BULLETIN OF MARINE SCIENCE, 66(2): 445–456, 2000 NEW SPECIES PAPER
A NEW SPECIES OF CYERCE BERGH, 1871 (MOLLUSCA, SACOGLOSSA, POLYBRANCHIIDAE) FROM THE PACIFIC COAST OF COSTA RICA
Ángel Valdés and Yolanda Camacho-García
ABSTRACT A new species of the genus Cyerce is described from the Pacific coast of Costa Rica, constituting the first record of the genus from the eastern Pacific. This new species, Cyerce orteai, is clearly distinguishable from other species of the genus, by both its anatomy and external morphology. Cyerce orteai has very elongate radular teeth, with two differ- entiated sets of denticles. Externally, this species is characterized by having numerous cerata bearing small rounded tubercles. The digestive and reproductive systems of this species are described anatomically.
Recently, sacoglossans with bifid rhinophores and flat, leaf-like dorsal cerata with marginal defensive glands have been placed in the family Polybranchiidae (Jensen, 1996; 1998). This family includes five valid genera: Polybranchia Pease, 1860, Caliphylla A. Costa, 1867, Cyerce Bergh, 1871, Mourgona Ev. Marcus and Er. Marcus, 1970 and Sohgenia Hamatani, 1991. Until now, Polybranchia viridis (Deshayes, 1857) is the only member of the family Polybranchiidae reported from the Pacific coast of the Americas (Bertsch and Smith, 1973). This paper describes a second species of this family, based on several specimens collected from the Pacific coast of Costa Rica, belonging to the genus Cyerce Bergh, 1871. The material studied is deposited at the Instituto Nacional de Biodiversidad de Costa Rica (INBio) and the California Academy of Sciences, San Francisco (CASIZ).
SPECIES DESCRIPTION
Family Polybranchiidae O’Donoghue, 1929 Genus Cyerce Bergh, 1871 Type species.—Cyerce elegans Bergh, 1871, by subsequent designation by Swennen (1961).
Cyerce orteai new species (Figs. 1–5)
Type Material.—HOLOTYPE.—Playa Cabuya, Cabuya, Cóbano, Puntarenas, Costa Rica, 29 January 1999, 13 mm preserved length (INBio CRI001500521). PARATYPES.—Playa Cabuya, Cabuya, Cóbano, Puntarenas, Costa Rica, 29 January 1999, three specimens 8–16 mm preserved length (INBio CRI001500644); two specimens 12–14 mm preserved length (CASIZ 115230). Estación San Miguel, Reserva Natural Absoluta de Cabo Blanco (ACT), Cabuya, Cóbano, Puntarenas, Costa Rica, 25 January 1999, one specimen 9 mm preserved length (INBio CRI001500646); three specimens 7–14 mm preserved length (INBio CRI001500647). Playa Ocal del Peñon, Santa Teresa, Cóbano, Puntarenas, Costa Rica, 29 January 1999, two specimens 13–18 mm preserved length (INBio CRI001500648).
445 446 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000
Figure 1. Photographs of living animals: A, Adult specimen, paratype (INBio CRI001500647); B, Juvenile specimen, paratype (INBio CRI001500644).
External Morphology.—The body is oval, flattened, wider in the middle region (Figs. 1,2A). The head is short and wide (Fig. 2B), having two short triangular lateral prolonga- tions. Dorsally, the head has two long, enrolled, bifurcated rhinophores, and two oral tentacles. The rhinophores and oral tentacles point forward in the living animal. They are covered with numerous, small tubercles. The central part of the body is somewhat de- pressed, the pericardium being elevated over the rest of the dorsum. There are two main posterior hepatic channels, ramified into smaller, secondary branches that connect into the cerata, without extending into them. Five smaller hepatic branches emerge antero- laterally, three on the right side and two on the left side, in all specimens examined (Fig. 3A–B). Some of the secondary branches are ramified and joined again to form a network. The cerata are arranged dorso-laterally and may total up to 98 in the larger specimens. VALDÉS AND CAMACHO-GARCÍA: NEW SPECIES OF CYERCE FROM COSTA RICA 447
Figure 2. External morphology: A, Dorsal view of a living paratype (INBio CRI001500646), scale bar = 5 mm; B, Detail of the head region in a preserved paratype (INBio CRI001500648), scale bar = 2 mm; C, Ventral view of anterior region of the foot of a paratype (CASIZ 115230), scale bar = 1 mm. Abbreviations: a = anal papillae, f1 = anterior region of the foot, f2 = posterior region of the foot, h = head, hb = hepatic branch, ot = oral tentacle, pc = pericardium, r = rhinophore.
