are about five whorls. The growth lines 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 315 are a few slight, opaque crests. There are about 25 flat, spiral striae, and the pitted grooves are much narrower than their interspaces. There are many fine secondary grooves between the primary ones (Fig. 25). The inner lip bears a glaze which goes onto the columella. The strong fold of the latter is set off by a deep, round spiral groove (Fig. 24). The outer lip is broken. Remarks.- The short spire, large aperture, sculpture, and the original description and figures of the shape and the columella agree so well with the present shell, that I dare to identify it with vagabundus, as long as the radulae from the localities so far apart are not known. As the Brazilian and the present material come from considerable depths, their specific identity is not quite improbable.

12. "Acteon" spec. 904 Figs. 26, 27 Material.-P-904: 13° 13' N, 77° 45.5' W, 366 m, 9. VII. 1969, one empty shell. Description.- The glossy white shell is 10 mm high and 6 mm in diameter; the aperture measures 6.2 mm, the spire 2.5 mm. There are six whorls. The outer lip is broken. The 23 flat striae are a little broader than the pitted grooves, and the whorls are distinctly narrowed towards the deep anterior suture. The callus is wide at the top of the columella, which bears a very weak fold. Remark.-I cannot identify the broad callus covering the place of a pos- sible chink with anything seen in other species, but from a single empty shell I do not want to create a new species. None of Dall's descriptions ( 1927: 19-21) of his seven unfigured species agrees with the present shell.

12. "Acteon" spec. 1261 Figs. 28-29 Material.-P-1261: 07° 13' N, 77° 50' W, 600-825 m, 15. VII. 1970, one empty shell.-P-754: see below.

Description.- The white, shining shell measures 8.3 X 4.8 mm, the aper- ture, 5 mm, and the spire, 2 mm. There are five whorls; the suture is marked by a shoulder. The apex is a little corroded. The approximately 40 irreg- ular flat striae are about twice as broad as the grooves. They have scalloped borders forming bridges between the pits, which correspond more or less to the slight growth lines. The outer lip is broken. The thin glaze on the body gets stronger on the columella and covers the insignificant umbilical chink completely. The columellar fold is near its top. 316 Bulletin of Marine Science [24(2) Remark.-"Acteon" melampoides Dall (No.7) is much more globose and has a shorter and broader spire. Of Dall's seven unfigured species (1927: 19-21), none can be applied to the present shell. Two rather worn specimens from P-754 (11° 36.9' N, 68° 42' W, 684- 1574 m, 26. VII. 1968) have proportions similar to those of spec. 1261, and their columellar fold is of the same shape. They measure 15.8 mm and ]4 mm X 7.8 mm, respectively.

14. "Acteon" (Lissactaeon) exilis (Jeffreys, ]870) Fig. 30 References.-Watson, ]886: 625 (Acteon exilis).-Dall, 1889: 39; 1903: 84-85 (Actaeon exilis).-Dautzenberg, 1889: 20, pI. 1, fig. 1 (A. exilis). -Pilsbry, 1893-95: 156, pI. ]9, figs. 5, 6 (Actaeon exilis).-non Pilsbry, 1893-95: 157, pI. 19, fig. 4 (A. nitidus).-Locard, 1897: 79, pI. 3, figs. 1-3 (A. exilis).-Dautzenberg & Fischer, 1896: 399 (A. [Lissactaeon] exilis); 1897: ]42.-Thiele, 1931: 378 (A. [Lissactaeon] exilis).- Lemche, 1948: 36,71 (synonyms of A. exilis).-Zilch, 1959-60: 8 (Creni- labium exilis).-Clarke, 1962: 39 (A. exilis). Material.-G-826: 25° 35'-25° 37'N, 80° 01'W, 269-271 m, 7. VII. ]967, one empty shell and a fragment. Further Distribution.-North Atlantic to Mediterranean; Azores, 160- 2679 m.

Description.-The present delicate white shell measures 2.6 X 1.2 mm. It has 3.5 whorls; the body whorl occupies 2 mm, the length of the aperture is 1.5 mm. The protoconch is big, the suture slightly channelled. The columellar fold is inconspicuous. The upper part of the body whorl and the second whorl bear irregular transparent growth lines. The anterior part of the body whorl has about six narrow spiral grooves. Discussion.-Zilch (1959-60) reestablished the synonymy of the Recent subgenus Lissactaeon Monterosato, 1890, with the fossil genus Crenilabium Cossmann, 1889, though Dautzenberg & Fischer (1896: 398) had stressed that Acteon exilis Jeffreys, ] 870, has no crenulated inner lip, and main- tained the genera separate. So did Thiele (1931: 378), who had described a further species from a 2.5 X 1 mm shell with 2.5 whorls from East Africa (1925: 225, pI. 30, fig. 13). Pilsbry (1893-95: 157) identified A. nitidus Verrill, ] 885 (No. 26) with A. exilis Jeffreys; his figures 5 and 6 are from Dautzenberg, 1889. The latter species has much weaker spiral and growth lines, and Dautzen- berg & Fischer (1896: 399) maintained both species separate. 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 317

C. ACTEONlDAE WITH UNKNOWN RADULA NOT IN THE PRESENT COLLECTION + 15. "Acteon" senegalensis (Petit, 1852) References.-Petit de la Sausseye, 1852: 262, pI. 8, fig. 3.-Pilsbry, 1893- 95: 152, pI. 18, figs. 90, 91. Distribution.-Mouth of Gambia River, W. Africa. Remark.-In 1885, von Maltzan described a species of Acteon from Goree, Senegal, and called it Actaeon (Amathis) senegalensis, spec. nov. This unfigured species was allotted to Leucotina by Pilsbry (1893-95: 172).

+ 16. "Acteon" danaida Dall, 1881 References.-Dall, 1889: 42, pI. 17, fig. 12.-Pilsbry, 1893-95: 160, pI. 20, fig. 32.-Marcus, 1970: 924, figs. 3, 4. Distribution.-Cuba; off Tortugas, 620 m; Brazil, off Alagoas, 09° S, 34° 51' W, 370 m. + 17. "A cteon" amabilis Watson, 1883 References.-Watson, 1886: 629, pI. 47, fig. 4.-Pilsbry, 1893-95: 154, pI. 20, figs. 27, 28.-Dautzenberg & Fischer, 1897: 142.-Strebel, 1905: 577. Distribution.-West of Azores; Canaries, 823-2061 m. + 18. "Acteon" hebes Verrill, 1885 References.-VerrilI, 1885: 428, pI. 44, fig. 15.-Dall, 1889: 41; 1927: 19.-Pilsbry, 1893-95: 159, pI. 19, fig. 12. Distribution.-Off North Carolina and Georgia, 4707 m. Remark.-Dall identified hebes with melampoides (1889), but in 1927 called two young specimens A. hebes.

+ 19. "Acteon" deUcatus Dall, 1889 References.-Dall, 1889: 41, pI. 17, fig. 5.-Pilsbry, 1893-95: 162, pI. 20, fig. 35. Distribution.-Gulf of Mexico; Cuba; off Barbados, 133-731 m. Remark.-The species is probably synonymous with "A." turritus (No. 10).

+ 20. "Acteon" monterosatoi Dautzenberg, 1889 References.-Dautzenberg, 1889: 20, pI. 1, figs. 2a-2d.-Pilsbry, 1893-95: 155, pI. 19, figs. 1-3. 318 Bulletin of Marine Science [24(2) Distribution.-Azores, 1287 m.

+ 21. "Acteon" semisculptus E. A. Smith, 1890 References.--Smith, 1890: 298, pI. 24, fig. 8.-Pilsbry, 1893-95: 152, pI. 18, fig. 97. Distribution.--St. Helena.

+ 22. "Acteon" grimaldii Dautzenberg & Fischer, 1896 Reference.-Dautzenberg & Fischer, 1896: 397, pI. 15, figs. 1, 2. Distribution.-Azores, 2187 m.

+ 23. "Acteon" azoricus Locard, 1897 Reference.-Locard, 1897: 85, pI. 3, figs. 8-11. Distribution.-Azores, 1258 m.

+ 24. "Acteon" torrei Aguayo & Rehder, 1935 Reference.-Aguayo & Rehder, 1935: 268, pI. 24, fig. 8. Distribution.-Cuba, 16 m.

+ 25. "Acteon" finlayi McGinty, 1955 References.-McGinty, 1955: 81, pI. 2, fig. 10.-Rios, 1970: 130, pI. 49. Distribution.-Florida; Cuba; Brazil, to Rio Grande do Sul, 40-80 m. Remark.-The figured shells may just as well be MysoutJa cumingii (No. 4), but the growth lines appear a little stronger. Without the soft parts, the identity cannot be stated.

+ 26. "Acteon" (Lissacteon) nitidus Verrill, 1882 References.-Dall, 1889: 39 (syn. of exilis).-Pilsbry, 1893-95: 157, pI. 19, fig. 4 (A. nitidus).-Dautzenberg & Fischer, 1896: 399 (A. nitidus). -Lemche, 1948: 71 (syn. of A. exilis).-Clarke, 1962: 37 (A. nitidus). Distribution.-From Martha's Vineyard to Florida and the Gulf of Mexico, 275-2653 m.

Genus Leucotina A. Adams, 1860 + 27. Leucotina minuta E. A. Smith, 1890 References.-E. A. Smith, 1890: 298, pI. 24, fig. 9.-Pilsbry, 1893-95: 171, pI. 60, fig. 17. 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 319 Distribution.-St. Helena. Remark.-According to Thiele (1931: 233), Leucotina belongs to the Pyramidellidae. Genus Neactaeonina Thiele, 1912 References.-Thiele, 1912: 219.-Zilch, 1959-60: 10. The genus Acteonina d'Orbigny, 1850, was erected for a fossil species. Thiele (1912: 220) maintained it for fossil shells with unknown radulae, and so did Zilch (1959-60: 14). Thiele created Neactaeonina for Recent species with the radular formula 8.0.8 (Hoffmann, 1934-39: 1002). How- ever, Rudman (1971: 208) described a species with the formula 8.0.8 as Maxacteon and placed two other species with 11.0.11 and 13.0.13 teeth in the same genus (pp. 208-210, fig. 2, D, E, F). The teeth of their radulae differ from those of Neactaeonina cingulata (Strebel, 1908) (Thiele, 1912: 220, fig. 4) by the basal denticles, which are wanting in Neactaeonina.

+ 28. Neactaeonina chariis (Watson, 1883) References.-Watson, 1886: 633, pI. 47, figs. 7a, 7b (Actaeonina chariis). -Pilsbry, 1893-95: 174, pI. 49, figs. 5, 6 (Actaeonina chariis).-Dautzen- berg & Fischer, ]897: 142 (Actaeonina chariis). Distribution.-Azores, 1167-2101 m.

Genus Bullina Ferussac, 1821 + 29. Bullina exquisita McGinty, 1955 Rejerence.-McGinty, 1955: 82, pI. 2, fig. 9. Distribution.-Florida, Palm Beach, 91-109 m. Remark.-Rudman (1972b: 117) proposed a new family Bullinidae with features of both the Acteonidae and the Hydatinidae.

