ON THE (: ) OF THE WARM WATERS OF THE WESTERN ATLANTIC

EVELINE D. B.-R. MARCUS Caixa Postal 6994, 01000 sao Paulo, Brazil

ABSTRACT A key is given for the 16 western Atlantic of Anaspidea and two Middle Atlantic ones. These are eight (and two) species of , and punctatus armatus, longicauda, Stylocheilus citrinus, leachii pleii, dolabrifera, petalifera bon- airei, Petalifera ramosa, and engeli. Descriptions are given for Petalifera p. bonairei and Petalifera ramosa. A key to the subgenera of the Aplysia is also furnished. Comparative notes are made on anatomical and systematic characters, sole, pallial cavity, shell, and nervous system. My subdivision of the Notarchidae leaves only Notarchus in the Notar- chinae; the remainder of the genera are allotted to the Dolabriferinae. Three species of Aplysia were dredged in 348-357 m depth.

INTRODUCTION The present paper deals with seven western Atlantic genera and 16 species of the order Anaspidea. Taylor & Sohl (1962: 12) considered the order Anaspidea as consisting of the superfamily Aplysiacea, including and . However, the Akeridae are often allotted to the Cephalaspidea (Hoffmann, 1933: 308-309; Wirz, 1952: 167; Pruvot-Fol, 1954: 92; Marcus, 1967b: 613; 1970c: 939; and others). The strepto- neurous (Guiart, 1901: 116; Marcus, 1970c: 938, fig. 32) really occupies an intermediate position between Cephalaspidea with cephalic shield and pallial caecum and the euthyneurous Anaspidea with numerous gizzard teeth. However, the argument that Akera has a head shield like the Cephalaspidea, while the Anaspidea have tentacles and , favors the placing of Akera in the Cephalaspidea in any key. The Anaspidea can be separated by the length of the visceral loop of the nervous system into the suborder Longicommissurata (Pruvot-Fol, 1954: 95) with the family Aplysiidae, including the Aplysiinae and Dolabel- linae; and the Brevicommissurata (1954: 101) with the family Notarchidae, divided into Notarchinae and Dolabriferinae. In all six genera of the Brevicommissurata, the body may be beset with knobs or warts or branched papillae, but these protuberances can be so completely contracted that the appear quite smooth. The genus Notarchus, with a pallial cavity all around the visceral sac and with a quite narrow foot, differs widely from the rest of the Brevicommissurata, in which the pallial cavity does not extend under the visceral sac and which have a broader foot. 842 Bulletin of Marine Science [22(4)

Martin (1966: 72) distinguished only three subfamilies of the Aplysiidae: Aplysiinae, Dolabellinae, and Dolabriferinae, and alloted Notarchus, of which he gave beautiful photographs, to the Dolabriferinae.

ACKNOWLEDGMENTS Prof. Dr. Frederick M. Bayer, Rosenstiel School of Marine and Atmo- spheric Sciences of the University of Miami, entrusted me with hundreds of specimens of opisthobranchs collected by himself and other members of the school and on cruises of the R/V JOHN ELLIOTT PILLSBURY and R/V GERDA. Drs. Marc Kempf, Recife, and Pierre Montouchet, Campinas, gave me their opisthobranchs from the Brazilian coast. From Lt. Col. Corinne E. Edwards, Miss Kaoru Hiroki, Mrs. Helen Hughes, Dr. John Lewis, Dr. Donald Moore, Messrs. Edgardo Ortiz Corp's, Bruce Ott, Robert Austin Smith, Robert C. Work and Warren Zeiller I received specimens. To all of them I am grateful for the interesting material and much help. Mr. Robert Austin Smith kindly revised my manuscript and the bibliography. Dr. Eurico de Oliveira classified the algal contents of the stomachs of three aplysiids dredged in 348-357 m depth. The "FundaC;ao de Amparo a Pesquisa do Estado de Sao Paulo" (FAPESP) granted me the trip to Recife, Curac;ao, Puerto Rico, and Miami for collecting and comparing material (Proc. 71/961), for which I express my sincere thanks. Dr. and Mrs. Ingvar Kristensen invited me to their house during my stay at Curac;ao, and I am grateful for the very pleasant time at their home.

LIST OF SPECIES Order ANASPIDEA Suborder LONGICOMMISSURATA Family Aplysiidae Subfamily Aplysiinae Genus Aplysia Linne, 1767 1. Aplysia (Pruvotaplysia) parvula March, 1863 2. Aplysia (Varria) brasiliana Rang, 1828 3. Aplysia (Varria) cervina Dall & Stimpson, 1901 4. Aplysia (Varria) dactylomela Rang, 1828 5. Aplysia (Varria) donca Marcus, 1960 6. A plysia (V arria) morio Verrill, 1901 7. Aplysia (Aplysia) juliana Quoy & Gaimard, 1832 Aplysia (Varria) willcoxi Heilprin, 1886-of this species, I had no specimens. Suborder BREVICOMMISSURATA Family Notarchidae Subfamily Notarchinae Genus N otarch us Cuvier, 1817 8. Notarchus punctatus armatus Baba, 1938 1972] Marcus: Anaspidea of the Western Atlantic 843 Subfamily Dolabriferinae Genus Stylocheilus Gould, 1852 9. Stylocheilus longicauda Quoy & Gaimard, 1824 10. Stylocheilus citrin us (Rang, 1828) Genus Bursatella Blainville, 1817 11. pleii (Rang, 1828) Genus Dolabrifera Gray, 1847 12. Dolabrifera dolabrifera (Cuvier, 1817) Genus Petalifera Gray, 1847 13. Petalifera petalifera bonairei Engel, 1936 14. Petalifera ramosa Baba, 1959 Genus Phyllaplysia P. Fischer, 1872 15. Phyllaplysia engeli Marcus, 1955 Of the well-described worldwide species, I do not give new descriptions nor all the localities.

KEY TO THE ANASPIDEA Figs. 1-8 1. Pleurovisceral nerve cords long, visceral ganglia far behind pedal ganglia . Longicommissurata, Aplysiidae, 2 Pleurovisceral nerve cords quite short, visceral ganglia over pedal ganglia . . Brevicommissurata, Notarchidae, 3 2. Animals with dorsocaudal, oblique disc ______Dolabellinae (, Indopacific, Fig. I) No dorsocaudal disc Aplysiinae, Aplysia (Syphonota, Indopacific) 3. Dorsal slit in anterior half of body, sole quite narrow, pallial cavity surrounds entire visceral mass Notarchinae, Notarchus Dorsal slit in middle or posterior half, pallial cavity dorsal and lateral to visceral mass . Dolabriferinae, 4 4. Foot narrower than greatest diameter of body, no shell .___ 5 Underside set off by a ridge along widest part of body 7 5. A row of small papillae around the broad sole . B ursatella No such papillae, long and slender tail Stylocheilus, 6 6. Color yellow to orange with white spots . S. citrinus Color greyish with dark stripes and eyespots S. longicauda 7. Normal, primary shell present . . 8 Shell absent or a small, secondary disc . Phyllaplysia 8. Shell foramen closed _.. . . . Dolabrifera Shell foramen wide . . . Petalifera, 9 9. Only two denticles on the outer side of the lateral teeth of the ______. .. . ._. Petalifer a pe tali fera bo nair ei More than two dentic1es Petalifera ramosa 844 Bulletin of Marine Science [22(4 )

KEY TO THE SUBGENERA OF Aplysia 1. Parapodia joined high up posteriorly 2 Parapodia joined low down posteriorly _ 3 2. Sole narrow, no sucker Aplysia (Pruvotaplysia) Engel, 1936) 1. A. parvula Sole broad, its hind end producing a temporary sucker _ Aplysia (Aplysia) Linne, 1767 7. A. juliana 3. Long and slender tail, shell with platelike, flattened apex ______Aplysia (Neaplysia) Cooper, 1863 (Pacific) No long tail, apex a reduced spiral A plysia (V arria) Eales, 1960