The cerata are elongate, wider near their apexes, and are connected into the body by a short stalk. They are thick and covered with small rounded tubercles. The anal papillae is situated antero-laterally on the right side, just anterior to the peri- cardium. The tail is short and triangular, and visible when the animal is in motion. Ven- trally, the foot is divided into two regions by a transverse groove (Fig. 2C). The general color of the body is pale cream with a number of irregular olive green speckles and some brown spots, that give the body a pale green appearance. The pericar- dium and the anal papillae are opaque white. In some specimens the pericardium may have some olive green pigment. The head is dark brown with opaque white spots and two large cream areas around the eyes. The rhinophores and oral tentacles are the same color as the head. The tubercles covering the rhinophores and tentacles are pale cream or opaque white. 448 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000
Figure 3. Dorsal vessels of preserved specimens: A, Paratype (INBio CRI001500648); B, Paratype (CASIZ 115230).
The cerata are dark brown in the larger specimens (Fig. 1A) and pale green in the smaller ones (Fig. 1B). The tubercles are translucent white to cream, with an opaque white spot on the tip. The base of each ceras is reddish. The edge of each ceras is sur- rounded by small, opaque white spots, that could be defensive glands. Ventrally, the foot is translucent white with opaque white spots. The anterior, differentiated region of the foot has a reddish labium at its anterior edge. Anatomy.—The radular formula is 10 × 0.1.0 in one of the specimens examined (CASIZ 115230). The radula consists of a single series of 10 teeth, six in the descending limb and four in the ascending limb (Fig. 4A). The teeth are very elongated, having several conical denticles on each side, all of them oriented towards the base of the teeth. There are 16–17 denticles on the descending limb teeth (Fig. 4B) and 13–14 on the ascending limb teeth (Fig. 4C). In addition, there are five to six other denticles, on each side, near the cusp of each tooth. These denticles, clearly different from the others, are curved backwards (Fig. 4D). The teeth have two different regions, one of them bears the denticles and the other attaches the teeth to the odontophore. Both regions are separated basally by a short basal prolongation (Fig. 4B). The descending limb ends at the ascus sac that contains a number of small, preradular teeth (Fig. 4E). The oral glands consist of a dense mass surrounding the pharynx anteriorly (Fig. 5A,D). The pharynx is muscular and oval. There are a pair of retractor muscles connected later- ally to the pharynx. The central nervous system is situated over the esophagus. The esopha- gus opens dorsally into the pharynx. The esophagus is a long duct that branches at 1/3 of its length into the blue esophageal pouch (Fig. 5A). The elongated, whitish pharyngeal pouch is approximately twice the length of the pharynx. The two salivary vesicles join the esophagus, each of them connected to a salivary gland through a long separate duct (Fig. 5B). There are two esophageal glands that connect to the esophagus directly behind the salivary vesicles. The long intestine runs forward and ends at the anal papilla. The large digestive gland represents approximately 85% of the viscera. It is greenish and extends to the sides of the body. VALDÉS AND CAMACHO-GARCÍA: NEW SPECIES OF CYERCE FROM COSTA RICA 449
Figure 4. Radular morphology of a paratype (CASIZ 115230): A, Complete radula, scale bar = 430 μm; B, First tooth of the descending limb, scale bar = 150 μm; C, Teeth of the ascending limb, scale bar = 200 μm; D, Detail of the denticles of a descending limb tooth, scale bar = 100 μm; E, Ascus sac, scale bar = 100 μm.
