On Some Opisthobranchs from Florida

Home , Berghia, Ercolania, Rictaxis, Trapania

ON SOME OPISTHOBRANCHS FROM FLORIDA

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

ABSTRACT Eleven species of opisthobranchs from Florida are treated. Bosellia marcusi and Trapania dalva are species new for science. Cylindrohulla ulla Marcus, 1970, is the type-species of the new genus Ascohulla and new for North America. Ercolania funerea, Elysia ornata, E. elena, Discodoris mortenseni, Trinchesia perca, and Berghia verrucicornis are recorded from Florida. Doto chica is discussed. Several species have their names changed due to recent work of Baba and Hamatani, Habe, Tardy, Edmunds, and Marcus. These are "Acteon" punctostriatus, now Rictaxis punctostriatus, "Stiliger" funereus, now Ercolania funerea, "Carriona" perca, now Trinchesia perca, and Berghia "coerulea," now B. verrucicornis.

INTRODUCTION During three visits to the Rosenstiel School of Marine and Atmospheric Science of the University of Miami in October 1969, November 1970, and January 1971, I collected a number of opisthobranchs for study. I also received several specimens and color photographs from Lt. Col. Corinne E. Edwards, USAF, Mrs. Sally Diana Kaicher, Mrs. Patricia Torrance, and Dr. M. Patricia Morse. These collections contain two species new for science and several species new for Florida. The names of some species had to be changed also. My sincere thanks go to Prof. Frederick M. Bayer for his friendship and hospitality and for the many trips he and Mr. Robert Work undertook with me; to Lt. Col. Corinne Edwards, who accompanied me when collect- ing; to Mr. Dennis Opresko, who gathered the Ascobulla and Rictaxis; to Prof. S. A. Gerlach, for a specimen of Elysia ornata from the Red Sea; and to Dr. Joseph P. Rosewater and Mr. Walter J. Byas, who gave me some dry acteonids with radulae. Mr. Robert Austin Smith kindly revised my manuscript and bibliography. The material has been sent to the Smithsonian Institution, Washington, D. C.

LIST OF SPECIES Order CEPHALASPIDEA Superfamily ACTEONACEA Family Acteonidae 1. Rictaxis punctostriatus (c. B. Adams), comb. nov. Figs. 1-6. Superfamily CYLINDROBULLACEA Family Cylindrobullidae 2. Ascohulla (gen. nov.) ulla (Marcus, 1970). Figs. 7-17. 1972] Marcus: Opisthobranchs from Florida 285

Order ASCOGLOSSA Superfamily ELYSIACEA Family Stiligeridae 3. Ereolania junerea (A. Costa, 1867). Family Elysiidae 4. Elysia ornata (Swainson, 1840). Fig. 18. 5. Elysia dena Marcus, 1970. 6. Bosellia marcusi, spec. nov. Figs. 19-22. Order DORlDOlDEA Suborder EUDORIDACEA Tribe CRYPTOBRANCHIA Family Dorididae (Subfamily Discodoridinae) 7. Diseodoris mortenseni Marcus, ] 963. Tribe PHANEROBRANCHIA Superfamily Suctoria Family Goniodorididae 8. Trapania dalva, spec. nov. Figs. 23-26. Order DENDRONOTOlDEA Family Dotoidae 9. Doto ehica Marcus, ] 960. Figs. 27-36. Order EOLlDOIDEA Suborder ACLEIOPROCTA Family Cuthonidae 10. Trinehesia perea (Marcus, 1958). Family Aeolidiidae 1 I. Berghia verrueieornis (A. Costa, 1864). 1. Rictaxis punctostriatus (c. B. Adams, 1840) Figs. 1-6 References.-Tornatella puncto-striata Gould, 1870: 224, fig. 515. Actaeon punctostriatus Pilsbry, 1893-1895: 157, pI. 18, figs. 98,99 (from d'Orbigny), pI. 19, fig. 22 (from Gould), fig. 23 (from Verrill, variety). Aceton punctostriatus Abbott, 1954: 275, pI. 26, fig. T. not Acteon punctostriatus Marcus, 1958a: 32, figs. 1-10. ? Acteon punctostriatus Marcus, 1970b: 924, fig. 5. Rictaxis punctostriatus Marcus, 1971b, in press; 1972: 178-183, figs. 2, 30-37.

Material.-Florida: SE of West Point on Biscayne Bay shore of Key Biscayne, salinity normal, algae and mud, November 1970, five specimens. Marco, USNM No. 53731, two dry shells with radula. Indian River, USNM No. 358116, two dry shells with radula. Maryland: Solomons, in oyster aquarium of the Chesapeake Biological Laboratory, September 1969, eight animals and three spawns. 286 Bulletin of Marine Science [22(2)

FIGURE 1. Rictaxis punctostriatus (C. B. Adams); shells from Indian River, Marco, and Miami, Florida.

