Review of the family Phyllidiidae in the Atlantic Ocean (Nudibranchia, Doridoidea) Angel Valdes1.2 and Jesus Ortea1 'DepartamentodeBiologi'adeOrganismosy Sistemas,LaboratoriodeZoologia, UniversidaddeOviedo,33071 Oviedo, Spain. 2LaboratoiredeBiologiedesInvertebresMarinsetMalacologie,MuseumNational d'HistoireNaturelle, 55,RuedeBuffon, 75005 Paris, France. Abstract: ThespeciesofPhyllidiidae fromthe AtlanticOcean(Mediterranean Seaexcluded)areredescribedinthispaper. Inlightofthe studyoftype materialandnewlycollectedspecimens,weconcludedthatPhyllidiopsisgynenoplaBouchet, 1977,isajuniorsynonymofP. berghiVayssiere, 1902. Also, anatomical studies supported the placement ofP. papilligera Bergh, 1890, in the genus Ceratophyllidia Eliot, 1903, and Phyllidiopsis molaensis Meyer, 1977,inthegenusPhyllidiellaBergh, 1869.Twonewbathyalspecies,Reticulidiagofasin.sp.andPhyllidiopsisbouchetin.sp.,aredescribed.Theformeris thefirstrecordofthisgenusintheAtlanticOcean. Key words: Nudibranchia,Phyllidiidae,taxonomy,newspecies,AtlanticOcean A review of the literature shows that few nominal MATERIAL AND METHODS species ofthe family Phyllidiidae, all lacking bright colors, Most ofthe specimens studied were collected during have been described in the Atlantic Ocean (Mediterranean several scientific expeditions {Z-Thalassa, Biacores, Sea excluded). This contrasts with the richly and brightly colored phyllidiid fauna of the Indo-Pacific recently re- Seamount I, and Seamount IP) organized by the Museum National d'Histoire Naturelle, Paris, France (MNHN) with viewed by Brunckhorst (1993). the vessels of the French agency for oceanographic The first report ofthis family in Atlantic waters was the description of Phyllidiopsis papilligera by Bergh research (IFREMER, Institut Francois de Recherche pour (1890), based on a single specimen collected at 182 m l'Exploitation de la Mer). Also,MtNyHpeNmaterial was exam- depth from the Gulf of Mexico, during the Blake ined from the collections of and the National Museum ofNatural History, Washington D. C. (USNM). Expedition. P. papilligera was later redescribed by Marcus and Marcus (1962, 1967) and Thompson (1980). Several Features of living specimens were recorded from original notes and drawings of collectors. Several speci- years later, Vayssiere (1902b) described Phyllidiopsis berghi, a species of uniform white color collected at 1480 mens have been dissected and particularly interesting soft m depth from the Bay of Biscay, by the Talisman parts have been critical point dried for scanning electron micrography (SEM). Expedition. In 1977, two new species ofPhyllidiopsis were reported from the Atlantic Ocean. One ofthem, P. gyneno- SPECIES DESCRIPTIONS pla Bouchet, 1977, was described on the basis of a single specimen collected at 525-600 m, near the Azores by the Genus Phyllidiella Bergh, 1869 Biacores Expedition. In the same paper, Bouchet (1977) Type species Phyllidia pustulosa Cuvier, 1804, by subse- redescribed P. berghi. The other species, P. molaensis, was quentdesignation (Brunckhorst, 1993). described by Meyer (1977) from shallow waters of the Caribbean coast ofPanama. Phyllidiella molaensis (Meyer, 1977) In the presentpaper, we review all the species report- (Figs. 1A, 2) ed in the Atlantic Ocean, and describe two new species. The Mediterranean species, Phyllidiaflava Aradas, 1847, PhyllidiopsismolaensisMeyer, 1977: 305-306, fig. 4. and Fryeria bayi Bouchet, 1983, have not been included because they were recently reviewed by Brunckhorst Material Examined (1993). USNM 760616, HOLOTYPE (by original designation), AmericanMalacologicalBulletin,Vol. 13(1/2)(1996):l-9 1 2 AMER. MALAC. BULL. 13(1/2) (1996) Fig. 1. Dorsal viewsofpreservedtypespecimens(scalebars= 5 mm). A.Phyllidiellamolaensis, holotype(USNM760616). B.Phyllidiopsisberghi, holo- type (MNHN). C. P. berghi, holotype ofPhyllidiopsisgynenopla (MNHN). D. Phyllidiopsisboucheti, holotype (MNHN). E. Ceratophyllidiapapilligera, neotype(USNM856418).F.Reticulidiagofasi. holotype(MNHN). Galeta