KEY TO WESTERN AND MIDDLE ATLANTIC SPECIES OF Aplysia (Varria) 1. Black rings on outer side of para podia 4. dactylomela No black rings 2 2. aperture closed 6. morio Man tie aperture open 3 3. Mantle aperture a flat hole 4 Mantle aperture tubular, on tip of papilla 5 4. Opaline gland multiporous 5. donca Opaline gland uniporous . fasciata Poiret, 1789* 5. Scattered, round, dark spots, diameter 2-3 mm, on parapodia, caecum curved 3. cervina No round spots, caecum straight .______6 6. Only tip of caecum visible on surface of digestive gland . ______wilcoxi, Heilprin, 1886t Caecum straight on surface of digestive gland 7 7. Para podia with smooth edges 2. brasiliana Parapodia with fimbriated edges . winneba Eales, 1960*

1. Aplysia (Pruvotaplysia) parvula Morch, 1863 References.-Eales, 1960a: 287, figs. 10, 11 (synonyms and bibliography); Marcus, 1967a: 154; 1970a: 26; Barash & Danin, 1971: 167. Material.-Brazil: Bahia, Olivem;a, 17. I. 1969; Bahia, Salvador, Ondina, 18. X. 1968; Espirito Santo, Guarapari, 6. XII. 1968, a total of 25 speci- mens in the lower tidal zone, Pierre Montouchet leg.

Further Distribution.-From 400N to 40·S in all warm and temperate seas. Remark.- The Israeli locality of Barash & Danin (1971) seems to be the first indication for the Mediterranean.

• For the two Middle Atlantic species of Aplysia. see Eales (1960a: 315, 352). t As I had no specimens of Aplysia willcoxi, it is not discussed in this paper; for a description, see Eales (1960; 350). 1972] Marcus: Anaspidea of the Western Atlantic 845

1

8

FIGURES 1-8. Aspect of living animals: 1, Dolabella; 2, Aplysia; 3, Notarchus punctatus armatus (from Baba, 1938); 4, Stylocheilus longicauda; 5, Bursatella leachii pleii; 6, Dolabrifera dolabrifera; 7, Petalifera ramasa (from Baba, 1959); 8, Phyllaplysia engeli. (Shell stippled.)

2. Aplysia (Varria) brasiliana Rang, 1828 Fig. 9 References.-Marcus, 1957b: 4, figs. 1-23; 1959: 8, figs. 11, 15-18; Eales, 1960a: 297, figs. 14, 15 (bibliography). Material.-PILLSBURY Sta. 580: 21°0S'N, 86°23'W, 348-357 m, one speci- men, 23 mm long, preserved, 22. V. 1967, together with A. donca and A. moria. 846 Bulletin of Marine Science [22(4 ) Further Distribution.-From New Jersey to Brazil, Sta. Catarina; West Africa, Ghana; St. Helena. Remarks.- The occurrence of Aplysia species, algal feeders, in depths beyond 200 m is exceptional. However, the three dredged species, A. brasiliana, A. danca, and A. moria, had their digestive tracts filled with freshly ingested algae: Dictyota, Acrochaetium, Polysiphonia, Hypoglos- sum, and Dasya, kindly classified by Dr. Eurico C. de Oliveira, Departa- mento de Botilnica do Instituto de Biociencias da Universidade de Sao Paulo. Prof. Frederick M. Bayer recalls the trawls coming up from 150- 200 fathoms filled with bushels of benthic algae-very big and healthy (letter of March 1, 1972). Eales (1960a: 377) stated that the feeding habits of Aplysia confine the genus to the littoral and sublittoral zones of the land masses and islands of the world. The larvae generally settle in the zone of red algae, and the growing animals migrate shorewards; the mature animals gather between tide marks for spawning. As they are mostly caught on the beach from 0-2 m, an indication of the depth is in most cases omitted. There are a few data, e.g., Vayssiere (1913: 201-203) gave 20-30 m for andA.jasciata,andWhite (1955: 176,177) mentioned 28 mforthesame two species. Bergh (1905: 12) had A. velifer (a synonym for A. dacty- lomela) from 40 m; Aplysia moria was reported from 42.4 m (Marcus, 1967c: 200). M. C. Miller (1960: 165) found spawning in 18-29 m in Manx waters, and Carefoot (1967: 335, ff.) studied a sub- littoral population of the same species living and spawning in about 13 m depth, not migrating shoreward. However, specimens of the intertidal populations were larger, as explained by the greater abundance of their preferred algae. We observed juvenile, recently metamorphosed speci- mens of Aplysia brasiliana (1957b: 10), from 0.9 to 8 mm long when alive, among algae in the eulittoral of the Island of Sao Sebastiao, Brazil, as well as slugs of all sizes up to 27 em in the same habitat.

3. Aplysia (Varria) cervina (Dall & Stimpson, 1901) Fig. 68 References.-MacFarland, 1909: 38, figs. 15-35, 39-42 (Tethys cervina); Marcus, 1957b: 3, figs. 1-10; 1964: 198; Eales, 1960a: 299, figs. 15, 16. Material.-Brazil: Pernambuco, Ponta da Pedra, Pierre Montouchet leg., 30. VIII. 1970, one specimen; Praia da Piedade, 23. XII. 1971, 5 speci- mens at low-water level (E. M.). Further Distribution.-Brazil, Sao Paulo, Alagoas; West Indies; Central America; southern United States. 1972] Marcus: Anaspidea of the Western Atlantic 847

FIGURES 9-14. Radula: 9, Aplysia brasiliana; 10, Aplysia donca; 11, ; 12, Stylocheilus longicauda; 13, Bursatella leachii pleii; 14, Notarchus punctatus armatus.

4. Aplysia (Varria) dactylomela Rang, 1828 Rejerences.-Marcus, 1957b: 14; 1959: 8, figs. 12, 13; 1970a: 26; Eales, 1960a: 307, frontispiece and figs. 1-8 (synonymy and bibliography). Material.-Brazil: Pernambuco, littoral, Marc Kempf and E. M. leg. XII. 1971 and I. 1972. Further Distribution.-Circumtropical, 0-40 m (A. velifer, Bergh 1905: 12). 5. Aplysia (Varria) donca Marcus, 1960 Fig. 10 Rejerence.-Marcus, 1960: 251, figs. 1-5. 848 Bulletin of Marine Science [22(4) Material.-PILLSBURY Sta. 580: 21°05'N, 86°23'W, 348-357 m, one speci- men, 56 mm long, preserved, 22. V. 1967, together with A. brasiliana and A. moria. PILLSBURYSta. 583: 21°10'N, 86°18'W, 27 m, one specimen, 23. V. 1967. GERDASta. 1037: 24°50'N, 80037.5'W, 46 m, one specimen, 26. II. 1969. Further Distribution.- Texas, in tide pool. Remark.-For the occurrence in great depth, see Aplys,ia brasiliana.