The further distribution cannot be given, as the shells are indistinguishable from those of Brazilian material, whose radula is that of a true Acteon. Remarks.-Habe (1956: 99) raised Dall's subgenus Rictaxis (1871) to generic rank by reason of its radular formula 5.0.5. As the present material has this formula (Fig. 6), it must be transferred to Rictaxis (Marcus, 1971b, in press). The Brazilian material, classified as A. punctostriatus, whose shells are indistinguishable from the North American ones, has a true Acteon-radula, and hence represents a separate species, which now has received the name Acteon pelecais (Marcus, 1971b, in press). The proportions of the shells in both species have the same range of variability, and the height of the columellar fold and the extent of the spiral striation vary in both.

Ascobulla, gen. nov. Diagnosis.-Cylindrobullidae with head shield, typical shell, and ascoglossan radula. Type-Species.-Cylindrobulla ulla Marcus (1970a: 25). The two species of Cylindrobulla known for their anatomy, the genotype C. beaui Fischer, 1856 (Marcus, 1970a: 22) and C. £lila, differ considerably in the shape of their radular teeth. These are diaphanacean in C. beaui (Marcus, 1970a: figs. 30, 31) and ascoglossan in £lila (Fig. 11). 1972] Marcus: Opisthobranchs from Florida 287

FIGURES 2-6. Rictaxis punctostriatus (C. B. Adams): 2-4, medium, narrow, and widest shells, respectively, from Solomons, Maryland; 5, jaw platelets; 6, half-row of radula.

Similar differences occur between the radulae of the Acochlidioidea Microhedylidae, of which Ganitus Marcus (1953: fig. 25) and Paraganitus Challis (1968: fig. C) have ascoglossan teeth, while Parhedyle tyrtowii (Kowalewski, 1901: fig. 39), Unela (Marcus, 1953: fig. 4) and Hedylopsis Thiele, 1931 (Swedmark, 1968: fig. 2) have Toledonia- and Newnesia-like teeth (Hoffmann, 1938: figs. 682 G, P). The radula of ulla and that of C. japonica Hamatani (1969: 172, fig. 1) are so far different from that of C. beaui that these two species must be removed to the new genus Ascobulla. A head shield and the radula of C. bealli are cepha1aspidean characters incompatible with the diagnosis of the Ascoglossa, while the radula of Ascobulla and its pectinate gill, as well as the closed male duet in bealli and ulla, are typically ascoglossan. Baba (1966: 201), Kay (1968: 130), and Franc (1968: 844) placed the 288 Bulletin of Marine Science [22(2)

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FIGURES 7-1 I. Ascobulla ulla (Marcus): 7, I-mm shell; 8, apical view of 6-mm shell; 9, labial cuticle; 10, preradula and four oldest teeth; 11, largest teeth.

Cylindrobullidae at the basis of the Ascoglossa. Also Burn (1966: 52-53) treats them as a superfamily at the root of the Ascoglossa, within the suborder Juliacea (Boettger, 1962). In an earlier paper (Marcus, 1970a: 4, 25) they constituted a superfamily of the Cephalaspidea, and I retain them in this place.

2. Ascobulla ulla (Marcus, 1970) Figs. 7-17 References.-Marcus,1956: 119, figs. 1-12 (Cylindrobulla spec.); 1970a: 25, fig. 33 (c. ulla). Material.-Florida: Key Biscayne, bay shore, on outer side of mangrove, among algae, 20. XI. 1970, 22 specimens. Brazil: Pernambuco, 1972] Marcus: Opisthobranchs from Florida 289