D. LIST OF UNFlGURED ACTEONlDAE WITH UNKNOWN RADULA A. pusillus (Forbes, 1843) (Pilsbry, 1893-95: 156); Gulf of Mexico and Caribbean A. globulinus (Forbes, ]843) (Pilsbry, 1893-95: 155); Azores A. semicingulatus DaH, ]927: ]9; Florida A. particolor DaH, 1927: 20; Florida A. juvenis Dall, 1927: 20; Florida A. liostracoides DaH, 1927: 20; Florida A. propius DaH, 1927: 20; Florida A. parallelus DaH, 1927; off Georgia and Florida A. lacunatus DaH, 1927; off Georgia. 320 Bulletin of Marine Science [24(2) Superfamily Philinacea Family Scaphandridae

A. SPECIES TREATED Genus Scaphander Montfort, 1810 The type-species of the genus, S. lignarius (Linne, 1758) has been studied anatomically by G. O. Sars (1878), Vayssiere (1879-80: 95-100), Pel- seneer (1894: 9-10), Bergh (1901: 263-272), Perrier & Fischer (1911: 72-126), Lloyd (1952: 26-31) and others. The spoon-shaped shell at- tenuated behind and with a wide anterior border is typical for a number of species. These are distinguished, as only the empty shells are known, mainly by their ratio of width to length and the characters of their spiral striation. Dall (1881: 99), describing S. watsoni, thought it "possible that this will prove to be a Philine when the animal is known, but the form and aspect are those of a Scaphander." Pruvot-Fol (1954: 73) deplored the "habit that has so long prevailed, to dissolve the soft parts to preserve the shells." The "bewildering variation in shell characters for young stages of wat- soni" (Bullis, 1956: 10) refers also to ratio and number of striae of the shells of other species. The ratio of width to length is in the present paper calculated for the entire length of the shell including the posterior extension of the outer lip, while Bullis considered only the length of the axis. The counting of the striae is subject to different results due to the inequidistant grooves, and because intermediate ones arise between them. It also depends upon the sector of the body whorl that is counted: they are more numerous on the outer lip than farther inward. The aspect of the punctation is dependent upon the angle of the illumination. My attempts to classify the empty shells of the present collection failed many times, as I could not distinguish the shells of the lots of S. watsoni from those of S. darius. These could only be separated by the quite different male organs (Figs. 41, 64). The radula, with the formula 1.1.1 for all species, does not offer specific characters, nor does the degree of "pinching together" of the unpaired gizzard plate (Bullis, 1956: fig. 5,b,c). In three specimens of lot G-606, one had an open unpaired plate, the second had it half open, and in the third it was tightly pinched together. The lateral gizzard plates also show considerable intraspecific variation (Bullis, 1956: 4). The penial organ of S. darius is armed with a circle of big warts and a bulb covered with small warts. It is of such different aspect according to the degree of retraction, invagination, or evagination (Lloyd, 1952: 28) that this generally so reliable character is only to be used for its armature, not for its shape. I prepared 28 organs of S. darius for comparison, to find out that the apparent differences are due to age and state of eversion. The nature of the armature, indicated as cartilaginous cones with growth lines 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 321 for !ignarius by Vayssiere (1879-80: 99), is in the present species muscular with a thin cuticula; the growth-line-like aspect is brought about by circular muscle fibres (Fig. 66). A character in common for the species I studied is the presence of the Blochmann's glands in the mantle border (Perrier & Fischer, 1911: 118- 123, figs. T, U). Hoffmann (1934-39: 299, fig. 199) erroneously called them Bohadsch's glands, but on pp. 426 and 430 he marked them properly as Blochmann's glands. Also the intrapallial gland is always present, though of varying size (Hoffmann, pp. 454, 456, designated it as interpallial). Several species of the genus Scaphander were allotted to other genera or subgenera by reason of the shapes of their shells. Dall (1890: 16) created the Section Bucconia for the globose S. nobilis and included the fossil Scaphander grandis Aldrich. "Bucconia is a thin-shelled Sabatia without a body callus." In 1927 (p. 26) Dall simply applied the generic name Scaphander to nobilis and his new species stigmatica. Bullis (1956) char- acterized the subgenus Bucconia by a pillarlike structure supporting the posterior extension of the outer lip, and included Scaphander stigmaticus Dall, 1927, and S. mundus Watson, 1883. This pillarlike structure is not always distinct. It occurs also in S. (Sabatina) bathymophilus (Dall). Habe (1955: 69,70) allotted several Recent Pacific species to the genera Bucconia and Sabatia. Zilch (1959-60: 29) retained the subgenus Buc- conia. The radulae of Thiele's Roxania simillima and R. (Sabatia) supra- cancel/ata (1925: 243,244, pI. 34, figs. 11, 12) are so far different from those of all known Scaphander species that they certainly belong to other genera. Dall originally described the species bathymophila (1881: 98) as A tys ? In 1889 (p. 53) he assigned it to the fossil genus Sabatia Bellardi, 1867; though its general form is that of a Scaphander, the minute sculpture and the characteristics in detail are distinct. Dall made Sabatia a subgenus of Scaphander. In] 908 he changed the position from the fossil Sabatia to the new Section Sabatina, as the "Recent species, without exceptions, differ from Bellardi's fossil one. For the globose Recent species, therefore, I propose the name Sabatina with S. planetica Dall (1908: 241) as type." I follow Dall and consider Sabatina as a subgenus of Scaphander, as it is inappropriate to identify fossil shells, whose anatomy never will be known, with Recent forms. Thiele (] 925: 244) allotted the subgenus Sabatia to Roxania Leach, to which he assigned Atys supracancel/ata Schepman (1913: 470, pI. 32, fig. 4). Thiele figured the radula of the latter (1925: pI. 34, fig. 12), which is quite different from that of bathymophilus, and did not mention the presence or shape of gizzard plates in this or in his new species Roxania simillima from Southwest Africa (1925: 243, pI. 32, fig. 17, pI. 34, fig. 11), which also has a different radula. Bullis (1956: 2) considered bathymophilus as "a new and distinctive genus of the Scaphan- 322 Bulletin of Marine Science [24(2)

FIGURES 31-34.-31, Scaphander lignarius (Linne), diagram of male copulatory organ (after Lloyd, 1952) .-32, Scaphander? gracilis Watson, shelI.-33, Sca- phander punctostriatus (Mighels), sheI1.-34, Scaphander clavus Dall, male copulatory organ. (I, flagellum.) dridae." Thiele (1931: 391-92) synonymized Sabatina with Sectio Sabatia s. str. of Sabatia. Locard (1897: 49) accepted Monterosato's broad concept of the Bu1- lidae against Pilsbry's restricted conception of Bulla only (1893-95: 326- 350). Locard (1897: 54, pI. 1, figs. 19-22) united quite different types of shells in his genus Bulla. His western and central Atlantic species known only for their shell with punctate spiral sculpture are insperata with puncto- striate sculpture, from 25t 8-2635 m in the Canaries, and Bulla semilaevis Seguenza, 1879 (Watson, 1886: 638; Locard, 1897: 57, pI. 1, figs. 3-6), sculptured with very few striae, from 823-1829 m in the Azores. Locard's Bulla millepunctata (p. 52, pI. 2, figs. 3-6) from the Azores (Fig. 56) is so much like Scaphander stigmaticus that it is evidently the same species. Locard's Bulla pinguicola Jeffreys (Locard, 1897: 58, pI. t, figs. 26, 27) might be a globose Scaphander with the greatest breadth behind the middle. 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 323 As the species had not been described by Jeffreys, Dall identified it from the type-specimen with his Bulla abyssicola (1881; 1889: 56, pI. 17, fig. 11; 1908: 244: Leucophysema abyssicola). Dall's figure of a 12.75-mm- long, globose shell with dark spiral bands and striae differs widely from his later description (1927: 25) of abyssicola based upon six shells from off Fernandina, 4.5 mm long and white, which he considered as probably young of abyssicola. Dall's figure (1889) is much more like Hydatina than like a Bulla or a Scaphander. Dall did remark: "as the soft parts are yet un- known, it is possible that this species is more nearly related to the preceding family" (the Scaphandridae). Zilch (1959-60: 38) placed Leucophysema as subgenus to Bulla, with the type-species abyssicola. The genus M eloscaphander Schepman (1913: 464) with the species sibogae (pI. 31, figs. 5-9) and M. grandis Minichev (1967: 131-134, figs. 25-29) is well characterized by its exserted spire. The families Scaphandridae and Philinidae are distinguished by: a free shell (Scaphandridae) and a shell covered with the mantle (Philinidae); the animal almost completely retractile into the shell (Scaphandridae), or much larger than the shell (Philinidae); pallial caecum present (Scaphan- dridae), or absent (Philinidae). The radula and gizzard plates are very similar in both families. The present collection contains a great number of lots consisting of empty shells only. In some exceptional cases it was possible to identify them with figures of previously described shells, but the classification remains un- certain. The greater number might be S. watsoni or darius. The Atlantic species of Scaphander may be separated by the shapes of their shells, ovoid in bathymophilus, mundus, nobilis, stigmaticus, spoon- shaped with narrow apex in lignarius, watsoni, and darius; clavus, gracilis and punctostriatus are intermediate. The punctations in the grooves be- tween the striae are uninterrupted in watsoni, loisae and darius, interrupted in the other species.

KEY TO THE SPECIES OF Scaphander WITH KNOWN MALE ORGAN 1. Prostate gradually narrowing towards prostatic duct ...... 2 1. Prostate set off from prostatic duct .. 4 2. Penial papilla smooth ...... _.. bathymophilus (Figs. 67-80) 2. Penial papilla warty .__...... _ 3 3. The warts are spiny cones; shell spoon-shaped lignarius (Fig. 31) 3. The warts are round knobs; shell ovoid __.. nobilis (Fig. 50) 4. Penis stout ...... darius (Fig. 62) 4. Penis flagelliform . .. ______5 5. Penial base smooth watsoni (Fig. 40) 5. Penial base spiny .. .. clavus (Fig. 34) 324 Bulletin of Marine Science [24(2) TABLE 1 COMPARATIVECHARACTERSOF WESTERNANDMIDDLEATLANTICSPECIES OF Scaphander