6. Aplysia (Varria) moria Verrill, 1901 Figs. 11, 52 References.-Eales, 1960a: 328, figs. 28, 29; Marcus, 1967c: 200. Material.-PILLSBURY Sta. 580: 21°05'N, 86°23'W, 348-357 m, one speci- men, 90 mm, 22. V. 1967, together with A. brasiliana and A. dactylomela. Further Distribution.-From Rhode Island to Texas; Bermuda; down to 42.4 m. Remarks.-The radulae of Aplysia are almost unspecific (Figs. 9-11), so they are of little use for classification, especially if the animals are of dif- ferent sizes, as are the present three. In that of A. moria (Fig. 11) the lateral teeth from the 11th to the 14th on one side have abnormal, irregular cusps. The mantle foramen of full-grown animals is a good character, but in quite young specimens it is always wide open. In the present large slug, it is completely closed as in the original moria; none of the other western Atlantic species of Aplysia shows this character. The three specimens of Aplysia from P-580 belong to three different species. The two accompanying spawns are also different. The diameter of the string is 2 mm in the bigger spawn, which possibly comes from the largest . The smaller spawn measures 1 mm in diameter. Both these spawns might be from specimens and species not captured. The spawns of full-grown individuals of A. brasiliana have a diameter of 2 mm.

7. Aplysia (Aplysia) juliana Quoy & Gaimard, 1832 Figs. 37, 44 References.-Marcus, 1957b: 15, figs. 24-37; 1958a: 10, figs. 15-18; 1958b: 42, figs. 33, 34; 1959: 8, fig. 14; 1967a: 155; 1970a: 26; Eales, 1960a: 364, figs. 44-46 (synonymy and bibliography). Material.-Venezuela, Los Roques, in tide pool, 4. VIII. 1963, one speci- men, Robert C. Work leg. Further Distribution.-Worldwide in warm seas. 1972] Marcus: Anaspidea of the Western Atlantic 849 8. Notarchus punctatus armatus Baba, 1938 Figs. 3, 14, 31, 38, 45, 53, 61, 75 References.-Baba, 1938: 4, figs. 2A-H (Notarchus indicus armatus); 1949: 28, 127, text figs. 12A, 13, pI. 5, fig. 18; Marcus, 1962a: 457, figs. 4-6 (Notarchus punctatus armatus); Abe, 1964: 24, pI. 5, fig. 15 (Notar- chus punctatus armatus). Material.-GERDA Sta. 1089: 24°N, 82°W, 60 m, 27. IV. 1969, four speci- mens, 8.5 to 13.5 mm long. Further Distribution.-Middle Japan, 8-10 m; Florida, Islamorada, 73 m. Remarks.- The present animals are smaller than our previously described ones (1962a: 458: 32 mm). They have no shell. The anterior border of the foot is bilabiate (Fig. 31): between the lips the superior foot glands open into a furrow. As we described previously (1962a), the purple and opaline glands are present (Fig. 38). Corresponding to the character of the subfamily, the opaline gland is simple and multiporous. What Hoffmann figured after MazzareIli (1934: 470, 472, fig. 346 L), is not Bohadsch's gland but the clustered gland (Marcus, 1962a: fig. 6,b), now called prostatic (Thomp- son & Bebbington, 1969: 349, fig. 3A). The penial papilla and the collar are covered with high, round tubercles without spines (Fig. 61), while in the larger specimens they were beset with spines without warts. The present specimens were probably in the developmental stage when captured. VayssU~re(1885: 77-99, figs. 86-89) described and figured the male organ of Notarchus punctatus Philippi, 1836, with the penial papilla beset with spines, similar to our previous material.

9. Stylocheilus longicauda (Quoy & Gaimard, 1824) Figs. 4, 12, 32, 46, 54, 62 References.-Marcus, 1963: 11, figs. 10-21 (bibliography); 1967a: 41-42 (discussed), 159, figs. 16, 17; 1970a: 27; MacFarland, 1966: 27, pI. 6, fig. 9, pI. 8, figs. 16-25, pI. 9, figs. 8-12 (Aclesia rickettsi); Farmer, 1967: 341; Bertsch, 1970: 2. Material.-Brazil: from 02°20'S to Recife, 29 animals, Marc Kempf leg.: Espirito Santo, Guarapari, Pierre Montouchet leg., 6. XII. 1968, one speci- men with eye spots, 9 mm long; Bahia, Itaparica, Mar Grande, Pierre Montouchet leg., 22. 1. 1969, one specimen with black streaks. Cura\;ao: intertidal, among algae, I. 1972, the smallest specimen was 2.5 mm, pre- served, and had three peacock-blue eye spots (E.M.). Florida: Miami, 19. XI. 1970, one specimen, 10 cm long, in the Seaquarium, Warren Zeiller leg. Further Distribution.-Circumtropical, from 0-35 m. 850 Bulletin of Marine Science [22(4 )

FIGURES 15-16. Transverse section of sole: 15, Stylocheilus citrin us; 16, Peta- litera ramosa. (r, lateral ridge; s, sole.)

Remarks.-MacFarland (1966) described the parapodial cavity as much smaller than in Notarchus, smaller than in Aplysia, but considerably larger than in Phyllaplysia (taylori). He also described and figured the repro- ductive organs (pI. 9, figs. 8, 9). 10. Stylocheilus citrinus (Rang, 1828) Figs. 15, 39, 55, 69 References.-Marcus, 1962b: 16, figs. 1-6; 1967a: 40. Material.-Brazil: Ceara, Fortaleza, Farol, Pierre Montouchet leg., 21. III. 1969, one specimen, 7.5 mm, with light orange papillae and white glands. Further Distribution.-North and South Atlantic, East Indies, Pacific, generally on floating algae and driftwood, in equatorial waters. Remarks.-The reduced mantle shelf contains the purple glands in front, 1972] Marcus: Anaspidea of the Western Atlantic 851

17

19

FIGURES 17-19.-17, Part of egg string of Phyllaplysia engeli.-18-19, Bursatella leachii pIe;;: 18, section of purple gland; 19, dorsal view of opened pallial cavity. Ca, genital aperture; b, opaline gland; e, seminal groove; g, gill; h, heart; i, rectum; m, mantle shelf; p, purple gland; y, hypobranchial gland.) opening on the under side of its bulge (Fig. 39), and at its hind end the hypobranchial gland, which opens onto the under side.