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FIGURES 12-17. Ascobulla ulla (Marcus): 12. Diagram of reproductive organs; 13, tangential section of spermathecal epithelium; 14, transverse section of efferent duct and oviducal furrow; 15, transverse section of prostate; 16, transverse section of inner oviduct; 17, transverse section of spermoviduct. (a, ampulla; d, efferent duct; e, spermatheca; t, female aperture; h, hermaphrodite duct; io, inner oviduct; m, mucus gland; 0, ovotestis; p, penial papilla; q, prostate; sc, spermatocyst; so, spermoviduct; t, male atrium; v, vagina.) 290 Bulletin of Marine Science [22(2) Itamaraca, 8. VIII. 1968, Pierre Montouchet, three specimens; Oliven<;a, 17. I. 1969, Pierre Montouchet, one specimen. Further Distribution.-Brazil, Sao Paulo. Descriptive Notes.-The shells measure from 1 to 6 mm in length, with the greatest diameter in the upper third. The aperture is continued into a sutural slit about half around the body whorl. The callus in the inner lip forms a crescent-shaped crest around it (Fig. 8). In the smallest shells, the protoconch stands up in the center of the depressed spire (Fig. 7); in the larger ones, it is covered by the growth of the callus. The protoconch has two whorls, and there are about two adult whorls. Between the suture and the subsutural line there are radial cracks on the body whorl. The upper part of the mantle border is beset with a row of papillae similar to those figured for Volvatella (Baba, 1966: pI. 9, fig. 1). Our previous description of the shell needs no further additions. In the buccal apparatus treated with potassium hydroxide the cuticular lining of the buccal cavity, mentioned in our description (1956: 122), appeared as a brown funnel (Fig. 9). The radula of the I-mm specimen contains 15 teeth, the longest of which is 65p.,; the preradular tooth is 26p.,. The 5-mm animal has 28 teeth up to 183p., long. The number of teeth increases very slowly, but each tooth is longer than the one immediately anterior to it. Our first description of the reproductive organs of Cylindrobulla sp. (1956: 123) was based upon a single young specimen with a 2-mm shell. Now a series of sections of a 5-mm snail makes some corrections necessary and some amendments possible. The ovotestis is multilobate (Fig. 12, 0). The waIls contain the ovocytes; the lumen, the sperm. The hermaphrodite duct is a simple tube which leads the reproductive ceIls to a globular ampulla (a). From the ampulla the narrow muscular spermoviduct coils and then widens into a tube with glandular waIls and a star- shaped lumen (Fig. 17, so). This tube opens into a large chamber with ciliated, thin and richly folded epithelium (Fig. 16, io). The male duct going out from the chamber is surrounded by long and large prostatic cells (Fig. 15, q). It forms a ventrad loop and then descends into the cutaneous musculature (Fig. 14) in front of the female aperture. It runs forward, then winds (d) to the fundus of the deep male atrium which lodges the entire penial papilla (p). The male duct enters the long muscular penis and opens with a curved, cuticular stylet, 66p., long and 14p., wide at is base. The course of the male duct in the muscle layer is accompanied on the outside by a ciliated groove from the female aperture to the male pore (Fig. 14). In our previous paper, this furrow was the only pathway described for the sperm of the present species. In Cylindrobulla beaui we had 1972] Marcus: Opisthobranchs from Florida 291 found the internal duct (1970a: 24), and a shallow groove between head shield and foot. Beyond the exit of the male duct, the inner oviduct Cio) is continued with richly folded, thin, ciliated walls and opens into the wide female atrium. The latter is in broad connection with the voluminous female gland mass (m). This gland mass stains differently in its various folded sections. It is partly lined with long cilia. From the female atrium inwards, a narrow vagina (v) courses to the large spermatheca (e) apposed to the diaphragm. It is filled with irregularly heaped sperm. Most of the cells lining the spermatheca have two nuclei of equal size (Fig. 13). Halfway down the vagina (v) inserts the short duct to the pear-shaped spermatocyst (sc). The sperm in the latter are directed with their heads to the wall. Remarks.-Ascobulia ulia differs slightly from A. japonica (Hamatani, 1969). In the latter, the sutural slit continuing the aperture coils about 1% times (p. 173). In a letter of 26. VI. 1971, Hamatani informs me that the crest around the callus of the inner lip is inconspicuous. The protoconch "seems to be hidden half in the spire." The reproductive organs of japonica were not studied. Hamatani (1971) described another species of Cylindrobulia, C. californica, from the Gulf of California. Its sutural slit is closed at about two- thirds of the body whorl; for japonica, Hamatani now indicates "one time and more," so that the specific differences diminish. The variation of the curve of the ventral edge of the radular tooth is too great in Hamatani's figures 2 and 3 to allow for a specific distinction. The snow-white calcareous layer of the shell might be due to treatment. On plate 7, C. japonica, the shell appears less transparent than on pI. 6, C. californica.

3. Ercolania funerea (A. Costa, 1867) Rejerence.-Marcus, 1970a: 37, figs. 69-73 (Stiliger [StiligerJ funereus). Material.-Florida: Gulf of Mexico, Cedar Keys, 29°09'N, 83°05'W, near Seahorse Key, in tuft of Eudendrium, 16. X. 1969, one specimen. Remark.-The recent redescription of the type-species ornatus of the genus Stiliger Ehrenberg, 1831, by Baba & Hamatani (1970: 199-206, pis. 3,4) made an exact diagnosis of the genus and the family Stiligeridae possible, so that many changes of names become necessary. Ercolania is re- established to generic rank. It is distinguished by simple rhinophores from Stiliger with auriform rhinophores. Hence the present species with its simple, finger-shaped rhinophores (Marcus, 1970a: 38) comes under Ercolania. New for Florida. 292 Bulletin of Marine Science [22(2) 4. Elysia ornata (Swainson, 1840) Fig. 18 Synonyms.-E. grandifolia Kelaart, 1858; E. marginata Pease, 1871. References.-Baba, 1966: 201-202 (synonyms); Marcus, 1963: 20; 1970a: 43, figs. 80-83; Kenny, 1970: 85. Material.-Florida: Grassy Key, 22. X. 1969, two specimens. Red Sea: Sarad Sarso, Farasan Archipelago, 16°50'N, 41°32'E, XII. 1964, S. A. Gerlach leg., one specimen. Further Distribution.-Bermudas; Lesser Antilles; Pacific; Queensland; India. Description.-The living animals were up to 30 mm long. They were olive green when well fed, and orange when the green contents of the liver diverticula were eliminated. A black and orange band borders the parapodia. The rhinophores have a black band along their margin and an orange stripe below the tip. The outer sides of the parapodia and the middle of the back are stippled with black rings of variable size around the outlets of orange subcutaneous glands. In the animal from the Red Sea their diameter is up to 0.3 mm; in those from Florida, 0.25 mm. On the outer sides there are also white dots, concentrated near the middle and the hind end of the borders of the parapodia. The sole is light colored. The pericardial eminence is short. It receives two to three vessels on either side. The radular teeth (Fig. 18) are up to 205JL long; their base is l!.J of their length. They bear small denticIes, 2JL broad. The worn teeth are heaped up in the ascus. The male copulatory organ is stretched and slender in the studied specimen; the atrium has a diverticulum. Remarks.-Elysia ornata is similar to E. cauze Marcus, 1957. The latter is often beset with papillae; its dorsal vessels are different; the radular tooth is more compact, and it has no atrial diverticulum. New for Florida.