The reproductive system is diaulic (Fig. 5C). The ovotestis consists of five follicles. The ducts from the follicles join together and lead to the hermaphrodite ampulla which narrows before dividing into the male and female ducts. The fertilization region is indi- cated by the confluence of the female and albumen ducts. There is a short duct that leads to a genital receptacle and the vagina. The vagina opens to the left body wall. The albu- men gland is composed of many branches of follicles and covers the reproductive system. 450 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000
Figure 5. Anatomy of a paratype (INBio CRI001500644): A, Dorsal view of the anatomy, scale bar = 1 mm; B, Detail of the anterior region of the esophagus and salivary glands, scale bar = 0.5 mm; C, Reproductive system, scale bar = 1 mm; D, Anterior region of the digestive system, scale bar = 0.5 mm. Abbreviations: ag = albumen gland, am = ampulla, dg = digestive gland, e = esophagus, eg = esophageal gland, ep = esophageal pouch, fl = follicle, gr = genital receptacle, i = intestine, m = retractor muscle, mg = membrane gland, o = oral glands, p = penis, ph = pharynx, pp = pharyngeal pouch, pr = prostate, sg = salivary gland, sv = salivary vesicle, v = vagina.
The vas deferens and the penis are located anteriorly. The penis contains a long copula- tory spine. The tip of the penis is triangular. There are two long, convoluted prostates. A bursa copulatrix is not present. Etymology.—This species is named for our friend and mentor Jesús Ortea, who intro- duced us to the study of opisthobranch mollusks.
DISCUSSION
Phylogenies of the Sacoglossa by Jensen (1996) and Mikkelsen (1998) have shown the Polybranchiidae to be paraphyletic. Jensen (1996, 1998) stated that the monophyly of the group can be easily “forced” by applying extra weight to a few characters. Unfortunately, weighting these characters results in lower resolution at the bases of the non-shelled clades of the Sacoglossa. Until this question is definitively solved, with a more detailed phyloge- VALDÉS AND CAMACHO-GARCÍA: NEW SPECIES OF CYERCE FROM COSTA RICA 451 netic analysis of the polybranchiids and related genera, we will follow Jensen’s classifica- tion and maintain the usage of the family Polybranchiidae. According to Jensen (1998), the genus Cyerce is characterized by having numerous flat, leaf-like cerata without tubules of the digestive gland, bifid rhinophores and short, rolled oral tentacles. The anal papillae is situated antero-dorsally, there are dorsal vessels entering the pericardium, and the foot sole is divided by a transverse groove. Internally, species of Cyerce have a dense digestive gland and the penis armed with a copulatory spine. C. orteai has all of these external and internal features and clearly belongs in this genus. The genus Cyerce comprises eight valid species distributed throughout tropical or sub- tropical seas, including the Caribbean, Mediterranean, Western Atlantic and tropical Indo- Pacific. C. orteai is clearly distinguishable from other species of the genus. The main distinctive features of this species are the presence of numerous tubercles on the cerata, rhinophores and oral tentacles, and the long and slender radular teeth, with two different sets of denticles. This combination of characters is not present in any other species of the genus (see Table 1). The western Pacific species Cyerce nigricans (Pease, 1866), from the tropical Indo- West Pacific and Indian Ocean, and Cyerce kikutarobabai Hamatani, 1976, from Japan, have very dark cerata with whitish-orange spots or lines, and a submarginal orange band (Pease, 1866; Hamatani, 1976). Also, the head of these two species is very dark with orange spots or lines. In contrast, C. orteai has dark brown to pale green cerata, orna- mented with translucent white or cream tubercles. The radula of C. kikutarobabai has long and strong teeth with more than 25 large denticles on each side. This is different from C. orteai that has thinner teeth with less and proportionally smaller denticles. The radula of C. nigricans (described by Bergh, 1888, as Cyerce pavonina Bergh, 1888) is composed of very long, slender teeth, with conical denticles, very similar to those of C. orteai. Another Indo-Pacific species, Cyerce elegans Bergh, 1871, was described from Papua New Guinea. C. elegans is characterized by having somewhat transparent, beige cerata, surrounded by a cream line (Bergh, 1871; 1888). From this line several thin, pale cream lines emerge, and join together in the center of the