Length Breadth Ratio (mm) (mm) (%) Striae Sculpture Pinch bathymophilus (No. 39) 29 20 62.5-77 64-100 punctations +- touching one another mundus (No. 33) 33 23 69.7 85-120 distant im- + pressed dots nobilis (No. 35) 11-39 7-24 68.6-73.7 40-50 sunken dots + stigmatcus (No. 37) 38 26 68.4 105 interrupted ? gracilis (No. 34) 30 19 63.3-71 100 remote stip- ? plings clavus (No. 36) 18.5-30 14.5-17 60-68.5 85-100 rectangular + confluent punctations punctostriatus (No. 31) 35 21 60-63 distinctly ? punctured grooves lignarius (No. 30) 60 26 52-55 continuous watsoni (Dall, 1889) 8.75 49-60 25 not punctate ? watsoni rehderi (Bullis, 1956; 39.3 21.4 50-54.7 85-100 scalloped- +- Marcus, 1967) ) edged 1 J I. ,I! Iij j 1.'/'/ I i./ "LXllfll, grooves watsoni (No. 32) 26 12 44.2-49.1 55-88 clear-cut +- scalIoped grooves darius (No. 38) 7-23 3.4-12 40.6-59.1 35-100 scalloped- +- edged grooves 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 325 + 30. Scaphander lignarius (Linne, 1758) Fig. 31 References.-G. O. Sars, ]878: 292, pI. ]8, fig. 7, pI. 26, fig. 4, pI. XI, figs. 13a-h.-Pilsbry, 1893-95: 245, pI. 31, figs. 17,21-23, pI. 32, figs. 24, 25.-Vayssiere, 1879-80: 73-113, figs. 89-94, 99.-Bergh, 1901: 263-272, pI. 21, figs. 16-38, pI. 22, figs. 1-13, pI. 24, fig. 1, pI. 26, fig. 10.-Lemche, 1948: 19, 59, 86, figs. 6, 38.-Lloyd, 1952: 26-31, figs. 6, 7.-Clarke, 1962: 40. Distribution.-Atlantic, from Norway to Gibraltar; Mediterranean; Ca- naries, 36-2250 m. Remarks.-Vayssiere (1879-80) described the male organ as consisting of three parts, entally the cylindrical, glandular prostate, 8 mmlong and 3 mm wide; then a short duct with longitudinal folds; and, third, the wide male atrium with thin walls. This contains entally around the prostatic duct a fleshy mammilla covered with small papillae, 200fL long, almost cartilagin- ous, with growth striae. Unfortunately, only an unopened male organ is seen in his figure 86, while figures 96-98 were omitted. These should have illustrated: fig. 96: penial sheath opened, showing the prostatic duct and the papillous mammillae which substitute the penis; fig. 97: three isolated papillae; fig. 98: highly magnified tip of one of these papillae. Bergh (1901: 262-263,270, pI. 21, fig. 18) drew the penial cones and mentioned them in his diagnosis of the genus. Lloyd (1952) gave figures of the inner reproductive organs and of the male apparatus. As her thesis was not published, I have copied (Fig. 31) her figure for comparison with the other species. Lloyd described the different aspects of the retracted and inverted and the everted penis. In lignarius, the protruded organ is hammer-shaped, resembling that of Philine. While Vayssiere called the papillae on the penis almost cartilaginous with growth striae, Lloyd found them as conical nodos- ities, each with a basal gland.

31. Scaphander ? punctostriatus (Mighels, 1841) Fig. 33 References.-G. O. Sars, 1878: 292, pI. 18, fig. 6, pI. XI, figs. 14a,b.- Watson, 1886: 642.-Pilsbry, 1893-95: 246, pI. 31, fig. 16.-Locard, 1897: 45.-non S. punctostriatus DaH, 1889: 52 (= clavus).-Bergh, 1901: 272, pI. 22, figs. 14-17 (rhachidian plates).-Lemche, 1948: 19, 59, 87, fig. 7.-Dall, 1903: 86, pI. 72, fig. 4.-? Abbott, 1955: 281, pI. 26,O.-Bullis, 1956: 5,9, figs. Ie, 2C.-Barnard, 1963: 322. Material.- P-606: 18° 45' N, 87° 37' W, 715-787 m, 17. III. 1968. 326 Bulletin of Marine Science [24(2) Further Distribution.-Arctic Seas to West Indies; Norway, Mediterranean; Sargasso Sea; Azores; Cape Town; 37-2106 m. Remarks.-One empty and incomplete shell measures 33.5 mm long and about 21 mm in breadth. It is so worn that the spiral grooves cannot be counted. It seems to suit best Sars's figure 6 (pI. 18), due to the flattened apical attenuation. It has a broken and reconstituted outer lip, so that the shape of its apical extension is not safely recognizable. Abbott's figure differs from that of Sars by the lack of a posterior exten- sion of the outer lip beyond the apex, and by the roundish curve of the apical attenuation. The classification of empty shells is risky, but as the species has fre- quently been reported from the Caribbean, its identification is quite prob- able.

32. Scaphander watsoni Dall, 1881 Figs. 35-41, 81, 87 References.-Dall, 1889: 52, pI. 17, fig. 10.-Pilsbry, 1893-95: 248, pI. 31, fig. 18.-McGinty, 1955: 83, pI. 2, fig. 9.-Abbott, 1955: 281, pI. 26,M.-Bullis, 1956: 10 (watsoni watsoni), 13 (watsoni rehderi), figs. 1, 3-5.-Marcus, 1967a: 602, figs. 5-9 (watsoni rehderi).

Material.-P-1357: 150 14' N, 81026' W, 249-256 m, 31. I. 1971, six complete animals and six empty shells. Further Distribution.-From Cape Hatteras to Venezuela, Los Testigos, 54-915 m.

Description.-The shells measure from 13 X 5.8 to 26 X 12 mm. Their ratio is 44.6 per cent in the smallest, 44.2 per cent in one 26-mm shell, and 49.1 per cent in a 22 X 10 mm shell. The shells are white; only the three largest with the animals in them have a yellowish brown band along the columella. The flesh of the animals is whitish to light greyish brown. The depressed apex is surrounded by a keel; about one and a half whorls are visible, but the pit is closed by a callus which is continued onto the inner lip. A narrower columellar border is reflected. The posterior exten- sion of the outer lip hardly surpasses the length of the axis. The number of spiral striae, which in several lots of other species increases with size, is, in the present lot, highest (84-88 striae) in shells 20 to 22 mm, and lowest (55 striae) in the 13-mm shell. They are closer together towards the ends than in the middle. Hence an indication of the average number of striae per millimeter (Bullis, 1956: 10) is not feasible. The spiral sculpture consists of inequidistant, narrow, straight grooves (Fig. 35). Their struc- 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 327

35

FIGURES 35-41. Scaphander watsoni Dall: 35, sculpture of shell; 36, paired gizzard plate from inside (left) and outside (right); 37, unpaired gizzard plates from outside (left) and side (right); 38, two rhachidian teeth; 39, one lateral tooth; 40, tip of flagellum; 41, male copulatory organ. (t, flagellum.) ture appears different according to low or high power and the angle of illumination. Frequently the bottom of the groove is divided into puncta- tions corresponding more or less to the fine growth lines. The intrapallial gland is small, but distinct. The pallial caecum is short and flat; it appears triangular. The radula of the studied 26-mm animal has 21 rows. The brown lateral teeth (Fig. 39) are about 550J.L long and bear irregular denticulations on 328 Bulletin of Marine Science [24(2) both sides. The transparent and caducous rhachidian teeth are 160j.t in breadth. They have a short median cusp and ear-like angles (Fig. 38). The crop contains globigerines and Foraminifera arenacea, up to 5 mm in greatest diameter. The paired gizzard plates are 8 X 7 and 8 X 6.5 mm. They are not exactly mirror images of each other, but slightly different in shape (Fig. 36). They have a transparent middle area with a brown ring around it. The unpaired plate (Fig. 37) is strongly pinched together. Its measurements are 6 mm long, 2.5 mm high, and 1.2 mm broad. The male copulatory organ (Fig. 41) of the dissected 26-mm specimen is 12 mm long; the oblong prostatic gland is 4 mm long, its slender duct with inner, longitudinal folds, 5 mm, and the male atrium, 3 mm. The male organ agrees with that described for Scaphander w. rehderi (Marcus, 1967a: 603, figs. 8, 9) of a 33-mm animal. It is a little smaller, but the diverticulum on the flagelliform penis (1967a: fig. 8) occurs also on the present specimen (Fig. 40). There are no papillae nor spines. Discussion.-Bullis (1956) separated the subspecies watsoni and rehderi by reason of their different ratios (48-53 per cent and 53-57 per cent) of breadth to length of the shell. The ratio is intermediate in S. pilsbryi (Bullis, fig. 4), and there are slightly more striae per millimeter of length in speci- mens of 25 mm or larger of rehderi, but this character intergrades com- pletely. However, in Bullis's specimens of rehderi the unpaired gizzard plate had the dorsal edges pinched together. Also, this character is variable, as already shown in Bullis's figures 5b and 5c. The present animals have a slender shell like watsoni, numerous striae like rehderi, and a perfectly pinched gizzard plate like the latter. I think that this "bewildering varia- tion" forces the abandonment of the subspecies. McGinty (1955: 82, pI. 2, fig. 8) described S. pilsbryi from Pensacola, Florida, 120 m. He distinguished it from watsoni by the "less pinched-in" appearance of the apical end of the shell and clearly punctate spiral grooves. His description of the apical end evidently refers to a shorter posterior expansiOn.

33. Scaphander mundus Watson, 1883 Figs. 81, 84 References.-Watson, 1886: 643, pI. 48, fig. 2.-Locard, 1897: 44.- Marcus, 1966: 156, figs. 3-7. Material.-P-34: 03° 53' N, 02° 33' W, 1984-1948 m, 29. V. 1964, eight and one empty shells. Further Distribution.-Canaries and Cape Verde Islands; Ivory Coast; east of Ceylon; Am Islands (original locality); 1435-2334 m. 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 329

Remarks.-The eight shells measure from 15 X 12 to 33 X 23 mm, the ratio is 69.7-73.3 per cent. The greyish shells are more solid than those of the original description, as were also those from the Ivory Coast (Marcus, 1966). They have a fine spiral sculpture consisting of interrupted puncta- tions in narrow grooves. The shape of the punctations varies according to their position on the shell. There are about 150 grooves narrower than the higher intervals. The fine growth lines are distinct. Along the outer lip of one shell there is a varix-like different structure, and 7 mm inwards an older one of the same kind. These are possibly repairs of damage. The columella is reflexed and the inner lip covered by a callus which hides the apex and extends onto the two varices. In the broken specimens there are many tubes encrusted with foraminifers and inhabited by polychaetes. One shell in the same lot was much more delicate than the rest. It was more than 41 mm long, but was broken. It was white, and its roundish punctations were larger and farther apart in the spiral direction than those in the other specimens; the intervals between the grooves were smaller. This shell had a greater likeness to that of S. nobilis (No. 35), but on one empty and broken shell I do not dare to base a classification of a species known only from the western Atlantic.

34. Scaphander ? gracilis Watson, 1883 Fig. 32 References.-Watson, 1886: 645, pI. 48, fig. 4.-Pilsbry, 1893-95: 247, pI. 31, figs. 19, 20.-Locard, 1897: 47, pI. 1, figs. I5-18.-Clarke, 1962: 40. Material.-P-770: 12° 55'N, 71°46'W, 1318-1299 m, 28. VII. 1968, two empty shells. Further Distribution.-Azores, 2178-2995 m.