11. Bursatella leachii pleii (Rang, 1828) Figs. 5, 13, 18, 19, 33, 40, 47, 56, 63, 73 References.-Eales & Engel, 1935: 292,301; Marcus, 1955: 101,103, figs. 20-28; 1963: 15. Material.-Florida: Key Largo, D. R. Moore leg., 11. VII. 1966, one speci- men. PILLSBURY Sta. 767: 12°16-17'N, n003.3'W, 26 m, 28. VII. 1968, one specimen and photograph. Brazil: Pernambuco, Carne de Vaca, 1. 1. 1972, many specimens of all sizes and colors (E.M.). Further Distribution.-West coast of Florida to Brazil, Rio Grande do SuI. Remarks.- Though Bursatella has no shell, it has a distinct mantle shelf (Fig. 19), in which two dense groups of Blochmann's glands are noticeable. The one in front is the purple gland (p) opening to the dorsal side (Fig. 18). The other (y) between gill and anal papilla opens ventrally and corresponds to the definition of the hypobranchial gland (Franc, 1968: 640). Bebbington described the subspecies guineensis of Bursatella leachii (1969: 323-333) and distinguished it by its less woolly appearance and by its colors. The variability of villi and colors of specimens of Bursatella 852 Bulletin of Marine Science [22(4) from one and the same locality was often noted (e.g., Eales & Engel, 1935: 301); these are not safe distinguishing characters. Bebbington's figure II,b of the male organ, however, is far different from those of B. leachii (Fig. 63, present paper; Eales & Engel, 1935: figs. 9, 10; Marcus, 1955: fig. 28; and others) . Also, the central nervous system of the subspecies guineensis differs from that of Bursatella leachii pleii (Hoffmann, 1936: fig. 512 A, after Bergh, 1902a; Fig. 73, present paper). The pedal commissure is much longer in B. leachii guineensis than in pleii. For Bursatella (Barnardaclesia) cirrhifera, Macnae (1962: 198) indi- cated the pallial cavity as almost entirely dorsal, with only small extensions down the sides. From Recife, Marc Kempf reported a great concentration of Bursatella copulating and spawning in October 1968, on the Praia Carne de Vaca. In January 1972, I saw great numbers on the same beach at low tide, neither very large nor very small ones, very different in color, light grey with reddish brown villi, dark brown with black villi, and many other combinations, lying flat in concavities in the sand. The mating pairs were often of different colors and sizes. Unfortunately Henry's misinterpretation of the anal spout as an os- phradium (1952: 10, pI. 1, fig. A) has crept into Hyman's manual (1967: 410, fig. 165 F, 24). 12. Dolabrifera dolabrifera (Rang, 1828) Figs. 6, 20, 24, 34, 43, 48, 57, 66, 74 References.-Engel, 1936a: 26, fig. 16; Eales, 1944: 7, figs. 9, 10; Kay, 1964: 184, pI. 8, fig. 5; Marcus, 1962a: 452; 1970a: 26; 1970b: 190; Bertsch, 1970: 11 (range extension) ; Sphon & Mulliner, (in press) . Material.-Barbados: Bruce Ott leg., 1971, four specimens. PILLSBURY Sta. 568: R. Panama, Punta Paitilla, 13. V. 1967, shore collecting, 7 speci- mens and 3 kodachromes. Curac;ao: I. 1972, tidal zone, 8 specimens (E.M.). Further Distribution.--Circumtropical and circumsubtropical; recently re- corded from the Gulf of California (Bertsch, 1970) and the Galapagos (Sphon & MuIIiner, in press). Remarks.- The kodachromes show light green animals with brown stipples. The under side is also light green, with big round white patches and small brown dots between. The shape and the size of the oral lobes depends upon the degree of pro- jection or contraction (Fig. 24). As Engel showed in almost a hundred measurements and drawings of shells of Dolabrifera, the proportion of breadth to length varies from 40 to 1972] Marcus: Anaspidea of the Western Atlantic 853

23 FIGURES 20-23. Radula: 20, Dolabrifera dolabrifera; 21, Petalifera ramosa; 22, Petalifera petalifera bonairei; 23, Phyllaplysia engeli.

60 per cent (1936a: 33-36). The straight left border of the shell is char- acteristic of Dolabrifera. The number of radular teeth varies widely. Kay (1964) counted 32 X 50.1.50 in a preserved 38-mm-long animal. Engel (1936a: 38), in speci- mens about 70 mm long, found 160 teeth per half-row. In our description of a small animal (1962a: 453), we mentioned a few spine-bearing warts on the collar of the male organ, but the penial papilla itself was without spines. In full-grown specimens, both collar and papilla are strongly armed (Eales, 1944: 8, fig. 10; Kay, 1964). A specimen from Recife (Fig. 66) has a slender papilla with a few small spines near the tip and near the base, and more than one hundred spine- bearing warts in the sheath. Five slugs from Curac;ao, 40-50 mm long preserved, had no penial papilla, and the sheath had varying numbers of warts with or without spines. In several other lots at my disposal, the spines of the male copulatory organ are rather different. In an animal from Madagascar (1970b: 191) there is a stout penial papilla with a single row of spines and a few more near the tip. The sheath is armed with two rows of about eight spines each, on conical warts. 13. Petalifera petalifera bonairei Engel, 1936 Figs. 22, 25-30, 49, 58, 65, 72 Reference.-Engel, 1936a: 43, figs. 17-23 (synonyms and distribution of Petalifera petalifera). 854 Bulletin of Marine Science [22(4)

FIGURE 24. Dolabrifera dolabrifera, anterior end of preserved animals with different degrees of contraction. (0, oral lobe; r, lateral ridge; s, sole; t, tentacle).

Material.-Puerto Rico: Vega Baja, Puerto Novo Beach, 30. V. 1971, Alida and Edgardo Ortiz Corp's leg., on algae on rocks, 2-3 m, 6 specimens, orange with black markings. Barbados: Helen Hughes, leg., 2 specimens. Brazil: Bahia Salvador, Rio Vermelho, Pierre Montouchet leg., 20. I. 1969, 3 specimens. Further Distribution.-Bonaire (Engel, 1936a). Description.- The Puerto Rican specimens are from 3 to 18 mm long, and from 2 to 11 mm broad. Those from Barbados and Bahia are intermediate between these sizes. The specimens in all three collections are colorless with black markings (Figs. 25, 27, 28); the largest animal has grey longi- tudinal patches on the sale. Rhinophores and tentacles have black bases; the seminal groove is bordered with black, and so is the overlapping right parapodial rim. The black spots on the outer side of the parapodia (Fig. 28) correspond to the papillae, which are contracted in most animals. In the 18-mm slug there are further radial black stripes towards the borders of the foot. The head is not set off from the notum. The rhinophores (Fig. 30) are smaller than the tentacles, which lie over the well-developed trans- verse or oblique oral lobes. The tentacles are rolled in along their entire length; the rhinophores have a blunt, rolled tip. The anterior border of the foot is bilobed. The foot is set off against the parapodia by a sharp ridge 1972] Marcus: Anaspidea of the Western Atlantic 855