cerata. There is also a dark patch near the base of each ceras. This is very different from the cerata of C. orteai that have several lighter tubercles, but lack lines. The Caribbean species Cyerce antillensis Engel, 1927 and Cyerce habanensis Ortea and Templado, 1988, are externally very similar. The former was redescribed by Ev. Marcus and Er. Marcus (1963), who found only a few large cerata on each side, and numerous smaller ones in between them. The cerata are situated laterally, so the dorsum is visible in the living animal. They are almost transparent often with a dark base and a dark spot near the center. Cyerce habanensis has exactly the same external morphology (see Ortea and Templado, 1988). In contrast, C. orteai has up to 98 large cerata in the larger specimens, that are arranged dorso-laterally, so the dorsum is hidden by cerata in the living animal. Also, the cerata of C. orteai are darker (in the adult specimens), with several cream tu- bercles that are absent in both C. antillensis and C. habanensis. The radular morphology of C. antillensis and C. habanensis is also very different from C. orteai. The teeth of the Caribbean species are shorter and broader than those of C. orteai. Also, in C. antillensis and C. habanensis the teeth have more, smaller denticles than in C. orteai, and all of them have a similar morphology. 452 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000
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C VALDÉS AND CAMACHO-GARCÍA: NEW SPECIES OF CYERCE FROM COSTA RICA 455
Eastern Atlantic species of the genus include the Mediterranean Cyerce cristallina (Trinchese, 1881) and Cyerce graeca Thompson, 1988, the former also reported from the Caribbean Sea (Ev. Marcus and Er. Marcus, 1970). Schmekel and Portmann (1982) and Thompson (1988) redescribed C. cristallina which has large, very elongate, almost trans- parent cerata with red-brown markings and orange-red defensive glands in the apex. These cerata are diamond-like, narrower near the base and the apex. In contrast, the cerata of C. orteai are always wider near the apex. The radular teeth of C. cristallina are also very elongate, but they do not have two differentiated sets of denticles, as in C. orteai (see Schmekel and Portmann, 1982; Thompson, 1988). The body shape of both species also appears to be different. Cyerce cristallina is a larger, more elongate species, with the pericardium situated very anteriorly and visible in the living animal. Cyerce graeca is very easily differentiated from C. orteai, by its oval cerata and short radular teeth (see Thompson, 1988). The Cape Verde Islands species, Cyerce verdensis Ortea and Templado, 1990, is similar to C. orteai in the shape of the cerata and in having papillae, or tubercles, on the cerata and rhinophores (Ortea and Templado, 1990). However, the tubercles of C. verdensis are smaller than those of C. orteai. The main difference between these two species is the radular morphology. The teeth of C. verdensis are much shorter and have fewer denticles than those of C. orteai. P. viridis (Deshayes, 1857) is the only member of the family Polybranchiidae reported from the Pacific Coast of the Americas. Bertsch and Smith (1973) redescribed this spe- cies based on specimens from Mexico. The main difference between C. orteai and P. viridis is that the highly branched tubule of the digestive system enters each ceras in the later. The radular teeth of P. viridis, described by Ferreira and Bertsch (1975) are much shorter than those of C. orteai. Also, in the former species the teeth have 14 short, square denticles, very different from the conical denticles of C. orteai. The diet of C. orteai remains unknown. All the specimens were collected under rocks, never in the immediate proximity of algae.
ACKNOWLEDGMENTS
We are very grateful to S. Ávila, F. Alvarado, A. Berrocal, F. Maramotti, A. Alvarado, K. Quesada, M. Calderón and S. Aucoin, who assisted during the field work. S. Ávila and F. Alvarado also helped in several other technical aspects of this paper. R. Johnson made constructive comments on the manuscript. The SEM work was carried out in the facilities of the Unidad de Microscopía Electrónica (Universidad de Costa Rica) and the California Academy of Sciences. The Reserva Absoluta de Cabo Blanco (ACT) provided logistic support during the field work. This paper and the field work was founded in part by INBio (through the Netherlands Development Assistance pro- grams), the California Academy of Sciences, and the SEUI of the Ministerio de Educación y Cultura of Spain.