5. Elysia elena Marcus, 1970 References.-Marcus, 1970a: 49, figs. 81,89-90; 1970c: figs. 3-6 (Elysia elena). Material.-Florida: St. Petersburg, Boca Ciega Bay, just below low tide, on muddy bottom, Dave Mason, 4. XII. 1970, two specimens. Tierra Verde toll gate, Dave Mason, 14. III. 1971, 4 specimens. 1972] Marcus: Opisthobranchs from Florida 293 Further Distribution.-Curac;ao; Barbados. Remark.-A comparison with Elysia serca Marcus, 1955, has been pub- lished (1970c: figs. 3-6). Though even the smallest teeth in the material from Tierra Verde have no denticles, the material is considered to belong to E. elena. The species is new for Florida.

6. Bosellia marcusi, spec. nov. Figs. 19-22 Material.-Florida: Grassy Key, 22. X. 1969, on Halimeda, nine specimens and one spawn. Miami, Bear Cut rocks, at low tide, on Halimeda opuntia, Corinne E. Edwards col., 15. II. 1972,8 specimens. Description.-The living animals were 3-4 mm long and 1 mm broad. The fore end was rounded, the rhinophores stood out, and the tail was pointed (Fig. 19). Preserved, the slugs are oval and measure 2.0 and 1.2 mm, respectively (Fig. 21). The color in life was light yellow, mostly hidden by the green contents in the ramifications of the digestive gland. There were tiny yellow dots and blue spots on the back, and white blotches along the border and on the rhinophores, and three larger ones along the middle of the back. The areas around the big eyes, the length of which is 60JJ-, and the underside of the body are colorless. The neck is slightly narrowed in life, the head set off by transverse furrows in the preserved state. Dorsal vessels were not noticed. The rhinophores were slender and rolled in; in the preserved animals they are completely retracted. The genital apertures open into furrows on the right side. At the entrance of the mouth lie two melanophores. The epithelium of the ventral side contains many blue-staining glands. Around the mouth cavity there are bundles of blue-staining buccal glands. Behind the mouth is a bulky, multicellular, reddish pedal gland. The small pharynx is 230j.t long, including the ascus, and 150j.t high. The radula forms a spiral of 18 slender teeth and a heap of about 10 teeth in the conspicuous ascus. The largest tooth is 50JJ- long with an 18JJ- long base. Its edge bears a few blunt denticles near its tip (Fig. 20). There is no crop, contrary to Bosellia mimetica Trinchese, 1891 (Marcus, 1970a: 51). The narrow oesophagus opens into a relatively small stomach. A short intestine leads to the anus which opens into the anterior transverse fold. The follicles of the ovotestis are fewer and bigger than those of Bosellia mimetica. There are five or six, about 250JJ-Iong, against 200JJ-in mimetica, which has more than 15 follicles (Marcus, 1970a: 51). The female cells are developed in the fundus, the sperm in the ectal half (Fig. 22). The male duct passes through a muscular penial papilla bearing a curved stylet. 294 Bulletin of Marine Science [22(2)

FIGURES 18-22.-18, Elysia ornata (Swainson), radular tooth.-19-22, Bosellia marcusi, spec. nov.: 19, living slug; 20, radular tooth; 21, clarified slug; 22, sagittal section of region of receptaculum seminis. (b, bursa copulatrix; c, brain; d, efferent duct; f, female aperture; g, albumen gland; i, diverticulum of digestive gland; m, mucus gland; n, kidney; 0, ovotestis follicle; p, penial papilla; r, receptaculum seminis; s, sole glands; x, strand from receptaculum; y, pharynx; z, duct from receptaculum.) ]972] Marcus: Opisthobranchs from Florida 295 The penial sheath opens into the right transverse furrow behind the right rhinophore. The ramified albumen glands consist of large cells, 30,u to 50,u in diameter. The female aperture lies in a deep dorsal furrow on the right side, 0.3 mm behind the male pore. This furrow is visible in the living and in the preserved specimens (Figs. 19, 2], f). In the hind end of the body lies a vesicle filled with sperm (Figs. 21, 22, r), from which a duct runs forward along the midline. Its size is 130,u to 21O,u in length, 85,u to 175,u in breadth and height. My material is not sufficient to analyze this organ. Its lining consists of vesicular cells measuring ] 2,u to 25,u, with dark-staining nuclei of 3,u to 7,u. The nuclei are larger than those of the surrounding tissues. They are comparable to those in the kidney and in the salivary glands. The nuclei in the kidney are paler. From this lining of the receptaculum, a broad strand goes forward and almost touches the fundus of the kidney. A tube (z) with thin muscular walls leaves the anteroventral side of the vesicle and winds forward in the midline. I could not trace its further course; it does not seem to be connected with the albumen glands as are some similar vesicles in B. mimetica, described as ampullae by Renate Becker (1960: 200). One spiral spawn on Halimeda, 5 mm in diameter, is composed of seven whorls. The eggs lie in a single row; the capsules measure 240,u X 260,u, the yolk, 230,u, and the embryo, 120,u. The shape of the spawn is identical with that of the spawns figured by Portmann (1958: fig. 5). Discussion.-The occurrence of Bosellia on Halimeda on the coast of Florida was to be expected, but to discover a species different from that found at Cura~ao and in the Mediterranean was a surprise. At first sight the eyes were much bigger, 60fL in length, against those of mimetica with 35,u. The two pigment cells in the sides of the mouth are wanting in mimetica. In the ascus the teeth are heaped up, while the radula is straight at the end in mimetica. B. mimetica has a big muscular crop, missing in B. marcusi. The female duct of mimetica opens into a ventral furrow quite near the male pore; in marcusi it goes into a dorsal furrow farther behind. Though smaller in size, preserved 2 mm long against the 4 mm of the preserved mimetica, B. marcusi has larger and fewer cells. This is most obvious in the eyes, the follicles of the ovotestis, and the albumen glands. Such a difference of size of the cells occurs also in other genera, e.g., turbellarians of the genus Chordarion (Marcus, 1945: 27-29). Andre Franc (1968: 846) includes Bosellia in the Polybranchiidae, though the diagnosis of that family on the same page reads: "papilles dorsales grandes, aplaties, foliacees." I dedicate this species to the memory of my dear husband Ernst Marcus. 296 Bulletin of Marine Science (22(2)