Descriptive Notes.-The white shells measure 30 X 19 and 12 X 8.2 mm. The smaller one is very young and fragile, the larger one is rather solid. Its spiral sculpture consists of about a hundred punctate grooves. The inner lip is covered by a strong callus which is continued onto the reflexed columella. Remarks.-The decision as to which of the known species these shells are to be ascribed was difficult, as there is another, incomplete, shell in the collection (P-606), only slightly different, and I identified that one with the figure of punctostriatus (No. 31). The present shells might also be a variation of punctostriatus, though they have a rounder shoulder than the other specimen. 330 Bulletin of Marine Science [24(2)

FIGURES 42-50. Scaphander nobilis Verrill: 42, shell; 43, sculpture of shell; 44, rhachidian tooth; 45, lateral tooth; 46, pallial caecum; 47, anterior part of digestive tract and male organ; 48, gizzard opened from anterior end; 49, gizzard plates; 50, male copulatory organ. (a, male atrium; b, cerebral ganglia; c, pallial caecum; g, gizzard; i, infrapallial lobe; k, gill; 0, oesophagus; q, prostate; r, salivary gland; s, suture.) 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 331 35. Scaphander nobilis Verrill, 1884 Figs. 42-50, 81-83 References.-Verrill, 1884: 209, pl. 32, figs. 18 a-d.-non Scaphander nobilis Dall, ] 889: 53 (= stigmaticus) .-Dall, 1890: 16 (Bucconia no- bilis).-Pi]sbry, ]893-95: 249, pI. 32, figs. 31, 32 (Scaphander nobilis). -DaH, 1903: 86, pl. 64, fig. ]06 (Scaphander nobi/is); ]927: 26 (Scaph- ander nobilis).-Thiele, ]925: 3]9 (Scaphander [Bucconia] nobilis).- Abbott, 1955: 281 (Scaphander nobi/is).-Bullis, 1956: 6, figs. 2A, 2B (S. [Bucconia] nobilis).-Zilch, 1959-60: 28 (S. [Bucconia] nobilis).- Clarke, 1962: 40 (Scaphander nobilis). Material.-Brazil, off Rio Grande do SuI: 34° 25' S, 51° 25' W, 1080- 1140 m, I. 1972, one complete specimen, received from Dr. Eliezer de C. Rios, Rio Grande.-P-680: 08° 42' N, 55° 48' W to 09° 17' N, 5SO48' W, 3012-3237 m, 13. VII. 1968, one broken shell.-P-884: 11° 30' N, 60° 14.5' W to 11° 44' N, 60° 11.2' W, ]463-1847 m, 1. VII. ] 969, one broken specimen.-P-1181: 18° 51' N, 73° 30' W to 18° 46.8' N, 74° 35.9' W, 2550-2180 m, "where Bayer was completely coated with mud," 1. VII. 1970, five empty and partly broken shells. Further Distribution.-From Martha's Vineyard to Delaware Bay and 38° 44' N; Gulf of Mexico, 28° 47' N, 87° 50' W; Canaries Basin, 24° 35.5' N, 17° 04.7' W, 1667-2823 m. Description.- The white shell from Rio Grande do SuI is 11 mm high and 7 mm in greatest diameter, which lies behind the middle. The Antillean empty shells measure 19 x 14, 35 x 24; and 33, 36, and 39 mm in length; the ratio of the complete shells is 73.7 and 68.6 per cent. The shells were in part still covered with the periostracum, which had become dirty white. The spiral sculpture conforms to that of the complete specimen from Brazil. The columella is distinctly reflexed. The single shell of P-680 is too far broken to permit a safe classification. The sculpture (Fig. 43) consists of 40-50 spiral rows of roundish or oblong flat pits, separated by radial bars narrower than the pits. The high spiral striae between the rows of pits are also mostly narrower than the pits. New rows of pits arise between the original ones. The smooth outer lip is a little higher than the insunken apex, which is closed by a thin callus. The slightly concave columella bears a narrow reflexed callus (Fig. 42). The animal is white with a brownish digestive gland. It is about 9 mm long. The head shield is 4.2 mm in breadth and 1.8 mm in anteroposterior direction. It is slightly notched in the midline of the anterior and the posterior borders. The quite short pallial caecum (Fig. 46) is curled. The intrapallial gland is small. The radula has 14 rows, 1.1.1; the curved laterals (Fig. 45) have broad 332 Bulletin of Marine Science [24(2) lobes on the outer side and a row of fine denticles on the cutting edge. Their length is about 31O,u. The rhachidian tooth (Fig. 44) measures 40,u X 40,u. It has a tiny pointed median cusp and two prolonged blunt lateral angles, the inner sides of which bear some irregular spines. The salivary glands are short. The gizzard plates have a transparent central area like a window through which the contents of the gizzard are visible. In the present specimen it showed foraminifers. The paired plates measure 4.0 x 3.3 mm. The trans- verse muscle bands are inserted on the convex inner sides and do not extend over the concave outer sides. The unpaired plate is slightly pinched together. It is 3 mm long and 1 mm wide. The male copulatory organ (Fig. 50) of the single small specimen is not fully developed. The prostate is composed of small glandular sacs enveloped in a thin, common muscular layer. From the coalescence of their lumina arises the short prostatic duct, whose walls bear longitudinal epithelial folds. In the male atrium, the duct ends with a papilla covered with soft knobs. Probably these will develop to the soft spines which occur on the penial papilla of other species of the genus, e.g., clavus and darius (Figs. 34, 60-66). Also on the walls of the atrium there are some knobs.

Diseussion.- The present Brazilian shell was compared with the descrip- tions and figures of Verrill (1884), Pilsbry (1893-95), and Bullis (1956), and was found to conform with the shape, though not with the size, of S. nobilis. Verrill gave a detailed comparison with S. punetostriatus (No. 31) and figured also the radula and the gizzard. The description of the sculpture of the shell of nobilis corresponds exactly to the present shell, while the figure (18b) of the rhachidian tooth, though not drawn in detail, seems different. S. punetostriatus from Iceland to the Azores and South Africa (Barnard, 1963: 322), and from Maine to Culebra Island and Barbados, has a longer and heavier shell, narrowed posteriorly. It is covered with a brown perios- tracum. Only its rhachidian tooth is in Bergh's figure 15 more like that of the present animal than that of Verrill's figure. Bullis (1956) examined three specimens of S. nobilis from the Gulf of Mexico. He did not succeed in finding the rhachidian tooth of the radula, and even "definitely denies" its existence, in accordance with Cooke (1895: 230, fig. 137A) for S. lignarius. Sars had figured it for S. !ignatius in 1878. Sars's figure was copied in several treatises. The central tooth of S. punetostriatus was drawn by Bergh (1901: pI. 22, fig. 15), and for watsoni rehderi by Marcus (1967a: fig. 7). It was Bullis, who "was in error," not Dall, who, as Bullis said, "claimed that the rhach- idian tooth of nobilis is proportionally larger than in the typical species of Seaphander," and it was Bullis who misinterpreted Verrill's figure 18b of 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 333 the rhachidian tooth of nobilis. It is true that the central teeth of Scaph- ander are known to be caducous (Bergh, J901: 262; Schepman, J913: 465; and others), and in Bergh's material they had all fallen off, but Bergh found them in the radular sac. This is the first record for Brazil.

36. Scaphander clavus Dall, 1889 Figs. 34, 81, 85 References.-Dall, 1889: 52 (Scaphander punctostriatus var. clavus, par- tim).-Pilsbry, 1893-95: 246 (S. punctostriatus var. clavus).-Bullis, 1956: 8, fig. 2,1 (S. clavus).-Marcus, 1967a: 599, figs. J-4 (S. clavus). Material.-P-673: or 56.5' to 08° 08' N, 54° 39'-36' W, 1042-1070 m, 11. VII. 1968, one specimen and one empty shell.-P-607: 18° 30' N, 8r 37' W, 731-787 m, 17-18. III. 1967, two empty shells. Further Distribution.-Northern Gulf of Darien: 357-1050 m; ? Azores (see Marcus, 1967a: 599).

Description.-The roundish shells measure 22 X 14.5 and 27.5 X 17 mm, respectively; their ratio of width to length is 65.9 and 61.8 per cent. The smaller empty shell is worn smooth. Its spiral grooves are only recognizable on the inner lip, which is covered farther inward by a strong callus. The largest shell contains the animal. It has a callus filling the apical pit and covering the inner lip. There are about 120 spiral striae, separated by straight grooves, of different width and distance from one another, accord- ing to the degree of their development. A few grooves are formed of con- fluent punctations, never of interrupted punctations. All striae are filled with the light brown periostracum, which is thicker and darker near the fore end of the shell. The whitish, 25-mm animal is not longer than the shell. The head shield is notched in front, 7 mm Jong and 12 mm in breadth; the posterior lobes are separated by a slight notch. The flat pallial caecum is 7 mm long; it forms a complete circle and is longer than that of S. punctost,.iatus (Perrier & Fischer, 1911: 75, fig. L). As in other species of Scaphander, the gland of the mantle border consists of Blochmann's glands (Perrier & Fischer, 1911: 101, pI. 7). The intrapallial gland is like that in S. lignarius and S. punctostriatus (Perrier & Fischer, 1911: 118, figs. T, U). The radula of the present specimen was lost. It was described and figured for material from the Gulf of Darien (Marcus, 1967a: 601, figs. 6,7). The paired gizzard plates lay one on the dorsal, the other on the ventral, side of the gizzard, but this position may have been that of the moment of fixation. Their measurements are 9 mm in length, 7 mm in breadth. The 334 Bulletin of Marine Science [24(2) pinched unpaired plate was on the left side; it is 6 mm long and 1.2 mm in breadth. The male organ (Fig. 34) corresponds to our first description. It is 15 mm long; the prostate is 4 mm long, its duct, 3 mm, and the atrium, 8 mm. From the sinuous bulb hangs a 2.3-mm-long, slender flagellum with a few spines on its tip. Remarks.-The single specimen of Scaphander loisae Bullis (1956: 8) differs from clavus by a minute apical pit, which is broad and generally filled with a heavy callus in clavus. Bullis (pp. 8-9) described the broadly eroded striation of dead shells of clavus with no trace of periostracum as inequidistant, and its squarish or rectangular punctations as variable in size. As long as no further specimens of loisae are found, I consider it as in- sufficiently described.

37. Scaphander stigmaticus Dall, 1927 Figs. 51-56 References.-Dall, 1889: 53 (S. nobilis, part.); 1927: 26 (S. stigmatica). -Locard, 1897: 52, pI. 2, figs. 3-6 (Bulla millepunctata).-von Martens, 1903: 15, pI. 5, fig. 20 (Atys millepunctatus).-Bullis, 1956: 6, figs. 2D,E (Scaphander [BucconiaJ stigmatica). Material.-P-1138: 20° 51.8' N, 74° 22'-18.6' W, 2758 m, 12. I. 1970, one dry shell. Further Distribution.-South of Cuba, 3000 m, the single, dead type-speci- men. Off Senegal and Azores (Locard), 1495-4255 m, many samples; Cameroons (von Martens). Remarks.- The incomplete dry shell (Fig. 51) measures 36 mm in length. The greyish periostracum is preserved; it shows some broad, longitudinal stripes accompanying the outer lip. The distinct spiral sculpture consists of about 105 grooves produced by round punctations touching one another. The interspaces are wider than the grooves in the middle of the shell (Fig. 52) and quite narrow in the anterior part (Fig. 53). The posterior exten- sion of the outer lip resembles that of nobilis more than that of Bullis's photographs of the type-specimen. It has a pillarlike support. From Bullis's figures 2D and E, I infer that part of the extension is broken off, so that it would have been broader and more' like that of nobilis, if it had been complete. However, the sculpture of the present shell has "much more closely spaced punctations" than nobilis, as Bullis stated for stigmatieus. The localities are so near together, that I risk the identification with Dall's species. On the other hand, the shape of the shell of "stigmatiea" in Bullis's 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 335

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FIGURES 51-56. Scaphander stigmatic us Dall: 51, shell; 52, sculpture near middle of shell; 53, sculpture near anterior end; 54, outlines of type-specimen (from Bullis's photographs); 55, outlines of present shell; 56, outlines of "Bulla millepllnctata" Locard (from Locard, 1897). photographs (Fig. 54) and that of the present broken shell (Fig. 55) con- form exactly to the drawings of Locard's Bulla millepunctata (1897: 52, pI. 2, figs. 3-6) from the Azores, Sudan, Sahara, and Senegal (Fig. 56). The superimposed traces of the outlines of the figures are identical. The ratio of Locard's snails (40 X 26 mm) and of the type-specimen in Bullis's figures (39.1 mm), as well as that of the present one, as far as recognizable, are comparable, and so is the sculpture. Schepman (1913: 466) is right not to consider the sculpture in Scaphander as a very constant character. The punctations differ considerably in some parts of the shell, and in differ- ent specimens of one and the same lot. However, these differences can be regarded as individual, "otherwise nearly every specimen should be a species" (Schepman, loco cit.). Bulla millepunctata Locard, 1897, was by von Martens (1903) con- sidered as Atys. Thiele tentatively united it with Scaphander nobilis, as S. stigmaticus had not yet been separated from nobilis. Thiele stated that the species is widely distributed in the Atlantic. I am convinced that Bulla millepunctata Locard, 1897, from seven localities, is a senior synonym to Scaphander stigmaticus Dall, 1927. As 336 Bulletin of Marine Science [24(2) nobody has used the name for more than 50 years, it need not now be revalidated.