LITERATURE CITED
Bergh, R. 1871. Malacologische Untersuchungen, Band 2. Pages 49–118 in C. Semper, ed. Reisen im Archipel der Philippinen, pls 9–16. ______. 1888. Malacologische Untersuchungen, Band 3. Pages 755–814 in C. Semper, ed. Reisen im Archipel der Philippinen, pls 77–81. Bertsch, H. and A. A. Smith. 1973. Observations on three opisthobranchs (Mollusca: Gastropoda) of the La Paz area, Baja California, Mexico. Southwest. Nat. 18: 165–176. 456 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000
Ferreira, A. J. and H. Bertsch. 1975. Anatomical and distributional observations of some opistho- branchs from the Panamic faunal Province. Veliger 17: 323–330. Hamatani, I. 1976. A new species of Cyerce Bergh, 1871, C. kikutarobabai, from Yoron Island (Opisthobranchia: Sacoglossa). Publs. Seto Mar. Biol. Lab. 23: 283–288, pl. 1. Jensen, K. R. 1996. Phylogenetic systematics and classification of the Sacoglossa (Mollusca, Gas- tropoda, Opisthobranchia). Phil. Trans. R. Soc. London B 351: 91–122. ______. 1998. Sacoglossernes systematik, fylogeni og evolution (Mollusca, Opisthobranchia). Ph.D. Thesis, Zoologisk Museum, Copenhaguen. 71 p + 65 p. Marcus, Ev. and Er. Marcus. 1963. Opistobranchs from the Lesser Antilles. Stud. Fauna Curaçao 79: 1–76. ______and ______. 1970. Opisthobranchs from Curaçao and faunistacally related re- gions. Stud. Fauna Curaçao 122: 1–129. Mikkelsen, P. M. 1998. Cylindrobulla and Ascobulla in the western Atlantic (Gastropoda, Opisthobranchia, Sacoglossa): Systematic review, description of a new species, and phyloge- netic reanalysis. Zool. Script. 27: 49–71. Ortea, J. and J. Templado. 1988. Una nueva especie de Cyerce Bergh, 1871(Opisthobranchia: Ascoglossa) de la Isla de Cuba. Iberus 8: 11–14. ______. 1990. A new species of the genus Cyerce Bergh, 1871, from the Cape Verde Islands (Opisthobranchia: Ascoglossa). Veliger 33: 202–205. Pease, W. H. 1866. Remarks on Nudibranchiata inhabiting the Pacific Islands, with descriptions of two new genera. Amer. J. Conchol. 2: 204–208. Schmekel, L. and A. Portmann. 1982. Opisthobranchia des Mittelmeeres. Nudibranchia und Saccoglossa. Springer-Verlag, Berlin. 410 p. Swennen, C. 1961. On a collection of Opisthobranchia from Turkey. Zool. Meded. 38: 41–75. Thompson, T. E. 1988. Eastern Mediterranean Opisthobranchia: Oxynoidae, Polybranchiidae, Stiligeridae (Sacoglossa). J. Moll. Stud. 54: 157–172.
DATE SUBMITTED: May 14, 1999. DATE ACCEPTED: November 2, 1999.
ADDRESSES: (Á.V.) Department of Invertebrate Zoology and Geology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118. (Y.C.-G.) Instituto Nacional de Biodiversidad (INBio), Sub-Programa de Malacología, Apdo. 22-3100, Santo Domingo de Heredia, Costa Rica.
ADDENDUM
While this paper was in press, we have received another specimen of Cyerce orteai collected from Napili Bay, Maui, Hawaii (CASIZ 118799). The specimen measures 5 mm preserved length, and was collected at 4 m depth by Cory Pittman. In addition, Scott Johnson provided us with a picture of another, larger specimen from Oahu, Hawaii. Thus, the geographic range of Cyerce orteai would be much larger than originally thought.