27

~ f! 30 '1'\ FIGURES25-30. Petalifera petalifera bonairei: 25, preserved animal from Bahia; 26, underside of head of another specimen; 27, 6-mm-long preserved animal from Puerto Rico; 28, melanophores; 29, ventral side of shell and en- larged protoconch; 30, head of third preserved specimen from Bahia. along the widest part of the body. Its concave sides are unciliated, while the convex middle part, occupying about half the breadth of the body, bears cilia. The dorsal slit lies in the hind part of the body. In a 14-mm animal it extends from the sixth to the eleventh mm. The mantle foramen is almost as wide as the border of the shell. The transparent shell of the 18-mm specimen is 7 mm long, 6 mm wide (Fig. 29). Its apex with the protoconch is bent downwards. The mantle border covers only a quite narrow strip of the shell. The genital opening lies inside the mantle cavity, and the black seminal groove runs forward to the right . The osphradium is situated in front, under the mantle shelf. In the bottom of the pallial cavity lie the simple opaline gland cells, extending to its hind end, distinctly to the left of the furrow that separates the parapodium from the bottom of the cavity. The underside of the mantle shelf contains the purple glands, a number of Blochmann's glands. The jaw plates are composed of about 20-30 irregular rows of 50-100 blunt rodlets bearing up to six rather big spines on their tops (Fig. 49). In the 18-mm animal, the radula measures 1.5 by ].5 mm, and consists of 37 rows with 36.1.37 teeth. The rhachidian tooth (Fig. 22) has two denticles on either side of the median cusp. The laterals have two cusps, the inner 856 Bulletin of Marine Science [22(4 ) ones, two outer denticles. Farther outward they have a single outer denticle, and from the 30th on there are only the two main cusps, except for the outermost long and single-pointed teeth. In one of the studied specimens the first lateral of one side was misshapen: in nine rows it had a single cusp, and in the following nine younger rows, many outer denticles, up to eight. In another radula, the 20th tooth of one side had five inner denticles and two outer ones. The narrow oesophagus ends with a wide crop and enters the muscular gizzard. This is armed with several smaller and seven large conical teeth. The second gizzard has several rows of small spines. The male organ (Fig. 65) is strongly contracted. The penial sheath bears a number of spiny warts, while the papilla itself appears to be un- armed, as Engel (1936a: 48) described and figured it for several specimens of Petalifera petalifera from the Mediterranean. There is no penial collar. Kay (1964: 186) found the penis of P. petalifera "apparently armed only with warts." Remarks.-Engel (1936a: 45) described the mandibular elements un- treated and treated with Eau de Javelle, so that the differences in the aspect of their tips, simple or provided with small or large denticles, cannot be used as a distinguishing character for species or subspecies. Eales figured the male organ of P. petalifera from the Mediterranean (1960b: 12, fig. 3a) fully extended. Evidently the sheath is completely everted, so that the spines come to lie on the base of the penial papilla, whereas in the present, retracted organ they lie in the wall of the sheath. Engel's synonymization of Dall, 1900, with Petalifera petalifera is not acceptable, because Phyllaplysia taylori has a small, second- ary, concave shell (Beeman, 1963: 44, fig. 1; 1968: 94). A secondary shell occurs also occasionally in Phyllaplysia engeli (Marcus, 1957a: 56, fig. 4), while Petalifera has a complete primary, convex aplysiid shell with its protoconch (Fig. 29; and Marcus, 1967a: fig. 48). The species is new for Brazil. Discussion.- Vicente (1961), who did not know Engel's paper (1936a), separated two Mediterranean species of "Aplysiella" from east of Toulon (A. virescens) which he observed alive, and preserved specimens from Algiers (A. webbii). He said that the differences concern radula, shell, digestive apparatus, gonads, and nervous system (Vicente, 1961: 5, 14). To my view, the entire radula of webbii (op. cit.: fig. 4, right) is identical with that of Vayssiere's drawing (1885: pI. 3, fig. 74), while that of virescens (fig. 4, left) seems to be much more relaxed, the transverse rows and the single teeth are separated. The shells (fig. 3) are of absolutely the same shape. A copy of the outlines fits onto both, but the upper border of the preserved, nontransparent shell of webbii is broken. The differences 1972] Marcus: Anaspidea of the Western Atlantic 857 of the digestive apparatus (fig. 6) may be due to the degree of contraction during preservation. The reproductive apparatus (fig. 7) might be in dif- ferent functional phases. The spermatocyst (fig. 7, no. 5) varies in shape according to its filling, and the annexed gland (no. 3), the prostate of Thompson & Bebbington (1969: 349, fig. 3 A), is differently developed. The figures of the nervous system (figs. 9 A and B) evidently differ also due to different treatment, as I have often observed. Hence I consider both populations as conspecific, and their names, following Engel, as synonyms of Petalifera petalifera. Engel (1936a: 54) united most of the described species of Petalifera under the name of P. petalifera (Rang, 1828) as a "Rassenkreis," and pre- served their original names as subspecific, mainly due to their geographic distribution. I accept his name Petalifera petalifera bonairei for the present material from Puerto Rico and northern Brazil, because it has the typical radula of P. petalifera. The distribution of this cosmopolitan species over- laps with that of the later described P. ramosa Baba, 1959, occurring in Florida (Marcus, 1962a: 453) and in the present collections in the Carib- bean area, from lIoN, 600W to 21°N, 86°W.

14. Petalifera ramosa Baba, 1959 Figs. 7, 16, 21, 35, 41, 50, 59, 64, 70 References.-Baba, 1959: 337, fig. 1 A-E (P. ramosa); Eales, 1960b: 12, 6 figs. (P. habei), 112 (P. ramosa); Marcus, 1962a: 453, figs. 1-3; 1967a: 39, fig. 48 (P. ramosa). Material.-PILLSBURY Sta. 746: 24. VII. 1968, 11°54'N, 66°54.5'W, 23-27 m. PILLSBURYSta. 965: 20. VII. 1969, 17°25.5'N, 63°23'W, 24 m. PILLS- BURYSta. 979: 22. VII. 1969, 17°51.5'N, 62°38.7'W, 49 m. PILLSBURY Sta. 886: 7. VII. 1969, 14°09'N, 60057.1'W, 18 m. A total of 13 specimens. Further Distribution.-J apan; Florida. Description.-The preserved specimens measure from 10 X 6 X 4.5 mm and 18 x 7 x 5 mm to 20 x 13 x 8 mm and 21 X 11 x 7 mm. They are contracted in different ways, long and narrow or broad and flat, with a more or less pronounced neck. The papillae are generally contracted and form soft knobs; some may be distended and ramified. The pigment has disap- peared in the preserving liquids, Bouin and alcohol, except for traces on or around the notal papillae. The oral lobes are well developed (Fig. 35) and in very different positions according to contraction, either triangulate to the sides of the mouth, or drooping backwards, or extended transversely. Also the tentacles vary. They are rolled in, and in some animals appear as curved tubes beset with tubercles, while in others they are folded to the ventral side. The rhino- phores are far apart from one another, as long or longer than the tentacles, 858 Bulletin of Marine Science [22(4 )

FIGURES 31-36. Ventral view of anterior end: 31, Notarchus punctatus ar- malus; 32, Stylocheilus longicauda; 33, Bursatella leachii pleii; 34, Dolabrifera dolabrifera; 35, Petalifera ramosa; 36, Phyllaplysia engeli. (The dashed lines complete the outlines of the transverse sections of the animals.) (r, lateral ridge; s, sole.) 1972] Marcus: Anaspidea of the Western Atlantic 859 and may bear a ramified papilla besides many tubercles. The eyes lie behind, or beside, or slightly in front of, the rhinophores. The animals have a sharp edge between upper and under surfaces (Fig. 41). The upper surface corresponds to the outer side of the para podia and bears papillae. The underside is differentiated into an opaque glandular ciliated sale, about half the breadth of the body, and a transparent hypo- notum to its sides (Fig. 16). The anterior border of the hyponotum unites with the outer side of the tentacles (Fig. 35). The anterior border of the sale is thickened by a strip of glands which open into a transverse groove, producing the typical "bila- biate anterior foot border." The posterior end of the body is rounded or slightly pointed. The dorsal slit is situated near the middle, more in the posterior half of the body (Fig. 7). Its length is one-fourth to one-half of the body length. In some preserved specimens the mantle shelf overlaps the right parapodial flap. The shell foramen is wide: its border hardly covers the border of the shell. The shell is 6.5 mm long and 4.5 mm broad. Its shape is round anteriorly with a narrow, long apex and a distinct anal sinus. In several specimens the shell was lost, in others it is decalcified. The opaline gland is simple and multiparous (Fig. 41), corresponding to this character in the subfamily. Mazzarelli (1890, cited from Hoffmann, 1934: 472, fig. 346 F) evidently mistook the prostatic gland on the genital duct, which is very well developed in Petalifera, for the opaline gland. The jaw plates, 1.15 by 0.67 mm, are composed of hollow elements (Fig. 50) in about 25 rows, measuring 21,u X 8fL on the surface and up to 60fL high. Sometimes they have a row of fine denticles on one side. The palatal rods of the common shape with one cusp are 50fL-I00fL long (Fig. 59). The radula is 1.5 mm long, 2.0 mm wide. Its formula is 30-36 X 35- 54.1.35-54. The rhachidian tooth has three denticles on either side of the middle cusp (Fig. 21). Sometimes there is a pair of small accessory denti- cles on the main cusp. The angle of the feet of the rhachidian tooth varies. The length of the lateral teeth and the number of their denticles increase outwards; only the outermost teeth are reduced. There are 2-7 denticles on the inner side and 2-5 on the outer side of the pointed main cusp, which is bifid on some teeth, singly pointed on others. The gizzard has 9-12 large teeth and several smaller ones. There is no caecum. The stomach of one specimen contained many spicules of sponges. The male organ has a small collar with two strong spines on warts, and about six smaller spines on the muscular penis (Fig. 64). From the tip of the penis stands out a thin, ciliated appendage without spines. The penial sheath is wrinkled and does not bear any spines. The nerve ring differs from that of Petalifera petalifera (Vayssiere, 1906: 860 Bulletin of Marine Science [22(4 )

43

FIGURES 37-43. Transverse section on level of gill: 37, young specimen of ; 38, Notarchus punctatus armatus; 39, Stylocheilus citrinus; 40, Bursatella leachii pleii; 41, Petalifera ramosa; 42, Phyllaplysia engeli; 43, Dolab- rifera dolabrifera. (b, opaline gland; c, shell; g, gill; h, heart; n, kidney; p, purple gland.) (The opaline gland secretion is drawn black.)