FIGURES 23-26. Trapania dalva, spec. nov.: 23, dorsal view of living slug; 24, left side view of same; 25, radular tooth; 26, preserved slug.

7. Discodoris mortenseni Marcus, 1963 References.-Marcus, 1963: 30, figs. 36-39; 1970a: 65, fig. 116. Material.-Florida: Biscayne Bay, Norris Cut, January 1971, one specimen, preserved 15 mm long. Remark.- The species is new for Florida.

8. Trapania dalva, spec. nov. Figs. 23-26 Material.-Florida: Miami, Crandon Marina, seawall, on Zoobotryon, 22. X. 1969, one specimen. 1972] Marcus: Opisthobranchs from Florida 297 Description.-The living animal was 12 mm long, 4 mm broad, and 5 mm high (Figs. 23, 24). The color was cream with white bosses and brownish blotches. Preserved, the specimen is 5 mm long, 2 mm broad, and 2.4 mm high. The dark pigment is retained (Fig. 26); it occurs in the epithelium and in the subjacent layers of the body wall. The bosses contain opaque white granules, also preserved in alcohol. The sale and the foot corners are colorless; on tentacles, rhinophores, rhinophoral processes, extrabranchial processes, and gills, there are brown and white spots. The tentacles are short and slender. The rhinophores have five folia- tions on both sides, separated by a broad dorsal and a narrow ventral rhachis, and a long tip. The three stout gills were bipinnate in life; preserved, they are hardly indented. The smooth rhinophoral processes are curved backwards. Both these and the extrabranchial ones are pointed and 1 mm in length. The sale is retracted and narrow. Its anterior corners are tentacle-like in life (Fig. 24) and rounded in the preserved specimen (Fig. 26). The labial armature is composed of conical pointed platelets. The radula has 19 rows of teeth (Fig. 25), increasing to the foremost which are 80ft in breadth. They bear 18-22 denticles inwards to the main cusp, which is 50ft high, and one or two outer denticles, missing in the smallest and largest teeth. The reproductive organs of the single specimen were not dissected, to preserve the type-specimen as complete as possible after extracting the radula. Discussion..-T. dalva is distinguished from the known species of the genus (Kress, 1968a: 164) by its color pattern, somewhat similar to that of T. fusca (Lafont, 1874) (Haefelfinger, 1960: fig. 4, 6b). It differs from T. maringa Marcus, 1957 (p. 442) by the white bosses and spots, and because the dark pigment of maringa fades in alcohol while that of dalva is preserved. The radular teeth of the species of Trapania do not have outer denticles in T. fusca, T. graeffei (Bergh, 1880), T. tartanella (v. Ihering, 1886) and T. velox (Cockerell, 1901). Outer denticles occur in T. japonica (Baba, 1935); T. maringa Marcus, 1957; T. maculata Haefelfinger, 1960; T. lineata Haefelfinger, 1960; and T. pallida Kress (1968a, 1970). How- ever, this character cannot be used to separate the two groups, as in the present species outer denticles are wanting in some teeth. The color pattern of T. velox is almost identical with that of T. maculata (Haefelfinger, 1960: 227, fig. 6a), from which velox differs by the want of outer denticles. Haefelfinger (p. 226) called the tentaculiform foot corners labial tentacles. Burn's Drepaniella (1961a: 102), later renamed Eucrairia mapae 298 Bulletin at Marine Science [22(2)