Point, Panama, 15 m depth, 25 May 1971, 20 absent. mm preserved length, coll. Meyer. Ventrally, the oral tentacles are separated. The anteri- USNM 760617, PARATYPE, Portobelo, Panama, 9 m or and posterior ends of the foot are notched (Figs. 2B-C). mm depth, 10 September 1971, 1 spm, 19 preserved The branchial leaves are very thin andelongated, with alter- length, coll. Meyer. nating large and small ones (Fig. 2D). External Morphology Anatomy The color in life of this species was described by The moderately long oral tube dilates into a very Meyer (1977) as '...color black and white consisting of folded pharyngeal bulb (Fig. 2A). Two anterior retractor three sets of concentric circles along the sides, two more muscles insert at the point where the pharyngeal bulb and anteriorly and one setposteriorly, each setcomposed oftwo oral tube connect; another pair of muscles inserts in both concentric white rings each of which has a grey ring run- sides of the pharyngeal bulb. The esophagus is long, with ning through its middle and separated from one another by nomuscularorglandularportions. a black ring; white lines randomly traversing areas not cov- ered by the rings; rhinophores black with white tips...' In Geographical Range preserved specimens only brown shadows remain of the This species has only been recorded from the original black color (Fig. 1A). The dorsum bears simple, Caribbeancoast ofPanama. conical tubercles, decreasing in size toward the notal mar- gin. The mantle margin is as wide as the notum. The anus Discussion opens dorsally. Rhinotubercles (see Brunckhorst, 1993) are Anatomical features of this species resemble those VALDES AND ORTEA: PHYLLIDIIDAE IN THE ATLANTIC OCEAN 3 sta. 141 (45°59.00' N, 04°09.46' W), 1480 m depth, 30 mm August 1883, 16 preserved length, dissected. MNHN, HOLOTYPE ofP. gynenopla (by original designa- tion), Jean Charcot-Biacores sta. 159 (37°26.00' N, m 25°51.00' W), 525-600 depth, 31 October 1971, 48 mm preserved length, dissected. MNHN, Jean Charcot-Biacores sta. 59 (38°22.05' N, m 28°48.05' W), 560-580 depth, 14 October 1971, 2 mm B spms, 7 and9 preserved length. MNHN, Z-Thalassa sta. 435 (48°39.07' N, 09°53.02' W), m mm 1050 depth, 26 October 1973, 1 spm, 14 pre- served length, dissected. MNHN, SeamountIIsta. DW128, Gran Canada (28°08.30' Fig.2.Phyllidiella molaensis, paratype (USNM 760617). A. Dorsal view ofthe anatomy. B. Ventral view ofthe anterioredge ofthe foot showing the oral tentacles. C. Ventral view ofthe posterior edge ofthe foot. D. Detailofthebranchial leaves, dg,digestivegland; h, heart;i,intestine; o, esophagus; ot, oral tube; pb, pharyngeal bulb; r, retractormuscles; t, oral tentacles. described by Brunckhorst (1993) for the genus Phyllidiella, such as the structure ofthe digestive system with a moder- ately long oral tube which connects with a folded pharyn- geal bulb. Externally, this species has separate oral tenta- cles, simple tubercles on the dorsum, black rhinophores, and no rhinotubercles, a combination of characters only present in the species ofthis genus (Brunckhorst, 1993). Genus Phyllidiopsis Bergh, 1875 Type species Phyllidiopsis cardinalis Bergh, 1875, by monotypy. Phyllidiopsis berghiVayssiere, 1902 (Figs. 1B-C, 3, 4A-B, 4D) Fig. 3. Phyllidiopsis berghi. A. Dorsal view ofthe anatomy ofthe speci- men fromGranCanaria(MNHN). B. Detail ofthe anteriordigestivetract Phyllidiopsis berghi Vayssiere, 1902a: 1-2 (nomen nudum); dissected from the holotype of Phyllidiopsis gynenopla (MNHN). C. 1902b: 237-242, pis. 9-10; Bouchet, 1977: 48-49, figs. 16-17. Detail ofthe anterior digestive tract dissected from the holotype of P. Phyllidiopsis gynenopla Bouchet, 1977: 50-53, figs. 18-19, berghi (MNHN). D. Detail ofthe branchial leaves. E. Genital system. F. syn. nov. Ventral view ofpreserved holotype ofP. gynenopla showingtheoral ten- tacles, a, ampulla; bg, blood gland; dg, digestive gland; fg, female gland; Material Examined g, gametolytic gland; h, heart; i, intestine; o, esophagus; pb, pharyngeal bulb;pr, prostate;r, retractormuscles; sr, seminal receptacle;t,oral tenta- MNHN, HOLOTYPE ofP. berghi (by monotypy), Talisman cles;vg,vestibulargland. 