48, fig. 65) by longer connectives between the left pallial and visceral ganglia and between left and right visceral ones (Fig. 70). Remarks.-As in the previous Floridian material, the present radulae generally have more numerous denticles on the inner side of the lateral teeth than on the outer, contrary to the Japanese specimens. Eales (1960b: 14, fig. 3b) described and figured the male organ of P. ramosa from Japan. This has more spines on the collar than in the present animals. Eales did not mention a ciliated appendage on the papilla. Perhaps these slight differences 1972] Marcus: Anaspidea of the Western Atlantic 861 would justify a specific separation of the two forms, but I prefer to wait for more ample material before creating a new species.

15. Phyllaplysia engeli Marcus, 1955 Figs. 8, 17, 23, 26, 42, 51, 60, 67, 71 References.-Marcus, 1955: 105, figs. 29-37; 1957a: 55, figs. 1-16; 1970a:27. Material.-Barbados: Helen Hughes leg., 1970. Florida: Miami, Corinne E. Edwards leg., III. 1971; Long Key, Robert Austin Smith leg., II. 1972. Puerto Rico: Maguey Island, 22. I. 1972 (E. M.). Brazil: Pernambuco, Paracuru, 19. II. 1969; Itamaraca, 8. VIII. 1968; Espirito Santo, Guaraparf, 6. XII. 1968, Pierre Montouchet leg. A total of 23 specimens. Further Distribution.-From Florida to Brazil, Sao Paulo. Remarks.-The 130 specimens of Phyllaplysia engeli collected on the coast of Sao Paulo since our description of the shell in 1957 were now studied with respect to their shells, though these might be decalcified. The largest specimen, collected by Miss Kaoru Hiroki, was 26 mm long alive, 18 mm preserved; it had a cuticular shell 0.5 mm in diameter with a calcified central pad 0.3 mm in diameter. The next in size, 12 and 9 mm preserved, had no trace of a shell. Of the smaller ones a great number were examined, but only four of them, of middle size, had a distinct brown, cuticular shell. Evidently the shell is not quite as rare in Phyllaplysia taylori (Beeman, 1963: 44). Two specimens from Sao Sebastiao, S. P., spawned in the aquarium. The spawns are 8 and 18 mm long and 5-7 mm wide (Fig. 17). The mucous ribbon is 2JL-300JL wide; it lies flat on the alga in hairpin windings. The eggs measure about 60JL in diameter; I reckon about 20,000 to one spawn.

COMPARATIVE NOTES The external shape (Figs. 1-8) is no safe character for the Anaspidea. Notarchus is frequently globose in preserved state, Petalifera and Phyll- aplysia are generally flattened, while Aplysia, Stylocheilus, and Bursatella are often laterally compressed. Dolabrifera is intermediate; it has a strongly muscular body wall, much firmer than the larger Bursatella. The presence or absence of a shell is distinctive, if the shell is typically developed. The oral lobes are not set off from the anterior border of the tentacles in Aplysia (Eales, 1960a: fig. 19 B). They are separated from the tentacles in varying degree in the other species examined here (Figs. 31-36). Even in one and the same species (e. g., in Dolabrifera), the contraction or dis- tension of the buccal muzzle changes the aspect of the oral lobes from broad lappets to narrow ridges (Fig. 24). 862 Bulletin of Marine Science [22(4)

45 aID) 48 47

FIGURES 44-51. Elements of jaw plates: 44, Aplysia juliana; 45, Notarchus punctatus armatus; 46, Stylocheilus longicauda, near margin (above) and in middle of jaw plate (below); 47, Bursatella leachii pleii; 48, Dolabrifera dolabri- fera; 49, Petalifera petalifera bonairei; 50, Petalifera ramosa; 51, Phyllaplysia engeli.

In the thickened anterior border of the foot there is always a transverse furrow into which the anterior pedal glands open, but it is not always dis- tinct, nor is the median notch in the border. The lateral extension of the anterior border of the foot corresponds to the breadth of the ciliation of the sole (Figs. 37-43), which is, in all the present species, narrower than the greatest width of the body. In Aplysia, Notarchus, Stylocheilus (Fig. 15), and Bursatella, the parapodia arise immediately from where the sole ends on the sides, while in Dolabrifera, Petalifera (Fig. 16), and Phyllaplysia, a broad, unciliated border of the under side, a hyponotum, is set off from the upper side by a sharp ridge around the widest part of the body. In Dolabri- fera this is the edge of a flat frill (Fig. 6), which Pease (1868, cited from Engel, 1936a: 40) supposed to be adapted for swimming. All Brevicaudata may bear warts or papillose projections which can be completely retracted, so that the animals appear quite smooth. The relative size of rhinophores and tentacles is variable, as well as the position of the eyes in relation to the rhinophores. There are many terms applied to the space between the parapodia. Thiele 1972] Marcus: Anaspidea of the Western Atlantic 863