38. Scaphander darius Marcus, 1967 Figs. 57-66, 81, 86 References.-Marcus, 1967a: 603, figs. 10-17; 1970: 925. Material.-From Stations P-625, 696, 714, 727, 728, 737, 749, 756, 757, 758, 760, 766, 768, 913, 1368, and G-592, 683, 1024, 1026, 1034, 1090, from the western Atlantic and the Caribbean, between 25° 52' and 08° 35' N to 50° 02' and 81° 27.8' W, in depths from 15-18 m to 374-393 m. Collected in the years 1965, 1968, 1969; 49 animals and more than 30 empty shells. Further Distribution.-Gulf of Darien; northern Brazil, 47-174 m. Description.-The shells of the present material are from 7-23 mm in length and 3.4-12 mm in greatest diameter. The ratio of width to length varies from 40.6 to 52.3 per cent, independent of the size of the shell (Table 2). The color of the shells varies from pure white to rusty brown. Frequently there is a brown stripe accompanying the columella. The outer lip is hardly extended posteriorly. A thin callus closes the apex and covers the inner lip. The spiral striae have scalloped borders and continuous, more or less distinct punctations on the bottom of the grooves. They are distributed more closely towards the ends of the shell than in its middle. Their number varies from 45 in a 17-mm shell to 150 in one of 19 mm, and 110 in the 23-mm shell. There are fine growth lines. The flat pallial caecum is broad and blunt. It is from one-quarter to half a whorl long. The intrapallial gland is rather small. The radula (Figs. 57-59) comprises 16-24 rows of 1.1.1 teeth, of which the small, transparent rhachidian is frequently lost. The shape of the latter varies; the median cusp is quite small or almost as big as the lateral corners, which are blunt or pointed (Figs. 57, 58). The horny yellow lateral teeth are unspecific; they are denticulated on one or both sides, or not at all. The short salivary glands have blunt, widened ends. The paired gizzard plates of the largest animal are 10 X 8.5 mm; all plates have an almost transparent central area and a brown cap on the inner side. The unpaired plate is as long as, and narrower than, the paired ones; the degree of pinching together of its borders is very different, not correlated to size or ratio or number of striae. The male organ consists of the three typical parts: (1) a prostatic vesicle with high walls consisting of glandular pouches, (2) a narrow prostatic duct with longitudinal folds of the epithelium; these wind spirally in the ectal part, where the duct enters (3) the wide atrium. The spiny penis is 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 337 TABLE 2 VARIABILITY OF CHARACTERS OF Scaphander darius ----.-- Length Breadth Ratio Sla. no. (mm) (mm) ('}'o) Striae Pinched P-758 12.5 5.5 44.8 70 + P-737 23.0 12.0 52.2 110 + P-625 14.5 6.5 44.8 105 + P-1368 15.0 7.8 52 105 + 19.0 9.0 47.4 150 + P-728 18.5 9.0 49 110 + P-760 13.0 6.8 52.3 110 + 11.5 65 + P-757 16.0 6.5 40.6 115 + 11.5 5.0 44.4 65 + P-749 17.3 8.5 49.2 95 + P-913 15.5 7.5 48.4 115 + G-1024 17.0 8.5 50 45 G-1034 14.5 7.5 51.7 50 G-592 11.0 5.5 50 75 G-696 13.0 6.2 47.7 80 G-I090 10.5 5.0 47.6 70 9.0 4.2 46.7 7.0 3.4 48.6 P-766 12.2 5.5 45.6 80 P-756 16.0 7.5 46.8 100 P-727 19.5 9.5 48.7 100 P-714 13.0 6.5 50 110 9.0 4.5 50 55 P-768 15.5 7.2 46.5 110 14.0 6.8 47.1 --- Maximum and minimum values italicized. lodged in the atrium. The limit between the wall of the atrium and the penial papilla bears a few big knobs, 350fL wide, each provided with several small cones. The penial papilla is eversible, and so far retractile that its warts, about 150fL to 200fL in length, are turned inward, lining a lumen (Figs. 63, 64). In the spine-shaped warts of darius I did not find any glands. The prostatic duct opens at the base of the penial papilla. Ectally a pouch coated with a thin, folded epithelium enters the papilla; probably 338 Bulletin of Marine Science [24(2)

FIGURES57-62. Scaphander darius Marcus: 57, lateral and rhacbidian teeth of P-737; 58, same of 0-636; 59, rhachidian tooth of P-756; 60, male copulatory organ of P-756, to show levels of sections in Figures 61 and 62; 61, section through the level of prostatic duct; 62, section through level of the seminal vesicle. (a, male atrium; d, hemocoel; p, penial papilla; u, prostatic duct; v, seminal vesicle; w, big wart.) it serves as a sperm reservoir. The wall of the atrium is thickened by muscle pads which border the seminal groove. Discussion.- The shells of Scaphander watsoni, watsoni rehderi, and S. darius are so much alike (Figs. 86, 87), that it is not possible to classify 1974] Marcus: Cephalaspidea tram Atlantic Warm Waters 339

FIGURES 63-66. Scaphander darius Marcus: 63, male copulatory organ, re- tracted and inverted, of G-1026; 64, same, everted, of G-1034; 65, three papillae; 66, section of papillae. empty shells if they do not belong to a lot with complete animals. Dall (1889: 52) said that watsoni resembles lignarius (Linne) more closely than any other species, and Marcus (l967a: 604) compared also darius with lignarius. The mentioned difference of the overtopping outer lip (Marcus, 1967a: 605) is variable (see Pilsbry, 1893-95: pI. 31, figs. 17, 21, 23, pI. 32, fig. 24). The shells of watsoni and darius are neither distinguishable by ratio or punctation, nor are the animals by the extremely variable rhachidian tooth 340 Bulletin of Marine Science [24(2)

~'.. : .. : ' ',' , I. : i:: i iI' j 70

FIGURES 67-70. Scaphander (Sabatina) bathymophi/Ils (Dall): 67, shell with animal; 68, animal without shell, in dorso-right-side view; 69, anterior part of digestive tract with paired gizzard plate; 70, unpaired gizzard plate. (b, bursa copulatrix; c, cephalic shield; e, oesophagus; t, foot; g, intrapallial gland; h, heart; i, intestine; k, gill; m, mantIe border; n, kidney; 0, columellar muscle; p, pallial caecum; r, radula; s, salivary gland; Il, suture; z, gizzard.) or the pinching of the unpaired gizzard plate. Bullis's illustrations (1956) of the unpaired plate of S. watsoni rehderi (fig. 5,b,c) differ in the degree of pinching, and the present lots, from everyone of which I dissected at least one sample, show all intergrades, from broadly open, to walls touch- ing one another. So the "pinched together," that Bullis (p. 14) considered as a unique feature of rehderi, is not a subspecific nor a specific character. The only character that I found reliable is the male organ, protractile and beset with cones in darius, smooth with a noninversive flagellum in watsoni. Vayssiere's description of the almost cartilaginous papillae, with growth striae, on the penis of lignarius (1879-80: 99) does not hold for those of darius, the circular striae of which are produced by muscle fibres (Fig. 66). 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 341

FIGURES 71-80. Scaphander (Sabatina) bathymophilus (DaB): 71, lateral tooth; 72, rhachidian tooth; 73, male copulatory organ; 74, diagram of same to show levels of six sections, Figures 75-80; 75-80, cross sections at various levels of male copulatory organ.

39. Scaphander (Sabatina) bathymophilus (DaH, 1881) Figs. 67-82 References.-DaH, 1881: 98 (Atys? bathymophila) .-DaH, 1889: 53, pI. 17, figs. 9, 9b (Scaphander subg. Sabatia b.) .-Pilsbry, 1893-95: 256, pI. 32, figs. 27, 28 (S. subg. Sabatia b.).-DaH, 1903: 86, pI. 17, figs. 9, 9b (S. subg. Sabatia b.).-Thiele, 1925: 244 (Roxania subg. Sabatia).- DaH, 1927: 25 (Sabatia, Section Sabatina b.) .-Thiele, 1931: 391 (Saba- tina syn. to Sectio Sabatia s. str. of genus Sabatia) .-Johnson, 1934: ]47 342 Bulletin of Marine Science [24(2)

FIGURE 81. Ventral views of shel1s of: (top row, left to right) Scaphander (Sabatina) bathymophilus (Dall), Scaphander nobilis Verril1, and Scaphander mundus Watson; (bottom row, left to right) Scaphander clavus DaH, Scaphan- der darius Marcus, and Scaphander watsoni Dal1.

(Scaphander subg. Sabatia b.).-Bullis, 1956: 2 (Sabatia bathymophilus). -Zilch, 1959-60: 27 (Roxania subg. Sabatia).-Clarke, 1962: 40 (Sabatia bathymophilus) .

Material.-P-892: 14° 17' N, 60° 45.2' W, 1280 ill, 7. VII. 1969; and G-148: 2Y 04'-07' N, 79° 30'-29' W, 805 ill, 23. VI. 1963, from each one complete specimen.-P-187, 586, 607, 689, 1187, 1304, and G-109, 293, 1357; from 2r 26' to 08° 42' N, 50° 45' to 87° 37' W, 713-3964 m, West Atlantic, Straits of Florida, and Caribbean, 1963-70, a total of 32 empty shells. Further Distribution.-Gulf of Mexico and Caribbean basin; Florida, Fernandina, 294-5120 m. Description.- The largest of the strong shells is 39 mm high and 25 mm in greatest diameter. The 2.S-mm-long posterior extension of the outer lip is included in the indication of the height. The size of the present shells 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 343

FIGURE 82. Scaphander (Sabatina) bathymophilus (DaB), dorsal view of shell. lies between 39 X 25 mm and ]4.0 x 9.5 mm; the ratio of width to length varies from 64-77 per cent, independent of the size. The periostracum is ivory white to dark brownish grey, and the shell beneath is white. The shape is ovoid (Fig. 82), the greatest width a little anterior to the middle. The outer lip is produced beyond the apex and is reinforced on the inner side, though not by a pillar distinctly set off, as such is said to characterize the genus Bucconia (Dall, 1890: 16). The apical depression is filled with a heavy callus that is continued along the inner lip. It is thickened with an irregular spiral crest in its posterior third and beset with flattened pustules (Fig. 67). The columella is reflexed, and there is a slight umbilical depression. The sculpture consists of spiral grooves, densely set near the apex, farther apart in front, and still farther in the middle of the shell. The interspaces, the striae, are always wider than the grooves. The spiral 344 Bulletin of Marine Science [24(2)

FIGURE 83. Scaphander nobilis VerriIl, dorsal view of sheIl. grooves are formed by uninterrupted round punctations; their borders are sometimes scalloped. The flesh of the two preserved snails (G-148: 28 x 19.5 mm shell; P-892: 20 X 13mm) is light with a darker digestive gland. The thickened mantle border (Fig. 68, m) is whitish. The small intrapallial gland (g) is round and firm. The short pallial caecum (p) is a flat triangle. The slender salivary glands (Fig. 69, s) are long and fastened at their pointed ental ends. The crop contained foraminifers. The radula has about 17 rows of stout, yellow lateral teeth (Fig. 71), 450/L high. The delicate, transparent, caducous rhachidian teeth (Fig. 72) are of variable measure- ments, 105/L, l30/L, 150/L, and 160/L high, and 135JL, l05/L, 105/L, and 125/L in breadth. 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 345

FIGURE 84. Scaphander mundus Watson, dorsal view of shell.