4 AMER. MALAC. BULL. 13(1/2) (1996) Fig. 4. Scanning electron micrographs (SEM) using critical point drying technique. A. Pharyngeal bulb ofPhyllidiopsis berghi. B. Detail ofthe external glands ofthe pharyngeal bulbofP. berghi. C. Pharyngeal bulbofReticulidiagofasi. D. Muscularesophageal portionofdigestive tractofP. berghi. Scale bars= 1 mm(A), 100urn(B,D),200urn(C). N, 15°52.00' W), 470 m depth, 06 January 1993, 1 size (Fig. 3D). They are clustered very closely together. mm spm, 13 preserved length, dissected. MNHN, Seamount II sta. DW188, Hyeres Bank (3T30.00' Anatomy N, 28°5m9.m50' W), 310 m depth, 17 January 1993, 1 The pharyngeal bulb is very long (Figs. 3A, 4A) and spm, 7 preserved length. MNHN, Seamount II sta. DW200, Hyeres Bank (31°19.10' covered by numerousminute glands (Fig. 4B). Two anterior m retractor muscles insert at the point where the pharyngeal N, 28°3m6.m00' W), 1060 depth, 17 January 1993, 1 bulb andthe esophagus connect. The esophagus is also very spm, 8 preserved length. MNHN, Seamount II sta. DW241, Plato Bank (33°11.90' long, with a muscular portion near the end (Fig. 4D) where N, 28°50.30' W), 695 m depth, 31 January 1993, 2 the posterior retractor muscles are attached. The intestine mm presents several small tubercles on its anteriorportion. spms, 6 and9 preserved length. The genital system (Fig. 3E) has a vestibular gland. External Morphology The ampulla is smaller than the seminal receptacle in all specimens examined. From the gametolytic gland emerge The general body coloroflive animals is pale cream; three ducts, one ofthem connecting with the seminal recep- in larger specimens black pigment over several tubercles tacle, another withthe vagina, and the thinnestofthem with has been described (Bouchet, 1977). The dorsum bears the female gland. The prostatic portion ofthe deferent duct numerous rounded tubercles (Figs. 1B-C). Several larger tubercles are distributed among the others in adult speci- is long and folded. mens. The border of the mantle is narrow, nearly 1/10 of the notum width. The rhinophores are white. The anus Geographical Range opens dorsally in all specimens observed. Rhinotubercles This species, occurring in deep water, is only known are absent. in the North Atlantic (Fig. 5). It has been collected on the Ventrally, the edge of the mantle shows spicules in continental slope of France, in the Azores (slope of Sao reticular disposition. The anterior edge of the foot is not Jorge), on the seamounts of the Meteor Group (Hyeres notched and the oral tentacles are spherical and fused (Fig. Bank, Plato Bank), and in the Canary Islands (slope of 3F). The branchial leaves are wide, triangular, and equal in GranCanaria). VALDES AND ORTEA: PHYLLIDIIDAE IN THE ATLANTIC OCEAN 5 Phyllidiopsis boucheti new species (Figs. ID, 6) Material Examined MNHN, HOLOTYPE, Punta de la Rasca, Tenerife, Canary Islands, 400 m depth, 12 May 1988, 8 mm preserved length, leg. J. J. Bacallado; MNHN, PARATYPE, same mm locality, 1 spm, 38 preserved length, dissected, leg. J. J. Bacallado. External Morphology Both specimens were fixed, so that no external fea- tures ofthe living animals were available. The general body color is white in the preserved specimens (Fig. ID). The dorsum is entirely covered by numerous, minute, simply rounded tubercles. No gradation in size occurs in the tuber- cles near the notal margin. The rhinophores are white. Rhinotubercles are absent. The mantle margin is as wide as 1/3 ofthe notum. The anus opens dorsally in all specimens observed. Ventrally, the branchial leaves are triangular (Fig. Fig.5.DistributionmapoftheeasternAtlanticphyllidiidspecies. 