FIGURES 52-60. Palatal rods: 52, Aplysia moria; 53, Notarchus punctatus ar- matus; 54, Stylocheilus longicauda; 55, Stylocheilus citrinus; 56, Bursatella leachii pleii; 57, Dolabrifera dolabrifera, young (left) and old (right); 58, Petalifera petalifera bonairei; 59, Petalifera ramosa; 60, Phyllaplysia engeli. called it a branchial cavity. Eales (1944: 8) and Engel (1936b: 113) used mantle cavity for the entire space. Hoffmann (1933: 309) called the cavity under the mantle shelf a mantle cavity, and in the case of Notarchus, where it extends between foot and visceral sac (p. 254), a parapodial cavity. Perrier & Fischer (1911) said pallial cavity; I follow them. The distinction between mantle and pallial cavities is clear on the left side, where the mantle shelf separates them. On the right side the lining of the parapodium is continued into that of the pallial cavity which lodges the gill; only a furrow between the inner wall of the right parapodium and the floor of the cavity may be considered as the limit. The size of the pallial cavity does not separate the subfamilies equally. It is widest in Notarchus (Pelseneer, 1888: 83, text fig. 4), where it sur- rounds the visceral sac on all sides (Fig. 38) except in front. The small dorsal slit acts as a spout for swimming by jet propulsion (Martin, 1966; Haefelfinger & Kress, 1967: 130; Farmer, 1970: 74). The cavity is much smaller in Stylocheilus (Fig. 39), but still enables the animal to swim 864 Bulletin of Marine Science [22(4 ) (Engel, 1930: 211). Engel supposed that Stylocheilus swims in the same way as Notarchus, filling and contracting the spacious mantle cavity. How- ever, Macnae (1962: 197) observed it swimming butterfly-fashion with outstretched parapodia. Aplysia swims in the same manner, with flapping movements. For Dolabrifera, Pease supposed that it swims with movements of the frilled mantle border. Its strong musculature might favor this move- ment, but I think that the animals are too heavy for swimming. The pallial cavity of Phyllaplysia is wider that that of Stylocheilus (Fig. 42), but it evidently does not swim. Hoffmann (1934: 311) recorded that there are no glands of the mantle cavity in Bursatella, because there is no mantle. Engel (Eales & Engel, 1935: 288) could not detect an opaline gland in one specimen of Bursatella leachii africana; in a second he stated (p. 291) that, "if present, it is diffuse." However Bergh (1902a: 354) found the "An deutung einer Mantelplatte" and figured the "pigmented margin of the pericardium" (1902b: 170-171, pI. 2, fig. 1, C; 1907, pI. 3, f. 16 a) and described "on the floor of the gill cavity a white dotted thin layer, the subbranchial gland." Also, Bebbington observed the purple and the opaline glands (1969: 325, figs. 3, 4). They are conspicuous in favorable specimens in the present material (Fig. 19). Risbec (1928: 49) found a black mass with white granules between the opened parapodia of "Aclesia striata," now Stylocheilus risbeci Engel (1936a: 69). In one 20-mm specimen of Stylocheilus longicauda, I found both glands thick and hard with coagulated secretion (Fig. 39), while in the other animals of the same lot, I could barely detect their location. The same holds for Notarchus punctatus, in which I could hardly find the glands (Fig. 38), contrary to Bergh (1902a: 345), who described the strongly developed purple gland. The purple glands (Hoffmann, 1933: 414), which Blochmann called Mantelranddrtisen (1883: 80), are unicellular glands-of-Blochmann like the opaline ones, situated on the underside of the mantle fold. Pelseneer (1888: 80) called them hypobranchial, Eales (1960a: 270) subbranchial. In the shell-less Bursatella (Fig. 18) they open on the dorsal side of the vestigial mantle fold, in Stylocheilus on its right side. The opaline glands-called the anal gland by Pelseneer (1888: 80), Bohadsch's gland or glandola del veneno by Mazzarelli (1889: 580), and the hypobranchial gland by Bergh (1902a: 345) and MacFarland (1909: 17, 40)-are situated inside the cavity, behind the genital aperture, on the right wall and on the floor. In Dolabrifera and Phyllaplysia, they occupy the floor to its hind end. Perrier & Fischer (1911: 155, note 1) maintained that the gland of Bohadsch lies outside the pallial cavity in Aplysiids, not related to the hypobranchial gland, which "for all the world is a pallial gland" ("qui pour tout Ie monde est nne glande palleale"). Mazzarelli took the innervation of the opaline gland by the pedal ganglia as proof that it 1972] Marcus: Anaspidea of the Western Atlantic 865 belongs to the foot. MacFarland (1909; 53) saw also an abdominal nerve going to the opaline gland. The shape of the shell is rather constant in each species, but there is a certain variability in its proportions, e. g., Engel (1936a: 33-36, fig. 16) gave the measurements and drawings of about a hundred shells of Dolabri- fera, in which the width was from 40-60 per cent of the length. In Bouin- preserved animals it is decalcified. Bergh (1902a: 342) found the shell missing in a specimen of Aplysia punctatella (Bergh, 1902), though "the mantle was undamaged." We found the shell completely absent in a speci- men of Dolabrifera (Marcus, 1962a: 453). It is also missing in some of the present animals preserved in alcohol. In Petalifera, where the mantle border covers only the rim of the shell, the latter is sometimes lost. On the other hand, Vayssiere found a shell in Notarchus (1885: 84, fig. 81). In Phylla- plysia, a secondary shell mayor may not be present (Marcus, 1957a: 56; Beeman, 1968: 94). Bergh's Aplysiella viridis (1905: 25, pI. 3, fig. 3, pI. 8, figs. 28-37) was described from a single specimen. Bergh was not sure of the absence of a shell. Engel (1936b: 206, figs. 2, a-q) studied a second animal of the same lot and noted the likeness of the sale with that of Petalifera petalifera. The radular teeth of both specimens agree well. The laterals have long, bifid cusps with several outer denticles as in Petalifera, not the short, square cusps of Phyllaplysia lafonti (Engel, 1936b: 202, fig. 1, e) or P. engeli (Marcus, 1955: 106, fig. 33). I conclude that Aplysiella viridis Bergh, 1905, called Phyllaplysia by Engel, really belongs to Petalifera. Due to the very wide mantle foramen, the shell had been lost in several of the present specimens of P. ramosa. From the recent reexaminations of Phyllaplysia taylori DaB, 1900, by McCauley (1960), Marcus (1961: 10) and Beeman (1963; 1968), it is evident that this is a good species of Phyllaplysia with a secondary shell and not "a synonym or a far divergent race" of Petalifera petalifera (Engel, 1936a: 54). The jaw elements (Figs. 44-51) are tightly packed at their bases. They are flat on the outer margin of the jaw plates, and high, generally hollow columns near the middle. Their tips are smooth, or bear more or less distinct denticles. Specific differences, e. g., between Petalifera p. bonairei (Fig. 49) and P. ramosa (Fig. 50), are greater than those between the genera, e. g., P. ramosa and Notarchus (Fig. 45). The quadrangular hollow plates on the border of the jaw bear distinct denticles, which disappear towards the middle, where the elements are high. The same holds for the two species of Stylocheilus (Fig. 46). The jaw elements of Phyllaplysia (Fig. 51) have an oblique, coniform tip, sometimes bearing several points. The big, delicate rods of Bursatella (Fig. 47) have a vesicular top. The jaw rodlets of Dolabrijera, even of 60-mm specimens, are so delicate and small, that 866 Bulletin of Marine Science [22(4) they easily escape observation (Fig. 48). That is probably the reason why Bergh took the much heavier palatal rods for jaw elements (1872: 441, pI. 5, figs. 17, 21, 22). The radular teeth (Figs. 9-14,20-23) are rather uniform throughout the order. The number of denticles to the sides of the central cusp of the rhachidian tooth varies from one on each side in Phyllaplysia (Fig. 23) to three in Stylocheilus (Fig. 12). Sometimes there are secondary dentic1es on the cusp-many in Aplysia (Figs. 9-11), fewer in Notarchus (Fig. 14), Stylocheilus longicauda (Fig. 12), and Bursatella (Fig. B)-so that this character seemed possibly to separate the subfamilies. However, the cusp may be smooth in Stylocheilus citrin us (Marcus, 1962b: 18, fig. 3), and sometimes it has denticles in Petalifera ramosa (Marcus, 1962a: fig. 3) and in Dolabrifera (Baba, 1936: text fig. 4 A, a). The palatal rods (Figs. 52-60) generally stand isolated from one another. In preparations they are frequently all orientated so as to show their frontal or basal face and appear as slender rods. In side view they are flat hooks (Fig. 54). They are often stratified (Figs. 52, 57). The hooks of Bursatella are covered with a layer of different refraction around their tips (Fig. 56). Bergh (1972: 441, pI. 5, figs. 17,21,22) took the palatal rodlets of Dola- brifera (Fig. 57) for the jaw platelets, because they are so densely set in old animals. In small ones, they are far from one another and are typical, often stratified, flat hooks. Hoffmann (1938: 1048, fig. 719 D) misinterpreted Vayssiere's (1885) drawing of the palatal hooks of Petalifera petalifera (Aplysiella webbii, pI. 3, fig. 73, bis); he thought the indented border was the cutting edge; it is, however, the fastened base of the hook, and the smooth, pointed cusp is typically directed downwards (Fig. 58). The male organ (Figs. 61-67) and its armature are variable. This is partly due to the state of development of the organ. In the present young specimen of Notarchus, the male organ is covered with soft papillae (Fig. 61), but in old ones there are spines without basal knobs. The different forms in Dolabrijera are mentioned above. In Stylocheilus longicauda from northern Brazil, the penial papilla was slender with a few spines on warts on its base, and there were several big, spiny warts in the sheath. Specimens from Cura~ao (Fig. 62), like those of S. citrinus (Marcus, 1962b: fig. 6), had the papilla covered with spines not standing on warts, and, along the sheath, rows of 12 big spiny warts each, and some more spines at the ectal end of these rows (see also Engel, 1936a: figs. 37,41). The papilla of an animal from the Gulf of California had simple spines on one side and spiny warts on the other (Marcus, 1967a: fig. 17). The size of the spines that we considered to be specifically different in longicauda and citrinus (1963: 15) is evidently correlated with the size of the animals, not with the species. The penis of Phyllaplysia taylori (as zostericola) was described with and without spines (Marcus, 1962a: 453). 1972] Marcus: Anaspidea of the Western Atlantic 867

61

FIGURES 61-67. Male copulatory organ: 61, Notarchus punctatus armatus, young; 62, Stylocheilus longicauda; 63, Bursatella leachii pleii; 64, Petalifera ramosa; 65, Petalifera petalifera bonairei; 66, Dolabrifera dolabrifera from Recife; 67, Phyllaplysia engeli.