FIGURES 27-33. Doto chica Marcus: 27, anterior part of lO-mm slug, preserved; 28, 1.7-mm slug, preserved; 29, ceras of lO-mm slug, side view; 30, inner view of same; 31, radular tooth of ] O-mm slug; 32, single loose ceras from Boca Ciega; 33, ceras of 6-mm slug from Boca Ciega. (k, gill.) 1972] Marcus: Opisthobranchs from Florida 299 (Burn, 1961b: 51), is now (personal communication) considered as a subgenus of Ancula. Burn did not dissect his single specimen to study the radula, which is probably of the typical formula for Ancula: 1.1.0.1.1. Burn (1961 a: 103) mistook the short tentacles of Ancula fuegiensis (Odhner, 1926a: 45) for foot corners, in which case the species would not have any tentacles, but there are no Goniodorididae without tentacles. The bifurcate rhinophoral processes are not distinctive of Burn's species, as Ancula gibbosa (Risso, 1818), A. pacifica Bergh, 1881, A. fuegiensis Odhner, 1926, A. evelinae Marcus, 1961, and A. lentiginosa Farmer, 1964, all have two more or less united processes lateral to the rhinophores. Hence mapae is congeneric with them. This greater number of extrabranchial processes is variable, with five in gibbosa, 3-4 in pacifica (MacFarland, 1906: 148; 1966: 123). Thus Burn separates the subgenus Ancula (s. str.) for gibbosa and pacifica from the subgenus Eucrairia for the remain- ing four species with only one process.

9. Doto chica Marcus, 1960 Figs. 27-36 Synonym.-Doto fragilis umia Marcus, 1969: 27, figs. 38-39. References.-Marcus, 1960: 167, figs. 58-60; 1963; 39; 1970a: 77.

Material.-Florida, Gulf of Mexico: Cedar Keys, 29D09'N, 83D05'W, near Seahorse Key, on a tuft of hydroids of Eudendrium, which came up with a crab trap, 16. X. 1969, three specimens, 1.7, 6.0, and 10 mm preserved, two loose cerata of the 100mm specimen, six spawns.-St. Petersburg, Boca Ciega Bayway, west of second toll gate, on Thalassia, two specimens, 6 and 4 rom, and a loose ceras of a third; one ektachrome transparency, 30. V. 1969, Sally Diana Kaicher leg.