6C). The oral tentacles are long and fused together (Fig. 6B). The mantle margin has spicules in reticular disposi- tion. The anterioredge ofthe foot is not notched. Anatomy Discussion The esophagus is very thin and long (Fig. 6A). It has Vayssiere (1902a) introduced the name Phyllidiopsis a muscular area in the middle of its length, just behind the berghi but did not give a description {nomen nudum). intestine. The pharynx is elongate, but approximately half Several months later, Vayssiere (1902b) gave a exhaustive the length ofthe esophagus. description ofthis species with anatomical details. The reproductive system (Fig. 6A) is bordered by the Phyllidiopsis gynenopla was described as different digestive gland. The smooth prostatic portion of the defer- from P. berghi by Bouchet (1977), on the basis of the dis- ent duct is twice as long as the ampulla, and leads into the tinctive external color pattern and few anatomical details. shortand narrow deferent duct. Observation of the digestive tract of the holotypes of both nominal species showed no significant differences (Figs. 3B-C). Also, the external morphology is identical among the specimens observed. The difference in the external color can be explained by the difference in size (more than 30 mm) between the holotype of P. gynenopla and other material examinedhere assigned to P. berghi. Another species with color pattern similar to Phyllidiopsis berghi is P. blanca Gosliner and Behrens, 1988, from the Pacific coast of North America. However, that species is distinguished by the disposition of the branchial leaves, all equal in size in P. berghi, and with alternation oflarge and small in P. blanca (see Gosliner and Behrens, 1988). Also, the disposition ofthe dorsal tubercles is different in the two species. In P. berghi, they are very close together, and there are two different sizes of tuber- cles; in P. blanca they are scattered and all ofthem are the 1mm same size. Anatomical differences between these species are found in the different disposition of the ducts arranged Fig. 6. Phyllidiopsis boucheti. paratype (MNHN). A. Dorsal view ofthe anatomy. B. Ventral view ofpreserved specimen showing the oral tenta- from the gametolytic gland (three in P. berghi and one in P. cles. C. Detail ofthe branchial leaves, dg, digestive gland; i, intestine; o, blanca). esophagus;pb,pharyngealbulb;r,retractormuscles;t,oraltentacles. 6 AMER. MALAC. BULL. 13(1/2) (1996) Geographical Range This species, occurring in deep water, is only known from the southern slope of Tenerife, Canary Islands (Fig. 5). Discussion Phyllidiopsis boucheti is clearly different from P. berghi. The species differ in the size, shape, and distribu- tion of the dorsal tubercles; they are very minute and even in size in P. boucheti, and larger, and of two different sizes in P. berghi. Also, ventrally, the shape and disposition of the branchial leaves are different in the two species. The differences between Phyllidiopsis boucheti and P. blanca are the shape and disposition of the dorsal tuber- cles, very minute and close together in the former and large and scattered in the latter. Also, differences in the shape and disposition of the branchial leaves support separation ofthe two species. Brunckhorst (1990b, 1993) has suggested that the nominal species Phyllidiopsis berghi and P. gynenopla, from the Atlantic Ocean, and P. blanca, from the Pacific coast of Northern America, should be included in a new genus, because the published descriptions make no mention of anterior retractor muscles and muscular segments in the esophagus of these species. Detailed anatomical studies of our material shows that Atlantic Phyllidiopsis do have ante- rior retractor muscles and also that there is a muscular por- tion in the esophagus as in the Indo-Pacific species. New Fig. 7. Ceratophyllidia papilligera. A. Dorsal view ofthe anatomy. B. anatomical studies ofP. blanca will probably also show the Genital system. C. Detail ofthe branchial leaves. D. Ventral viewofpre- presence of these features. Although Atlantic and Indo- served neotype (USNM 856418) showing the oral tentacles, a, ampulla; Pacific