The gizzard plates correspond perfectly to the type of Scaphander. The paired ones (Fig. 69, z) of the largest animal are 8 x 7.3 mm. Its un- paired plate is 5.5 mm long and pinched together, so that its lateral borders are 0.5 mm from One another (Fig. 70). In the smaller snail, the paired plates measure 7 x 6 mm, the unpaired one, 5 mm in length, and the breadth of its outside opening is 0.8 mm. The male copulatory organ (Figs. 73-80) consists of a tubular prostatic ental part and a little narrower prostatic duct, together about 12 mm long. The duct enters with a bend (Fig. 74) into the small male atrium, 2 mm long. The opening of the prostatic duct lies on the tip of the papilla. In the atrium there are masses of small, longish nuclei which might be sperm (Fig. 76). In the prostatic duct there are big nuclei which might belong to special glandular cells (Fig. 75). The prostatic gland itself is subdivided into glandular pouches, whose nuclei are small and inconspicuous. 346 Bulletin of Marine Science [24(2)

FIGURE 85. Scaphander clavus Dall, dorsal view of shell.

Remarks.- The pustulate callus and the tubular prostate justify a sub- generic separation of Sabatina from Scaphander s. str. A distinctive name for the Recent forms, bathymophilus and "Sabatia" japonica Habe, 1952 (see Habe, 1955: 70, pI. 4, fig. 13), which have no special resemblance to the fossil type-species Sabatia isseli Bellardi, 1876 (Dall, 1889: 54), is preferable to uniting the Recent and the incompletely known fossil species generically.

ON THE GENUS Philine ASCANIUS, 1772 G. O. Sars (1878: 294, 298, 303), Pilsbry (1895-96: 3), Lemche (1948: 61), and Habe (1958: 120) arranged sections of the genus Philine 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 347

FIGURE 86. Scaphander darius Marcus, dorsal view of shell. according to the sculpture of the shell, whereas Thiele (1931: 393) and Pruvot-Fol (1954: 71) used the characters of the gizzard plates to separate sections or subgenera. I abstain from trying to classify the species described only from their shells (those of Dall even frequently unfigured), as the shells are shattered inside the animal in two well-defined species of the present collection, so that only the sculpture of the shells is recognizable on the fragments, not the shape. A good distinguishing character is furnished by the gizzard plates. Thiele (1931: 393) divided the genus Philine into sections. Philine (s. str.) has differently shaped unpaired and paired plates. In P. aperta and P. aperta guineensis, and in P. angasi (Rudman, 1970: 31, pI. 3,G, fig. 1A,B), where they were called cornered hat-shaped, they have a pair of pores in each plate. These pores are sometimes called perforations, though they do 348 Bulletin of Marine Science [24(2)

FIGURE 87. Scaphander watsoni Dall, dorsal view of shell. not reach the inner surface of the plates. In the mentioned species, the unpaired plate is smaller than the paired ones. Both aperta and angasi have the radular formula 0.1.0.1.0. The pores are given as a generic character in Pilsbry's diagnosis, but they are wanting in most other species. The paired plates are smaller than the unpaired ones in infundibulum, mera, and trachyostraca from the western Atlantic. Vayssiere was right when he pondered that if all species of the genus Philine were known, one might subdivide the genus into several, based upon some anatomical characters given by Sars (1878). Such great differ- ences are found in the structure of radula and stomach that it would be right to separate certain species totally from others. Thiele's section Her- mania contains the species with equally shaped and sized gizzard plates. 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 349 Of the western Atlantic species, this would refer to alba, amabilis, catena, fmmarchica, gibba, scabra, and thurmanni. Several of these have the outer side of the plate hollowed out on each side of a central, longitudinal, cigar- shaped bar, e.g., finmarchica (Fig. 90); auriformis Suter (Rudman, 1970: 24); P. d. kerguelensis Powell (1951: 178); thurmanni Marcus (1969b: 16); and P. powelli Rudman (] 970: 25). The plates of alba (Fig. 102) are very small, as are those (Fig. 99) of P. falklandica Powell (1951: 178). There are also several species without gizzard plates: sinuata, lima, and quadrata, and Pruvot-Fol (1954: 7]) proposed to separate them as Laona A. Adams, 1865, to which also P. pruinosa (Clark, 1827) belongs, with the exceptional radular formula 6.1.0.1.6. The radular formula does not offer good features for separation, as, for example, P. thurmanni thurmanni and P. thurmanni chilla are distinguished only by their marginal teeth, 1.1.0.1.1 and 0.1.0.1.0, respectively. A rhachidian plate occurs in P. gibba and P. falklandica (2.1.1.1.2), and Vayssiere noted it for P. quadrata (1913: 174). The transverse muscle layers of the gizzard might be distinctive. In finmarchica (Marcus, 1969b: fig. 13), P. cf. kerguelensis (Marcus, 1960: 890, fig. 24), and thurmanni there is a thick, continuous outer layer. In aperta, transverse muscle bundles are inserted on the sides of the plates and are on the outer side connected by a layer of connective tissue (Forster, 1934: 15, figs. 4,5; Brown, 1934: 189, fig. 18; Fretter, ]939: 604). An intermediate development is found in infundibulum (Fig. 96) and mera, while there is no distinct muscle layer in alba, nor are the small plates united by transverse muscle bundles. Many foraminifers and a little sand were the recognizable contents of the digestive tract in front of, and even behind, the gizzard. In several cases, where in the present collection the shell was damaged or the gizzard plates were lost, the shape of the penial papilla made it possible to determine the species. Perhaps the noncoiled but sausage-shaped prostatic gland (e.g., of lima, quadrata, gibba, falklandica, and alba) may serve to define a section. This accessory gland of the male apparatus is generally called a prostate, as in the other cephalaspideans. However, Forster (1934: 53) considered it to be a spermatophore-forming gland, as in one case he had found the masses of sperm in the glandular lumen included in a loose capsule secreted by the gland cells of the "prostate." None of the earlier authors, Vayssiere (1885), Guiart (1901), Bergh (1902, 1907), Pruvot-Fol (1930), Si (1931), Brown (1934), nor Hilda Lloyd (1952) mentioned spermato- phores in their detailed descriptions of Philine aperta. Lloyd (1952: 20) even observed, in a freshly dissected animal, a continuous stream of pros- tatic fluid pouring from the tip of the penis. The male apparatus of P. catena described by Vayssiere (1885: 35, pI. 350 Bulletin of Marine Science [24(2)

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40. Philine finmarchica M. Sars, 1858 Figs. 88-92 Reterences.-G. O. Sars, 1878: 296, pI. 18, figs. 10a-l0d, pI. XII, figs. la, Ib.-Pilsbry, 1895-96: 14, pI. 5, figs. 14-16.-Lemche, 1948: 64-66 (synonymy), 96 (references).-Marcus, 1969b: 10, figs. 13-16. Material.-P-672: or 37' N, 55° 22' W, 1220-1335 m, two damaged specimens. Further Distribution.-From Greenland to Cape Cod; Spitsbergen and Kara Sea to Murman and Lofoten, from 12-2220 m. Lemche supposed (1938: 13) that the records from great depths probably refer to dead specimens. Description.-The animals are about 18 mm long and 9 mm in breadth (Fig. 89). The shells are broken to small pieces, but their fine spiral striation is visible; it consists of grooves, not of pits (Fig. 88). The cephalic shield is 8 mm long, narrow and slightly pointed behind. The mantle shield is badly damaged in both specimens, so its shape is not recognizable, except that it is broader than the head shield. The proboscis (Hurst, 1965: 282) or buccal bulb (Rudman, 1970: 33) is everted in both animals, and one radula is missing. The radula has l8 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 353

FIGURES 88-92. Philine finmarchica M. Sars: 88, spiral sculpture of shell; 89, dorsal (left) and ventral (right) views of preserved specimen; 90, gizzard plate from outside (left) and inside (right); 91, radular tooth and denticles; 92, male copulatory organ and part of prostatic gland. (a, male atrium; d, ejaculatory duct; e, sperm duct; p, penial papilla; q, prostate; v, seminal vesicle.) 354 Bulletin of Marine Science [24(2) rows of 0.1.0.1.0 teeth. The pale lateral teeth (Fig. 91) measure 58511-. They are of the typical shape and bear 60-70 long and soft denticles. The large gizzard stands out of the damaged hind end in one of the specimens. It is 11 mm long and 5.5 mm in diameter, silky white all round, and the transverse muscle layer is not interrupted by the gizzard plates, as we figured it for finmarchica (1969b: fig. 13). The three brownish gizzard plates are slightly curved and of equal size, 10 x 3 mm; their ends are rounded (Fig. 90). Their outer side has a median, longitudinal, roundish "cigar-shaped" (Powell, 1951: 178) crest, flanked by flat margins with sharp borders. They have no pores. The center of the inner side is flat, with a median furrow; the margins slope evenly towards the borders. The contents of the stomach are broken foraminifers and small stones. From the male atrium, the male duct arises and forms a small seminal vesicle (Fig. 92, v). The ental end of the prostatic tube is connected with the atrium by a feeble strand. The middle section of the prostate is knobby, due to surrounding masses of prostatic tubules (q). The outermost, long, winding part is smooth and connects with the seminal vesicle. The penial papilla (p) is hammer-shaped, with two almost equal, smooth pointed tips. The wall of the atrium (a) bears some thickened pads. Discussion.-Lemche (1941: 21) considered P. finmarchica as a true Arctic species. As the present animals come from more than 1000 m depth, they had lived in temperatures similar to those in shallower water in higher latitudes. The present would be the first finding south of Nova Scotia, as the occurrence off Cape Cod, 29-164 m (Johnson, 1934: 149) was not mentioned by Lemche (1941: 21-22), and Franz says (1970: 175) it should be confirmed. The shell sculpture, radula, gizzard plates, and muscles conform to the descriptions of finmarchica and its synonyms. The structure of the prostate is by degree different from the knobby one figured (Marcus, 1969b, fig. 16) from the Labrador Sea. The penial papilla is not verrucose as in that material, but I call the present specimens finmarchica. Powell (1951: 178) described the shape of the three equal gizzard plates of Philine d. kerguelensis Thiele (1925: 279, pI. 32, figs. 22, 22A) from 47" S, 62° W, northwest of the Falkland Islands in 116-114 m depth. Their shape corresponds to the present ones, but neither size nor ratio are indicated. Thiele had based his species on two empty shells of 2.6-mm length, only. Powell's largest animal was 6 mm long and had a 4-mm-Iong shell, while the present animals are much bigger, 18 mm long, as Odhner's finmarchica (1907: 14). The small animals from the Maledive Islands (Marcus, 1960: 890, figs. 22-24) are 0.8-1.6 mm long; they have much broader gizzard plates, a ratio of length to width of 1.73: 1, against 3.3: 1 in the present material and 3 to 4: 1 in finmarchica (Marcus, 1969b: 10, fig. 14). 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 355 The second South Atlantic species of Philine with the radular formula 0.1.0. ].0 is P. aperta (Linne, 1767) (No. 44). Its gizzard is quite different from that of finmarchica and has an uninterrupted outer muscle layer, a narrow unpaired and two broader gizzard plates (Brown, 1934: fig. 17) with a pair of pores.