The division of the Anaspidea by reason of the shape of the shell and the concentration of the nervous system (Eales, 1944: 20) serves well for the separation of the Longicommissurata (= Aplysiidae) and Brevicommis- surata (= Notarchidae). In Eales's diagnoses (1944: 16,18), Notarchinae and Dolabriferinae are distinguished by the concentrated ganglia in the visceral loop of the Notarchinae and the distinct ganglia of the Dolabri- 868 Bulletin of Marine Science

FIGURES 68-75. Central nervous system: 68, Aplysia cervina; 69, Stylocheilus citrin us; 70, Petalifera ramosa; 71, Phyllaplysia engeli; 72, Petalifera petalifera bonairei; 73, Bursatella leachii pleii; 74, Dolabrifera dolabrifera; 75, Notarchus punctatus armatus (modified from Vayssiere, 1885). (d, pedal ganglion; u, pleural ganglion; v, visceral ganglion; z, cerebral ganglion. The visceral con- nectives are drawn black.) ferinae. However, in Stylocheilus, allotted to the Notarchinae (Eales, 1938: 90, fig. 8), there are three visceral connectives, which I also found (Fig. 69), longer than the connectives in the dolabriferine Petalifera ramosa (Fig. 70). The latter has two distinct connectives, while in the congeneric P. p. bonairei there is only one, and the visceral ganglia are contiguous (Fig. 72). In the notarchine Bursatella (Fig. 73) both visceral and the right pleural ganglia are almost confluent, but in the dolabriferine Dolabrifera (Engel, 1927: 97; and Fig. 74) all four ganglia are confluent, just as they are in the notarchine Notarchus, so that this distinguishing character does not work for the separation of subfamilies in the combination proposed by Eales. The visceral ganglia of the dolabriferine Phyllaplysia (Mazarelli, in Holtmann, 1936: fig. 512b; Marcus, 1957a: fig. 11) are distinct, though there may be only two connectives in the visceral loop (Fig. 71).

CONCLUSIONS The separation of the order Anaspidea, without the Akeridae, considered as Cephalaspidea, into the suborders Longicommissurata and Brevicommis- 1972] Marcus: Anaspidea of the Western Atlantic 869 surata (Pruvot-Fol, 1954: 95, 101) is obvious. The former correspond to the family Aplysiidae with the subfamilies Aplysiinae and Dolabellinae, of which the first consists of two genera, Aplysia with five subgenera, and the Indopacific Syphonota, probably only one species (Eales, 1960a: 270); the Indopacific Dolabellinae consist of a single genus, Dolabella, with two spe- cies (Engel, 1942: 233). The Brevicommissurata, family Notarchidae, are divided into the subfamilies Notarchinae and Dolabriferinae. According to the characters considered, the genera would be united in different ways. The ciliated sole over the whole breadth of the foot (Figs. 15, 38-40) connects Notarchus, Stylocheilus, and Bursatella against Dola- brifera, Petalifera, and Phyllaplysia, in which the ciliation of the sole is restricted to a middle band about half breadth of the body (Figs. 16, 41- 43). The differences between Notarchus on the one hand, and Stylocheilus and Bursatella on the other, all three generally allotted to the Notarchinae, are much greater than those between the latter two and the remaining Brevicommissurata. The extension of the pallial cavity (Fig. 38) under the visceral mass separates Notarchus from the rest of the Notarchidae, in which the visceral mass is fastened to the foot along its whole underside. So the attribute in the family diagnosis given by Franc (1968: 850), "une cavite profonde ou flotte Ie sac visceral a insertion anterieure," would cor- respond to my subfamily Notarchinae, as only Notarchus has this character. The presence of one to three connectives in the visceral loop separates Stylocheilus, Petalifera, Phyllaplysia, and Bursatella from Dolabrifera and Notarchus, in which there are no connectives and the visceral and pleural ganglia are all contiguous. However, Petalifera petalifera and P. ramosa differ from each other in that the former has one connective from the left pleural to the left visceral ganglion, while the latter has a second connective between the right and left visceral ganglia.

SUMARIO SOBRE ANASPIDEA (GASTROPODA: OPISTOBRANQUIA) DE LAS AGUAS CALIDAS DEL ATLANTICO OCCIDENTAL Se da una clave para las 16 especies de Anaspidea del Athintico occidental y las dos del Athintico medio. Estas son ocho (y dos) especies de Aplysia y Notarchus punctatus armatus, Stylocheilus longicauda, Stylocheilus citri- nus, Bursatella leachii pleii, Dolabrifera dolabrifera, Petalifera petalifera bonairei, Petalifera ramosa y Phyllaplysia engeli. Se dan descripciones de Petalifera p. bonairei y Petalifera ramosa. Tambien se da una clave para los subgeneros del genero Aplysia. Se dan notas comparativas de caracteres anat6micos y sistematicos, planta del pie, cavidad palial, concha y sistema nervioso. 870 Bulletin of Marine Science [22(4) Mi subdivision de Notarchidae deja s6lamente a Notarchus en Notar- chinae; el resto de los generos son asignados a Dolabriferinae. Tres especies de Aplysia fueron rastreadas a 348-357 m de profundidad.

ZUSAMMENF ASSUNG Bestimmungstabellen flir die 16 westatlantischen und 2 mittelatlantische Anaspideen. Dieses sind 8 (und 2) Arten von Aplysia, Notarchus punctatus armatus, Stylocheilus longicauda, Stylocheilus citrinus, Bursatella leachii pleii, Dolabrifera dOlabrifera, Petalifera petalifera bonairei, Petalifera ramosa, und Phyllaplysia engeli. Petalifera p. bonairei und P. ramosa werden beschrieben. Vergleichende Bemerkungen tiber anatomische und systematische Merkmale, Sohle, Pallialh6hle, Schale, und Nervensystem. Meine Einteilung der Notarchidae Hisstnur Notarchus in den Notarchinae. AIle tibrigen Gattungen kommen zu den Dolabriferinen. Drei Aden von Aplysia wurden in 348-357 m Tide gefischt.

RESUMO Chaves para as 16 especies dos Anaspidea do Atli'mtico ocidental mais 2 do Atlantico central. Sao estas 8 especies de Aplysia (mais 2), Notarchus punctatus armatus, Stylocheilus longicauda, Stylocheilus citrin us, Bursatella leachii pleii, Dolabrifera dolabrifera, Petalifera petalifera bonairei, Petalifera ramosa, e Phyllaplysia engeli. Petalifera p. bonairei e Petalifera ramosa Sao descritas. Notas comparativas dos carateres anatomicos e sistematicos, a sola, a cavidade paleal, a concha, e 0 sistema nervoso. A minha subdivisao dos Notarchidae coloca somente Notarchus nos Notarchinae, e deixa todos os generos restantes nos Dolabriferinae. Tres especies de Aplysia fcram dragadas de 348-357 m de profundidade.

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