Phyllidiopsis show differences, such as the longer dg, digestive gland; g, gametolytic gland; h, heart; i, intestine; o, esopha- gus; pb, pharyngeal bulb; pr, prostate; r, retractor muscles; sg, salivary pharyngeal bulb in the Atlantic species, we feel that this is glands;sr,seminalreceptacle;t,oraltentacles. not enough to consider the separation of the Atlantic and Indo-Pacific Phyllidiopsis as two different genera. Material Examined Etymology USNM 856418, NEOTYPE, Sofia Expedition sta. 11, The name Phyllidiopsis boucheti is in honor of our (26°16.43' N, 83°46.49' W), 77 m depth, 30 April friend and colleague, Dr. Philippe Bouchet (MNHN), who 1981, 19 mm preserved length. has made important contributions to the knowledge ofmol- USNM 856417, Sofia Expedition sta. 11, (26°16.43' N, lusks. 83°46.49' W), 77 m depth, 30 April 1981, 2 spms, 10 mm and 12 preservedlength. m Indian Keys, Cuba, 30 October 1993, 30 depth, 1 spm, Genus Ceratophyllidia Eliot, 1903 26 mm long, leg. J. Espinosa. Type species Ceratophyllidia africana Eliot, 1903, by monotypy. External Morphology The general color ofthe body is white, with rounded Ceratophyllidiapapilligera (Bergh, 1890) black spots of various sizes (Fig. IE). The body is oval- (Figs. IE, 7) shaped, with numerous stalked papillae on the dorsum. The placement ofthe black spots is independent ofthe papillae; Phyllidiopsis papilligera Bergh, 1890: 176-178, pis. 2, 7- their size is variable but usually very large. The notum is 14; Marcus and Marcus, 1962: 475-479, figs. 20-24; 1967: spiculose. The rhinophores are white with black lamellae; 99, fig. 123; Thompson, 1980: 93, fig. 12. each clavus has ten lamellae. Rhinotubercles are absent. 1 VALDES AND ORTEA: PHYLLIDIIDAE IN THE ATLANTIC OCEAN 7 Ventrally, the anterior border of the foot is notched equal in size in C. papilligera), and the pattern of color (Fig. 7D). The dark tubercles of the notal margin show (black rings in P. molaensis and black spots in C. papilli- through the ventral side. The oral tentacles are separated. gera) easily separate the two species. The branchial leaves are equal in size (Fig. 7C). According to Thompson (1980), the type material of Ceratophyllidia papilligera is untraceable; however, for Anatomy nomenclatural stability of this species we considered it At the point where the esophagus inserts into the desirable to designate a neotype. For this reason, we desig- USNM musculoglandular pharyngeal bulb, it connects with two nated as neotype one specimen, 856418, collected conspicuous and brown-colored salivary glands (Fig. 7A). in the Gulf of Mexico, off Florida, close to the original The esophagus is short, and no glandular portion has been record ofthis species. observed. The genital system (Fig. 7B) shows an ampulla larg- er than the gametolytic gland. No vestibular gland has been Genus Reticulidia Brunckhorst, 1990 observed. The gametolytic gland connects with two ducts, Type species Reticulidia halgerda Brunckhorst and Burn, one of them connecting with the vagina, and the other one 1990, by original designation. with the seminal receptacle and the female gland. Reticulidiagofasi newspecies Geographical Range (Figs. IF, 4C, 8) This species appears to be restricted to the Caribbean Sea and GulfofMexico. After its original description from Material Examined the Gulf of Mexico (Bergh, 1890), Ceratophyllidia papil- MNHN, HOLOTYPE, Seamount I sta. DW61, Josephine ligera has been reported from Jamaica (Marcus and Bank (36°40.02' N, 14°16.00' W), 200-205 m depth, Marcus, 1967; Thompson, 1980) and from the Virgin mm 07 October 1987, 11 preserved length. Islands andBahamas (Marcus and Marcus, 1962). MNHN, PARATYPE, Seamount II sta. DW256, Atlantis Bank (34°06.20' N, 30°16.00' W), 340 m depth, 02 mm Discussion February 1993, 1 spm, 9 preserved length, dissect- Eliot (1903b) described the new genus ed. Ceratophyllidia and included Phyllidiopsis papilligera as MNHN, PARATYPE, Jean Charcot-Biacores sta. 1 the type species. In