41. Philine infundibulum Dall, 1889 Figs. 93-97 References.-Dall, 1889: 57.-Pilsbry, 1895-96: 23.-Johnson, 1934: 149.-Marcus, 1967a: 606, figs. 18-22; 1970: 927 (Phi/ine d. infundib- ulum). Material.-From stations P-781, P-984, G-456, G-507, G-589, G-659, G-676, G-820, G-894, G-1015, G-I028, G-I035, from the western Atlan- tic, Straits of Florida, and the Caribbean, between 11" 30' and 27" 25' N and 73" 20' and 86" 19' W, in depths from 68-567 m, in the years 1965- 69, a total of 20 specimens. Further Distribution.-Bermuda; Straits of Florida; Antilles to Barbados; Gulf of Darien; northern Brazil, 03" 08' N, 48" 07' W; 85-724 m. Remarks.-As in our previous material (Marcus, 1967a: 606), the shells of all the present animals were shattered in the animal, so that only their sculpture could be determined (Fig. 95). The characters of the soft parts were compared with the earlier description. There is a certain variability in the radular teeth: the denticulation of the lateral was an irregular ser- ration in the previous specimens from the Gulf of Darien, while the present animals vary between a weak irregular serration and 70 regular denticles. Also the shape of the marginal tooth, especially its tip, shows slight differ- ences (Figs. 93, 94). In one specimen, a foraminifer more than 4 mm in diameter had passed through the gizzard unbroken. Dall (1889: 57) called P. infundibulum a rather common species, rang- ing from the Straits of Florida to Barbados, whence he had obtained nine samples in depths from 216-680 m. The single specimen we described (1967a: 606) was designated P. d. infundibulum, as Dall had not figured his species nor described the radula. The present rich collection from the same area and depths as Dall's original animals justifies the identification with his species. The gizzard plates of Philine berghi E. A. Smith, 1910 (synonyms: P. capensis Bergh, 1907: 27, pI. 5, figs. 11-15; P. berghii O'Donoghue, 1929: 10), are very similar to those of infundibulum, but in berghi the unpaired plate is smaller than the paired ones, and its radular formula is 2.1.0.1.2. The piece of grass sticking to the specimen from 531-567 m depth is not an indicator for shallow water, as fragments of grass are frequent even 356 Bulletin of Marine Science [24(2)

FIGURES 93-97. Philine infundibulum Dall; 93, denticJes of lateral teeth and marginal teeth; 94, lateral teeth; 95, sculpture of shell; 96, gizzard; 97, male copulatory organ. (a, male atrium; d, ejaculatory duct; e, sperm duct; p, penial papilla; q, prostate; v, seminal vesicle.) 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 357 in much greater depths, e.g., 2200-3629 m (Roper & Brundage, 1972: 41, figs. 7, 19,52,23). The distinction between Philine infundibulum and P. mera is not easy. The shape of the shell is variable, a little broader in infundibulum, whose ratio of breadth to length is 75 per cent in the original description, 84 per cent in our previous material, and 93-117 per cent in the present collection, against 66-84 per cent in memo The sculpture, which Dall (1889: 57) described as finely closely spirally striate for infundibulum, was in our previous material (1967a: 606) spiral only near the apex, curving outward and forming growth lines on the body whorl. The present peripheral fragments have only strong growth lines (Fig. 95), or also feeble spiral ones. The shells of P. mera have dense, slightly catenate spiral grooves and loose growth lines (Fig. 105). A distinct difference exists in the shape of the penis; it has two almost equal tips in infundibulum (1967a: fig. 22; and the present Fig. 97), and in mera a single main tip with a rudimentary second point (1969a: fig. 21; and the present Fig. 107). Hence, I maintain the two species separate.

42. Philine pc. falklandica Powell, 1951 Figs. 98-101 References.-Powell, 1951: 178, 195, fig. M, 94, pI. 7, fig. 24.-Rudman, 1972b: 172, 183, figs. la, Ib, 2c, 7a, 8a, 8b, 9b. Material.-Brazil, off Rio Grande do SuI, 33° 17' S, 50° 34' W, 173 m, 10. III. 1971., Oceanographic Institute of the University of Sao Paulo, eight specimens. Further Distribution.-From 47° 37' S to the Falkland Islands; 161-219 m. McMurdo Sound, 13-33 m under the ice shelf. Descriptive Notes.-The length of the animals is 7-17 mm. The shells are nearly all shattered. That of the 7-mm animal is 4.8 mm long and shows fine spiral lines in the older parts; the later parts have only radial growth lines. The radula has 2.1.1.1.2 teeth. The laterals are about 600fL high in the largest snail and bear two or three broad denticles on the medial edge (Fig. 100). The marginals are smooth hooks, 254fL and 196fL high, respectively. There is a row of small rhachidian teeth, 45fL broad and 34fL in anteroposterior direction, so weak that it was at first not noticed. The gizzard plates are of equal size and shape. They are not united by special muscle bands and fall off easily. They consist of brownish hori- zontal layers with a whitish incrustation. They have a curved tip (Fig. 99). The prostate is short and sausage-shaped (q, Fig. 101). It is connected 358 Bulletin of Marine Science [24(2)

101

~I02 FIGURES 98-102.-98-101, Philine pro falklandica Powell: 98, inside view of apex of shell; 99, gizzard plates; 100, radular teeth; 101, male copulatory organ, opened.-102, Philine alba Mattox, gizzard plates. (a, male atrium; 0, opening of penial papilla; p, penial papilla; q, prostate; r, thickened part of atrial wall.) with a small penial papilla (p) opening with a long slit (0). Part of the atrial wall is thickened and glandular (r). RemarkS.-Both P. alba and P. falklandica have similar small gizzard plates of equal size. As P. falklandica has a rhachidian tooth (Rudman, 1972b: 173) and curved tips of the gizzard plates (Rudman, 1972b: fig. 1b), against the straight ones of alba, I approach the present specimens to falklandica. However, the male organ is intermediate between that of alba (Marcus, 1967a: fig. 28) and that of falklandica (Rudman, 1972b: fig. 7a). Geographically, the latter is more probable than the former. 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 359

FIGURE 103. Philine alba Mattox: diagrammatic reconstruction of male copu- latory organ from serial sections. (a, male atrium; 0, U, lobes of penis; p, tip of penis; q, prostate; s, seminal groove; t, tip of prostate.)

43. Philine alba Mattox, 1958 Figs. 102, 103 References.-Mattox, 1958: 98, figs. 1-7.-Marcus, 1967a: 607, figs. 23- 28. 360 Bulletin of Marine Science [24(2) Material.-P-199: 27° 59' N, 79° 20' W, 311-329 01, 11. VIII. 1964.- G-158: 26° 32'-36' N, 79° 21'-24' W, 530-54001, 25. VI. 1963.-G-507: 25° 52'-53' N, 79° 19'-20' W, 366-384 01, 2. III. 1965.-G-798: 25° 26' N, 79° 17'-22' W, 392 01, 12. IX. 1966.-G-887: 24° 00' N, 79° 47' W, 274-325 01, 31. VIII. 1967.-G-820: 23° 05' N, 69° 01' W, 225-120 01, 12. VI. 1967.-P-1354: 14° 21' N, 81° 55' W, 192-263 01, 31. I. 1971. -P-I171: 23° 45' N, 79° 24' W, 51201, 27. VI. 1970.-Eight specimens from the Straits of Florida. Further Distribution.--California; Caribbean, 46-295 m. Descriptive Notes.-The length of the specimens is 19-440101. The largest shell (P-l171) is 23 0101long and 19.5 0101broad. It is rather solid, as was our previous Caribbean shell. Its growth lines are distinct, but spiral ones appear as denser lines in the rather transparent shell. The Hancock's organ is a simple narrow bulge. The labial cuticle is smooth and brown. The radula has 14-16 rows of 2.1.0.1.2 teeth. The gizzard plates (Fig. 102) have straight tips, contrary to those of falklandica. Their size is not correlated with that of the snails. One snail of 22 0101 length had plates 1.4 0101long, and in another of the same length they measured 2.1 0101. The male organ (Fig. 103) corresponds to the previous description (Marcus, 1967a: 608), with a folded atrium and no true penial papilla. It has the characteristic short, sausage-shaped prostate and folded penis (Fig. 103). Remarks.-The shape of the shell of Philinorbis Habe, 1950 (Zilch, 1959- 60: fig. 99) is rather similar to that of Philine alba, but the sculpture consists of many spiral stripes. As the soft parts of Philinorbis teramachii were not described, they cannot be compared. The gizzard plates of the largest present specimen were lost, because its digestive tract was full of fragments of foraminifers; the most frequent ones were brown Astrorhizidae, kindly classified by Walter Narchi.

++ Philine aperta guineensis Marcus, 1966 Fig. 104 References.-Marcus, 1966: 159, figs. 9-18; 1968: 1334. Material.-P-251: 04° 03' N, 06° 03' E, 27 01, 14. V. 1965.-P-63: 04° 35' N, 06° 40' W, 6401, 2. VI. 1964.-P-65: 04° 15' N, 07" 32' W, 46-49 01,2. VI. 1964.-P-17: 05° 35'-36'N, 00° 1O'-l1.5'E, 42 01,26. V. 1964. -P-250: 04° 06'-02' N, 05" 58'-06° 04' E, 24 01, 14. V. 1965. A total of 10 specimens. Further Distribution.-From Ivory Coast to Nigeria, 22-98 m. 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 361

FIGURE 104. Phi/ine aperta guineensis Marcus, male copulatory organ. (a, atrium; d, ejaculatory duct; p, penial papilla.)

Remarks.-The present specimens from the same area as the previous ones are 14-50 mm long. The paired gizzard plates are still broader than in the former specimens; in the wet plates, the ratio of length to breadth is 88 per cent in the 50-mm specimen, in others 73 per cent. The unpaired plate with 5.5 x 2.5 mm = 45.5 per cent ratio is inside the known variation. As the male organ was not figured before, I give a figure (Fig. 104). The hammer-shaped organ is much broader than in Philine aperta aperta.