Further Distribution.-Southwest Greenland, 60D27'N, VEMA station 17-29 RD, 366 m; Florida, Virginia Key; Puerto Rico, 4-5 m; Cura9ao, mangrove. Description.-Length of the specimens, preserved, is 10, 6, 6, 4, and 1.7 mm. Breadth of the largest specimen is 2.7 mm. The color of the four large animals and the ektachrome is yellowish with three irregular bands of black pigment, and black spots on the veil and the rhinophoral sheaths. The larger dorsal and lateral papillae have a dark core, and the tubercles on the cerata have a subepidermal heap of black granules under their tips. The sale and its frilled border are without pigment. The 1.7-mm slug is quite white. The color slide of the living animal shows many opaque white spots in the cerata and the rhinophores. The veil is evenly arched and expanded on the sides. A ridge goes from the base of the rhinophore over half the breadth of the veil. The rhinophoral 300 Bulletin of Marine Science [22(2) sheaths of the four large animals have wide scalloped margins lengthened in front. Around their bases there are some papillae. The sheath of the 1.7-mm slug is smooth. The rhinophores are completely retracted. The papillae in the middorsal row are scarce on the 10-mm specimen, and only one dorsal papilla occurs in one of the 6-mm slugs. The largest specimen has the inconspicuous scars of eleven cerata that have fallen off on either side, and the twelfth left ceras in place. Two of its 4-mm-high cerata are preserved (Figs. 29, 30). They bear seven rings of tubercles, with seven tubercles in the second ring from the top. The third and the following rings are interrupted on the inner side by the well- developed gill. The insertion of each ceras lies slightly above its lower end. In the small slugs the cerata are shorter, up to 2 mm high in the 4-mm animal, with a maximum of five rings. Gills are developed from the foremost to the penultimate pair. The jaws are extremely delicate. The radula in the largest slug consists of 115 teeth. They are of the typical shape for Doto, with two salient rows of 4-5 denticles on the sides over the not prominent cusp. The position of the lateral denticles varies and is repeated each fourth tooth. The follicles of the ovotestis of the 4-mm-Iong, sectioned specimen contain large, yolk-laden eggs in the fundus and sperm in the ectal part. In front of the ampulla (Fig. 36, a) the spermoviduct divides into the male and the female duct. The former immediately forms an ovoid prostatic section (q), in which the high glandular cells have apical nuclei. Where it narrows, the nuclei become basal. The narrow part is longer than the wide one. It is followed by a shorter muscular efferent duct (d) which pierces the quite short penial papilla (p) in an almost straight course. The male atrium (t) opens on a papilla in front of the female pore (t). The inner oviduct enters the seminal receptacle (r) with a slrongly ciliated valve (w), and is continued into a looping vagina (v), whosl~opening lies quite near the nidamental aperture (t). The way of the eggs after insemination in the receptaculum is down the vagina and into the folded female gland mass, first through the albumen gland, farther on through the mucus gland, whence the eggs are expelled through the female aperture. Between the entrance of the albumen gland and the exit of the mucus gland there is a small pouch (b) which corresponds to the bursa copulatrix of Doto uva Marcus (1959: fig. 160, b). The six spawns coiled on one twig of Eudendrium conform to the figures of the spawn of Doto fragilis (Forbes, 1838) (Alder & Hancock, 1851: pt. 5, fam. 3, pI. 5, figs. 5, 6). The length of the folded ribbon is 1 em; unfolded it might be 2.5 em. It is 1.5 mm in breadth. The diameter of the embryos is 0.1 mm, and the number of eggs in one spawn estimated at about 5000. 1972] Marcus: Opisthobranchs from Florida 301 Discussion.-Three characters led me to the present classification: the 1.7-mm specimen was an undoubted Dolo chica by the shape of its cerata; the previous 4.5-mm animal of chica had had three lateral papillae; and the radular tooth corresponds exactly to that of chica (Marcus, 1960: fig. 60). Also the identity of the largest slug with material previously called Dolo fragilis umia (Marcus, 1969: 27) was quite evident. The latter we had determined from two individuals without cerata. D. fragilis seemed to have the closest likeness by its number of cerata, the dorsal papillae, and the rhinophoral sheaths. This classification cannot be main- tained, as the cerata of the present specimens do not agree with those of fragilis, which have more numerous rings of light-colored tubercles, figured as pointed on the plate of Alder & Hancock. The indication of lighter or darker tips of the tubercles is only of restricted value, as Odhner (1936: 1120) said "the black spot on the tubercles is in preserved specimens often hidden under the opaque epidermis." In the present material, the smallest animal has white tubercles, the larger ones have a few black granules in the tissue underlying the epithelium of the tubercular tips. Unfortunately these observations invalidate the basic distinction of Odhner's key (1936: 1119-1121). The shape and density of the tubercles are also extremely variable. In the specimens from Seahorse Key they are globose and from far apart (Fig. 28) to crowded (Fig. 29), what Kress calls grape- shaped cerata ("traubenartig," 1968b: 246). In the slugs from Boca Ciega they are pointed and far apart (Fig. 33), and the loose ceras from Boca Ciega is grape-shaped with globose tubercles (Fig. 32). Our first specimen of Dolo chica, 4.5 mm long preserved, had three lateral "tubercles, recalling those of D. pinnalifida" (Marcus, 1960). The later ones, 2 mm preserved, and 8 mm alive, had no papillae-I want to use the term papillae for the knobs on the body, so as not to confuse them with tubercles on the cerata. In the present material, only the 6-mm and lO-mm slugs from Seahorse Key have papillae. The 6-mm individual has three dorsal papillae, and the 10-mm one has dorsal and lateral rows and several on the head and the rhinophoral sheaths (Fig. 27). Eliot (1906: 357; 1910: 124-125) considered his variety splendida, 5 mm long, "with imperfect papillae" as the young form, and his var. papillifera, 10 mm long, which bears dorsal and lateral rows of papillae, as the normal adult form of Dolo pinnalifida (Montagu, 1804). Thus, I think, I am right to unite my specimens as young and adult of Dolo chica. Alder & Hancock (1851: pt. 5, fam. 3, pI. 5) indicated dorsal yellow tubercular spots for Dolo fragilis (Forbes, 1838). A row of papillae on the middle of the back and a lateral one on either side under the cerata is characteristic of D. pinnatifida. Odhner (1926b: 29) mentioned a 10-mm specimen of D. cuspidata Alder & Hancock, which had some small spherical verrucae on the middle part of the back. 302 Bulletin of Marine Science [22(2)

FIGURES 34-36. Doto chica Marcus: 34, 4-mm slug from Boca Ciega; 35, fourth right ceras of same in side view; 36, diagram of reproductive organs. Ca, ampulla; b, bursa copulatrix; d, efferent duct; t, female aperture; g, albumen gland; h, hermaphrodite duct; j, rhinophore; k, gill; I, velum; m, mucus gland; p, penial papilla; q, prostate; r, receptaculum seminis; t, male atrium; u, anal tube; v, vagina; w, valve.) 1972] Marcus: Opisthobranchs from Florida 303 O'Donoghue (1921: 204) said that the oral veil of Data columbiana O'Donoghue, 1921, bears a few short papillae. Such were not developed in the specimens we studied (Marcus, 1961a: 36). Dr. M. Patricia Morse kindly presented me with a specimen and a color slide labeled Data fragilis from Millport, Scotland. This well-preserved sample, 8.5 rum long, has in the photograph white globose tubercles apart from one another on seven pairs of bright red cerata. In preserved state they are slender blunt cones in four to six circles almost as far apart from one another as the height of the cones. This animal can be identified with Kress's description of D. fragilis (1968b: 246, pI. 2, fig. 2). The much less developed gills of the present slug might be due to its smaller size, as Kress's specimens were up to 40 mm long, alive. The animal from Millport does not have any dorsal papillae either, nor does Kress mention them in her description of fragilis, though she does for pinnatifida (p. 246). I do not want to risk a classification of this single animal. The description and figures of fragilis given by Kress (1968a: 246, figs. 1, 4c, 5c, 5d, pI. 2, fig. 2) differ far from the original ones by the shape of the cerata, and the number and shape of the tubercles. The size of the gills is much larger than in Odhner's figure (1936: fig. 46b). Neither Odhner nor Kress mentioned lateral papillae; Odhner's description (p. 1120) includes the tubercular yellow spots on the back indicated by Alder & Hancock in the original diagnosis. Baba (1971) described a new subspecies of Doto fragilis from Sagami Bay, D. fragilis nipponensis, 10-15 rum long, uniformly ashy yellow preserved, with seven pairs of cerata with four or five circlets of elongated conical tubercles each, and a series of bluntly conical papillae on the median line of the back and on each side of the body. His figure (1 A) of the right side shows 16 lateral papillae. All these characters differ from those of the present material. Resuming the indications in the literature, the external characters of the species of Doto vary considerably (Eliot, 1910: 123-125; Kress, 1968a: 246), e.g., the color (Odhner, 1936: 1120), the shape of the rhinophores (Marcus, 1967: 216), and the number and shape of the cerata. The reproductive organs are known for only a few of the species. Perhaps they will be the basis for a safer classification in future (see Marcus, 1961a: 41).