a later paper (Eliot, 1903a), he (38°30.00' N, 27°14.05' W), 76-105 m depth, 08 mm described the new species Ceratophyllidia africana, includ- October 1971, 1 spm, 10 preserved length, dissect- ing a description of the genus Ceratophyllidia. According ed. to Brunckhorst (1990b, 1993), the former paper was MNHN, Seamount II sta. DW274, Atlantis Bank received in June 1902 but was not published until sometime (34°05.10' N, 30°13.60' W), 280m depth, 05 February mm between July and November 1903. Eliot's description of C. 1993, 1 spm, 3 preserved length. africana appeared earlier (March 1903) and therefore con- stitutes the original description of the genus External Morphology Ceratophyllidia with C. africana as the type species by The general body color is pale yellow in live speci- monotypy. The anatomical characters observed in C. papil- mens. The dorsum is heavily spiculose, covered by conical ligera present all the features characteristic of the genus tubercles, grading to smaller toward the notal margin (Fig. Ceratophyllidia described by Eliot (1903a, b) and IF). The mantle margin is as wide as the notum, and Brunckhorst (1993) based on C. africana. The structure of translucent. The rhinophores are pale yellow. Rhino- the digestive system, with two conspicuous salivary glands, tubercles are absent. The anus opens dorsally in all speci- and the external stalked papillae of C. papilligera confirm mens observed. its placement in the genus Ceratophyllidia. Ventrally, the anterior edge of the foot is notched Phyllidiella molaensis shows an external combina- (Fig. 8D). The large and conical oral tentacles are separat- tion of colors similar to Ceratophyllidia papilligera. ed. The branchial leaves are very thin, and triangular, show- Nevertheless, anatomical features such as the structure of ing alternation oflarge and small ones (Fig. 8C). The man- the digestive system (typical ofthe genus Phyllidiella in P. tle margin shows radial spicules. molaensis) and external differences such as the shape ofthe dorsal tubercles (conical tubercles in P. molaensis and Anatomy stalked papillae in C. papilligera), the disposition of the The pharyngeal bulb is embedded on its anteriorpor- branchial leaves (showing alternation in P. molaensis and tion in the transparent oral tube; its edge is surrounded by 8 AMER. MALAC. BULL. 13(1/2) (1996) forthe genusReticulidia. However, the dorsal external mor- phology of the Indo-Pacific species of this genus, Reticulidia halgerda Brunckhorst and Burn, 1990, and Reticulidiafungia Brunckhorst and Gosliner, 1993, with bright colors and dorsal ridges, is very different from our species, pale yellow with conical tubercles. The color dif- ferences of R. gofasi with the other species of the genus could be connected with the bathyal habitat of the former; the same occurs for other deep water phyllidiids such as Fryeria bayi, Phyllidiopsis berghi, andR boucheti. In the external morphology and anatomy, Reticulidia gofasi is clearly different from the other Atlantic phyllidiid species. The morphology ofthe oral ten- tacles, the shape and distribution of the dorsal tubercles, and the structure ofthe digestive system are very distinctive characters ofthis species. Etymology The name Reticulidiagofasi is in honor ofour friend and colleague, Dr. Serge Gofas (MNHN), who collected and made available to us most of the material of this species. ACKNOWLEDGMENTS 0.5mm We are very grateful to Dr. Serge Gofas (MNHN), Dr. Philippe Fig. 8. Reticulidia gofasi, paratype from Atlantis Bank (MNHN). A. Bouchet (MNHN), Dr. Terrence Gosliner (California Academy of Dorsal viewofthe anatomy. B. Genital system. C. Detail ofthebranchial Sciences, San Francisco), and Gilianne Brodie (James Cook University, leaves. D. Ventral view ofpreservedspecimen showingtheoraltentacles, Australia) for constructive comments on the manuscript. We also a, ampulla; bg, blood gland; dg, digestive gland; g, gametolytic gland; h, acknowledge the valuble comments