44. Philine mera Marcus, 1969 Figs. 105-111 Reference.-Marcus, 1969a: 5, figs. 13-21. Material.-Brazil, Rio Grande do SuI, 22° 39' S, 41 ° 33' W, 52 m, 10. III. 1971.-23° 40' S, 45° 40' W, 22 ill, 6. V. 1970, Plinio S. Moreira leg.- 29° 39' S, 48° 41' W, 124 m.-31 ° 19' S, 50° 22' W, 102 m.-31° 33' S, 40° 52' W, 209 m.-33 ° 17' S, 50° 34' W, 173 m. A total of 43 specimens. Further Distribution.-Brazil, off Ubatuba, 42-52 m, in mud. Description.-The largest animals are 10 mm long. Their shells (Figs. 105, 106) measure 7 x 5.5 mm. They are pale whitish, or pinkish with the dark liver shining through. The broad shell is semitransparent whitish; it has a breadth-to-Iength ratio of 66-84 per cent. The smooth outer lip is higher than the round apex. The sculpture consists of wide-spaced growth lines and dense spiral rows of flat pits. These pits are shorter and broader apically than anteriorly. 362 Bulletin of Marine Science [24(2)

105 106

1\0 109

FIGURES 105-111. Philine mera Marcus: 105,106, shell; 107, male copulatory organ; 108, radular teeth; 109, ventral view of preserved animal; 110, dorsal view of same; Ill, gizzard plates. (a, male atrium; d, ejaculatory duct; p, penial papilla.)

The radula (Fig. 108) has 15-20 rows of 1.1.0.1.1 teeth. The big lateral tooth (246,u) bears about 30 long denticles on its inner border. The small marginal tooth (130,u) is smooth and pointed, with a broad base. The three soft, whitish gizzard plates (Fig. 111) have yellow, cuticular layers and a white center; they have no pores. The paired plates are 3.1 mm long and 1.0 mm in breadth. The unpaired plate is 3.1 x 1.72 mm, its anterior end is blunt, the posterior one, narrowed. Several pebbles, up to 1.5 mm in diameter, were stopped in front of the gizzard. Others had passed through. 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 363 The male organ (Fig. 107) is similar to that of P. aperta (L.) in Lloyd's description (1952: 17, fig. 4). The shape of the penial papilla differs by a more slender and single tip, it has only a rudimentary second point. Remarks.-The shell of P. mera is somewhat similar to, but much broader than, that of P. sagra (d'Orbigny, 1841), from Cape Hatteras to the West Indies, of which the radula and gizzard plates are not known. Philine falklandica Powell, 1951 (p. 171, pI. 7, fig. 24) has a rather similar shell, but the radular formula is 2.1.1.1.2, and the gizzard plates are minute.

B. WESTERN ATLANTIC SPECIES OF Philine NOT IN THE PRESENT COLLECTION + 45. Philine aperta aperta (Linne, 1767) References.-G. O. Sars, 1878: 303 (key only), pI. XI, figs. 15a-h.- Pilsbry 1895-96: 10, pI. 3, figs. 47-56.-Lemche, 1929: 6 (distribution); 1948: 90 (synonyms).-O'Donoghue, 1929: 7, figs. 1-7.-Forster, 1934: 1-67, figs. 1-46 (histology).-Brown, 1934: 179-310, figs. 1-38 (anatomy and development) .-Fretter, 1939: 600-608, figs. 1-6 (histology) .-Lloyd, 1952: 8-26, figs. 1-5, 13, 17 (reproductive apparatus).-White, 1955: 168. -Hurst, 1965: 218-325, figs. 1-23 (buccal apparatus). Distribution.---Southern Norway; west coasts of Europe; Mediterranean; West Africa, from Morocco to Cape of Good Hope; Indian Ocean; 0-91 m. Remark.-White (1955: 169) mentioned that the gastric plates of her specimens from Cape Lopez are quite typical for P. aperta, hence for P. aperta aperta, of which O'Donoghue figured the gizzard plates from South Africa. + 46. Philine scabra (0. F. Muller, 1776) References.-G. O. Sars, 1878: 294, pI. 18, figs. 13a-c, pI. XII, figs. 4a-c. -Pilsbry, 1895-96: 12, pI. 5, figs. 1-3.-Lemche, 1948: 66, 91 (syn- onyms).-Marcus, 1966: 158, fig. 8. Distribution.-East Atlantic from Iceland to West Africa and Madeira; 20-216 m, living specimens; shells 2300 m (Lemche).

+ 47. Philine catena (Montagu, 1803) References.-G. O. Sars, 1878: 294, pI. 26, figs. 6a-c, pI. XII, fig. 6.- Vayssiere, 1885: 35-38, figs. 25-34.-Pilsbry, ] 895-96: 13, pI. 5, figs. 23-25.-Lemche, 1948: 67, 92 (synonyms) .-Pruvot-Fol, ]954: 66, figs. 15a-h. Distribution.-Mediterranean; boreal East Atlantic to Madeira; 1115..2019 m. 364 Bulletin of Marine Science [24(2) + 48. Philine lima (Brown, 1825) References.-G. O. Sars, 1878: 300, pI. 18, figs. 12a-f, pI. XII, fig. 8.- Lemche, 1948: 68, 93-94 (synonyms).-Pruvot-Fol, 1954: 72 (?Laona lima).-Marcus, 1969b: 12, fig. 17. Distribution.-From Greenland to Massachusetts and Iceland, Northern Norway and the Kara Sea, 4-800 m. + 49. Philine quadrata (S. Wood, 1839) References.-G. O. Sars, 1878: 299, pI. 18, figs. 9a-9d, pI. XII, figs. 7a-7d. -Pilsbry, 1895-96: 19, pI. 5, figs. 18, 19.-Vayssiere, 1913: 174.- Lemche, 1948: 68,95 (synonyms).-Pruvot-Fol, 1954: 72 (Laona qua- drata).-Marcus, 1969b: 12, figs. 18, 19.-Rudman, 1972a: 172, 177, figs. Ie, 7c, lOd. Distribution.-Eastern Atlantic from Kola Peninsula to the Mediterranean, Azores, and St. Helena; West Atlantic to North Carolina; 5-2150 m. + 50. Philine sinuata (Stimpson, 1850) Reference.-Franz & Clark, 1969: 69-71, figs. 1-8. Distribution.-New England, 3-13 m. + 51. Philine trachyostraca Watson, 1897 References.-Watson, 1897: 236, pI. 19, figs. 4, 4a.-Marcus, 1969a: 3, figs. 5-12. Distribution.-Madeira; Brazil, 20-91 m.

+ 52. Philine (d.) gibba Strebel, 1908 References.-Odhner, 1926: 17-18, figs. 12, B.-Marcus, 1969b: 13, figs. 20-22.-Rudman, 1972b: 174,175,177, figs. 2a, 2b, 7d, 7e, lOb. Distribution.-Southern West Atlantic, 45° S, McMurdo Sound, and South Georgia, 12-310 m. + 53. Philine thurmanni thurmanni Marcus, 1969 Reference.-Marcus, 1969b: 14, figs. 23-28.

Distribution.-Southern West Atlantic, off Argentina; 340 29' to 4r 02' S, 70-4116 m. + 54. Philine amabilis Verrill, 1880 Reference.-Verrill, 1880: 398. Distribution.-Western Atlantic, New England.

C. LIST OF WESTERN AND MIDDLE ATLANTIC SPECIES OF Philine KNOWN ONLY FOR THEIR SHELLS P. sagra d'Orbigny, 1841, West Indies and Brazil. P. candeana d'Orbigny, 1853, West Indies. 1974] Marcus: Cephalaspidea from Atlantic Warm Waters 365 P. planata DaH, 1889, West Indies. P. approximans Dautzenberg & Fischer, 1896, Azores. P. rugosula Dautzenberg & Fischer, 1896, Azores. P. miLne-edwardsi Locard, 1897, Azores. P. complanata Watson, 1897, Madeira. P. desmotis Watson, 1897, Madeira. P. Lucida DaH, 1927, West Indies.

RESUMO o trabalho representa 72 especies pertencentes a Familia Acteonidae e aos generos Scaphander e Philine, conhecidos do Atlantico Ocidental, de Cape Cod as Ilhas Malvinas, e, para leste, ate Azores e Camirias, que sao enumeradas, sendo as 25 das presentes cole~5es descritas e ilustradas. Somente de tres, Acteon candens, Scaphander nobilis e S. (Sabatina) bathymophilus, as partes moles for am descritas pela primeira vez; algumas espccies sao redescritas. Das especies conhecidas, 48 sao somente con- hecidas pelas conchas vazias, 9 delas ainda nao tern figuras, assim que a posi~ao generica das 48 permanece incerta. As partes moles conhecem-se agora de 26 especies dos grupos tratados. Acteon candens, Scaphander nobilis e Philine pr. faLkLandica sao novos para 0 Brasil. Acteon cumingii A. Adams, 1854, e a especie tipo do novo genero MysoufJa.

ZUSAMMENFASSUNG Die 72 Arten der Acteonidae und der Gattungen Scaphander und PhiLine, die vom westlichen und mittleren Atlantik von Kap Cod bis zu den Falk- land-Inseln, ostwarts bis zu den Azoren und Kanaren, bekannt sind, werden aufgezahlt, und die 25 Arten des vorliegenden Materials werden beschrieben und abgebildet. Nur fUr 3 Arten: Acteon candens, Scaphander nobiLis und S. (Sabatina) bathymophiLus, konnten die Weichteile erstmalig beschrieben werden. Mehrere Arten wurden wieder beschrieben. Von 48 Arten sind nur die leeren Sehalen besehrieben, 9 nieht einmal abgebildet, so dass ihre Gattunszugehorigkeit unsicher bleibt. Die Weichteile sind jetzt fUr 26 Arten der behandelten Gruppen bekannt. Acteon candens, Scaphander nobiLis und Philine pr. faLkLandica sind neu fUr Brasilien. Acteon cumingii A. Adams, 1854, ist der Typus der neuen Gattung MysoufJa.

LITERATURE CITED ABBOTT, ROBERT TUCKER 1955. American Seashells. (3rd printing.) D. Van Nostrand, New York, xiv + 541 pp., 40 pIs. AGUAYO, C. G. AND HARALD A. REHDER 1935. New marine mollusks from Cuba. Mems Soc. cub. Hist. nat. 'Felipe Poey,' 9: 263-268, pI. 24. 366 Bulletin of Marine Science [24(2)

BARNARD, KEPPEL HARCOURT 1963. Contributions to the knowledge of South African marine Mollusca, Pt. 4, Tectibranchiata and others. Ann. S. Air. Mus., 47(2): 201-360. BERGH, RUDOLPH 1901. Malacologische Untersuchungen, 5, 4. Abt., 3. Abschnitt, Bullacea, 1 & 2. In Semper, C., Reisen im Archipel der Philippinen. Zweiter Teil, Wissenschaftliche Resultate, 7: 209-312, pIs. 17-24. 1902. Malacologische Untersuchungen, 5,4. Abt., 4. Abschnitt, Ascoglossa (err. pro Acteonidae), Aplysiidae. In Semper, C., op. cit., 7; 313-382, pIs. 25-29. 1907. The Opisthobranchiata of South Africa. Trans. S. Afr. phil. Soc., 17 (1): 1-144, pis. 1-14. BROWN, HERBERT 1934. A study of a tectibranch gasteropod mollusc, Philine aperta (L.). Trans. R. Soc. Edinb., 58: 179-210. BULLIS, HARVEY, JR. 1956. The genus Scaphander in the Gulf of Mexico and notes on the western Atlantic species. Bull. Mar. Sci. Gulf Caribb., 6: 1-17, 5 figs. CLARKE, ARTHUR HADDLETON, JR. 1962. 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