10. Trinehesia perea (Marcus, 1958) Referenees.-Marcus,1958b: 45, figs. 81-87 (Catriona perea); Edmunds, 1964: 4, figs. 1 C, D, 2 A, 3 B (Catriona perea); Scbmekel, 1968: 447 (Trinchesia perea) ; Edmunds, 1970: 18, 30 (Trinehesia perea). Material.-Florida: Key Biscayne, Crandon Marina, 28. X. 1969, two specimens. 304 Bulletin of Marine Science [22(2) Further Distribution.-J amaica; Brazil, Sao Paulo. Remarks.-The denticles of the masticatory border and the radular teeth correspond to Edmunds's and Schmekel's definition of Trinchesia. This is a new record for Florida.

11. Berghia verrucicornis (A. Costa, 1864) Synonym.-Berghia coeru/escens (Laurillard, 1830), part. References.-Marcus, 1957: 477, figs. 237-246; 1959: 258, figs. 16, 17; 1961b: 149 (B. coerulescens); Tardy, 1962: 4, figs. 1-3 (B. verrucicornis); Edmunds, 1968: 212, figs. 7-9 (B. verrucicornis). Material.-Florida: Grassy Key, 21. X. 1969, two specimens under a stone in upper tidal zone.

Further Distribution.-From Cape Hatteras to Brazil, Sao Paulo; Atlantic and Mediterranean coasts of Europe. Description.-The living slugs were 12 and 10 mm long; preserved they measure 10 and 9 mm, respectively. They have a very slender tail, 2 mm long, bent upwards, and tentaculiform foot corners. The animals were transparent greyish with an opaque white pattern along the middle of the back to the tip of the tail and on the cerata. The dark liver duct appears bluish under the white pigment, which is preserved in alcohol. Orange lines run transversely in front of the rhinophores, the bases of which are orange. Orange crescents accompany the apices of the horseshoe-shaped insertion lines of the cerata. On the cerata there is no orange ring. The rhinophores bear irregular spherical knobs on the hinder side; in front they are transversely wrinkled. The cerata, 50-55 on either side, are arranged in nine groups. In the anterior horseshoes they stand in irregular lines; the hindmost groups are simple rows. The first arch contains 14 cerata. The anal opening and the nephropore lie close together near the top of the second arch in both specimens. The jaws are half as broad, 530fL, as they are long, 1 mm. The masticatory process is narrow and 0.7 mm long; it is beset with about 350 hair- shaped denticles, 6-8fL high. The radula has 19 complete and one or two developing teeth with 30-38 denticles on either side of the small median cusp, on a convex widening. The oldest tooth is 90fL broad, the largest, 230fL. Remarks.- The specimens were compared with Edmunds's detailed analy- sis of his material from Jamaica and Ghana (1968: 213). Their color pattern comes nearest to his specimen B from Jamaica, except for the colorless rhinophore clubs in my slugs, and the orange ring on the cerata 1972] Marcus: Opisthobranehs from Florida 305 in the Jamaican animal. The broad white dorsal ornament occurs also in the European and the Brazilian material. The latter is more similar to the Jamaican slug by the red color on tentacles and rhinophores and on the cerata. The species is new for Florida.

SUMARIO

SOBRE ALGUNOS OPISTOBRANQUIOS DE LA FLORIDA Se estudian once especies de opistobranquios de la Florida. Bosellia mareusi y Trapania dalva sonespecies nuevas para la ciencia. Cylindro- bulla ulla Marcus, 1970, es la especie tipo del nuevo genera Aseobul/a y es nueva para Norte America. Se reportan en la Florida Ereolania funerea, Elysia ornata, E. elena, Diseodoris mortenseni, Trinehesia perea y Berghia verrllcieornis. Se discute Doto chiea. Varias especies han cambiado sus nombres debido a trabajos recientes de Baba y Hamatani, Habe, Tardy, Edmunds, y Marcus. Estas son: "Acteon" punctostriatus ahora Rietaxis punctostriatus, "Stiliger" funereus ahora Ercolania funerea, "Catriona" perea ahara Trinchesia perea y Berghia "eoerulea" ahora B. verrucieornis.

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