ofEditorDr. R. Toll andtwoanony- heart; i, intestine; o, esophagus; ot, oral tube; pb, pharyngeal bulb; pr, mous reviewers. Material studied has been provided by Dr. Alan Kabat prostate;r,retractormuscles;sr,seminalreceptacle;t,oraltentacles. (USNM), Dr. Juan J. Bacallado (Museo Insular de Ciencias Naturales, Tenerife, Spain), and Dr. Jose Espinosa (Instituto de Oceanologfa, Havana,Cuba). several discs, around a hole through which the short esoph- Dr. RudovonCosel (MNHN)producedthephotographsofspeci- mens forthispaper. TheassistanceofD. Guillaumin(CIME, Dept. MEB, agus emerges (Figs. 4C, 8A). Also, two large retractormus- UniversitedeParis-VI)fortheSEMphotosisgratefullyacknowledged. cles insert on the pharyngeal bulb at both sides of the esophagus. The genital system (Fig. 8B) presents an ampulla LITERATURE CITED larger than the seminal receptacle. The prostatic portion of the deferent duct is quite short, with only one fold, and con- Bergh,R. 1890.Reportontheresultsofdredging,underthesupervisionof nects with the deferent portion by a short and thin duct. Alexander Agassiz, in the Gulfof Mexico (1877-78), and in the Caribbean Sea (1879-80), by the U. S. Coast Survey Steamer Geographical Range "Blake", Lieut.-Commander C. D. Sigsbee, U. S. N., and Reticulidia gofasi is only known in the Meteor CommanderJ. R. Bartlett, U. S. N., Commanding. 32. Report on Group seamounts (Atlantis Bank), Lusitanian seamounts thenudibranchs. Bulletin oftheMuseum ofComparativeZoology 19(3):155-181. (Josephine Bank), and the Azores (slope of Terceira) (Fig. Bouchet, P. 1977. Opisthobranches de profondeurde l'Ocean Atlantique: 5). II, Notaspidea et Nudibranchiata. Journal ofMolluscan Studies 43:28-66. Discussion Brunckhorst,D.J. 1990a. Descriptionofanewgenusandspeciesbelong- The structure of the pharyngeal bulb of Reticulidia ing to the family Phyllidiidae (Nudibranchia, Doridoidea). JournalofMolluscanStudies56:567-576. gofasi, with several discs and two large retractormuscles, is Brunckhorst, D. J. 1990b. Description ofa new species ofPhyllidiopsis identical to that described by Brunckhorst (1990a, 1993) Bergh (Nudibranchia, Doridoidea, Phyllidiidae) from the tropical VALDES AND ORTEA: PHYLLIDIIDAE IN THE ATLANTIC OCEAN 9 western Pacific, with comments on the Atlantic species. Journal Marcus, Ev. and Er. Marcus. 1967. Tropical American opisthobrar.chs. ofMolluscanStudies56:577-584. Studiesin TropicalOceanography, Miami6:1-137. Brunckhorst, D. J. 1993. The systematics and phylogeny ofphyllidiid Meyer, K. B. 1977. Dorid nudibranchs ofthe Caribbean coast of the nudibranchs (Doridoidea). Records ofthe Australian Museum, Panama Canal Zone. Bulletin ofMarine Science ofthe Gulfand supplement 16:1-107. Caribbean27(2):299-207. Eliot, C. 1903a. On some nudibranchs fromEast Africaand Zanzibar, 2. Thompson, T. E. 1980. Jamaican opisthobranch molluscs II. Journal of ProceedingsoftheZoologicalSocietyofLondon 1903:250-255. MolluscanStudies46:74-99. Eliot, C. 1903b. Nudibranchiata, with some remarks on the families and Vayssiere, A. 1902a. Sur les opisthobranches recueillis en 1883 par genera and description ofa new genus, Doridomorpha. Fauna l'expeditiondu "Talisman." ComptesRendusHebdomadariesdes and Geography ofthe Maldive and Laccadive Archipelagoes SciencesetMemoriesde la Societede Biologie etde ses Filiales 2(l):540-573. 134:296-297. Gosliner,T.andD. Behrens. 1988. Areviewofthegenericdivisionswith- Vayssiere, A. 1902b. Opisthobranches du "Talisman" campagne de 1883. in the Phyllidiidae with the description of a new species of In: Expedition scientifique du "Travailleur" et du "Talisman" Phyllidiopsis (Nudibranchia: Phyllidiidae) from the Pacific coast pendant les annees 1880-1883, pp. 221-271. Ministere de ofNorthAmerica. The Veliger30(3):305-314. l'lnstructionPublique, Paris. Marcus, Ev. and Er. Marcus. 1962. Opisthobranchs from Floridaand the Virgin Islands. Bulletin ofMarine Science ofthe Gulfand Dateofmanuscriptacceptance: 11 October 1995 Caribbean 12(3):450-488.