<<

Caribbean Journal of Science, Vol. 41, No. 3, 600-612, 2005 Copyright 2005 College of Arts and Sciences University of Puerto Rico, Mayagu¨ez

Ascidians from Bocas del Toro, Panama. I. Biodiversity.

ROSANA M. ROCHA*, SUZANA B. FARIA AND TATIANE R. MORENO

Universidade Federal do Paraná, Departamento de Zoologia, CP 19020, 81.531-980, Curitiba, Paraná, Brazil Corresponding author: *[email protected]

ABSTRACT.—An intensive survey of ascidian species was carried out in August 2003, in different environ- ments of the Bocas del Toro region, northwestern Panama. Most samples are from shallow waters (< 3 m) in coral reefs, among mangrove roots and in Thallasia testudines banks. Of the 58 species found, 14 are new species. Of the 26 sampling sites, the most diverse were Crawl Key Canal (9°15.050’N, 82°07.631’W), Solarte ,Island (9°17.929’N, 82°11.672’W), Wild Cane Key (9° 20؅40؅N, 82°10؅20؅W), Isla Pastores (9°14.332’N W), the bay behind the STRI Lab (9°21؅4.3؅ N, 82°15’25.6؅ W), and the entrance of Bocatorito Bay’82°19.968 (9°13.375’N, 82°12.555’W). Ascidians have been studied and reported from 31 locations within the Caribbean, and 139 species have been reported. This count will increase with the description of the 14 new species, and Bocas del Toro may be considered a region of high ascidian diversity since > 40% of the total known Caribbean ascidian fauna occurs there.

KEYWORDS.—Ascidian , Caribbean, checklist

INTRODUCTION found and discuss the relationship of this fauna with that of other Caribbean loca- tions. Warm Caribbean waters contain a wide variety of marine environments including different types of mangroves, shallow la- MATERIALS AND METHODS goons, open sea, shallow and deep coral reefs and rocky shores distributed among the islands and coasts of Central and South In the western Caribbean, the Bocas del America. Ascidians form a diverse yet Toro archipelago, on the northwestern poorly studied group in these Caribbean shore of Panama, is comprised of more waters. The most recent listing of ascidian than 68 islands and numerous mangrove species of the Caribbean was based on the keys. Ascidians were collected in this re- following studies: Plough and Jones (1939), gion between 03 and 12 August 2003. Habi- Van Name (1921, 1924, 1930, 1945), Millar tats surveyed included mangrove roots, (1962), Van der Sloot (1969), Millar and shallow coral reefs or coral formations on Goodbody (1974), Lafargue and Duclaux sandy substrates, and very shallow sea- (1979), Monniot, C. (1983a, b, c), Monniot grass meadows of Thalassia testudinum on and Monniot (1984), Monniot, F. (1983a, b, several of the islands of the archipelago c, 1984), Goodbody (1984a, 1984b,1993, (Fig.1, see list of stations in appendix). Most 1995, 1996, 2000, 2003), Goodbody and Cole of the locations surveyed were very shal- (1987) and Hernandez-Zanuy (1990). low (< 3 m) and sampled while snorkeling, In August 2003, the Smithsonian Tropical while others were sampled in deeper wa- Research Institute hosted a workshop to ters using SCUBA equipment. study the Bocas del Toro region, western were collected with their substrate (usually Panama. This region has a surprisingly rich bivalve shells on mangrove roots and small ascidian fauna and several new species pieces of dead coral on coral reefs) to insure were discovered, which will be described better anesthesia. Upon collection, samples elsewhere. Here we report the species were relaxed with menthol crystals after 600 ASCIDIAN BIODIVERSITY 601

FIG. 1. Bocas del Toro region showing diving sites (1-26). which they were fixed and preserved in same effort but each location was surveyed 10% seawater formalin. for a minimum of one hour and most sam- Most species were photographed prior to pling locations had few species (1-4). A sampling with a Canon Powershot 45 digi- quadratic regression of the species accumu- tal camera with underwater housing (See lation curve, based on the number of sites Photographic Identification Guide in this sampled, resulted in a significant model (R2 issue). = 0.97, p < 0.05). This model predicts that with 26 sites the curve reaches its asymp- RESULTS tote, and suggests that of the sites sampled, the ascidian community comprises ap- Forty-four known species and another 14 proximately 58 species (Fig. 2). new species were found during the survey. The most species-rich genera were Pyura Locations were not all surveyed with the (7 species) and (6 species, Table 1). 602 R. M. ROCHA ET AL.

depth, cover of algae and other inverte- brates.

DISCUSSION

At least for shallow-water ascidian com- munities in both mangroves and coral en- FIG. 2. The cumulative number of species across vironments, this survey found more species diving sites. The number of each site according to in Bocas than has been recorded in any Table 1 and Fig. 1. other site in the Caribbean other than Guadeloupe. Some identifications in Table 1 are tentative because SEM photographs of The most species-rich site was Crawl Key the spicules are not yet available and these Canal (18 species, 9°15.050’N, 82°07.631’W) are important for species’ identification. between Bastimentos and Popa islands We also believe that collecting in deeper (Table 1). This channel is subject to tidal environments will further increase the currents and links the more protected bay number of ascidians in the area, e.g., with the open sea. Here, sampling took sp. and turbinata were place on the coral reef that extends over a only found on deeper sites. The didemnids large part of the canal, including shallow to also need further study in Bocas del Toro deep locations. The second richest site, with since most colonies did not contain mature 14 species, was Solarte Island (9°17.929’N, gonads or larvae needed for identification. 82°11.672’W), a well conserved mangrove A comprehensive literature survey of the site situated inside a Conservation Unit. last 50 years showed that ascidians have Other rich sites were Wild Cane Key been recorded from 31 locations in the Car- (9°20Ј40ЈN, 82°10Ј 20ЈW, 10 species), Isla ibbean and Bermuda (Table 2, we included Pastores (9°14.332’N, 82°19.968’W, 9 spe- Bermuda since that ascidian fauna is simi- cies), the bay at the STRI Lab (9°21Ј 4.3Ј N, lar to that of the Caribbean). Most of the 82°15Ј 25.6Ј W, 10 species), and the entrance localities were not intensively surveyed of Bocatorito Bay (9°13.375’N, 82°12.555’W, and very few ascidians are known from 10 species). The most commonly found spe- them. A hundred thirty-eight species are cies were Microcosmus exasperatus (found in reported, which is a low figure compared 8 dive sites), sp., ab- with the survey of the Iberic Mediterranean dominalis and nigra (in 5 dive sites, with 101 species (Ramos-Esplá 1988), or the Table 1). monographs by Kott from Australia with Some species were exclusively found hundreds of species (Kott 1985, 1990, 1992, on mangroves, on corals or on Thalassia 2001). leaves (Table 1), suggesting a relationship In spite of the statement by Goodbody between diversity of environments and (1984) that “the ascidian fauna of the Car- high species diversity. Even the same type ibbean is well known” it appears that of environment was not uniform in the many new species of Caribbean ascidians whole area. Coral reefs, for instance, at are still to be described when more detailed times were comprised of mostly Porites and sampling is performed in different islands. other times Millepora or Acropora. Coral It emerges that the most diverse locations reefs in different locations, such as inside are Guadeloupe, Belize, Jamaica and Ber- the bay in very protected areas (Pastores muda which were surveyed more than island) or in channels close to the open sea once and/or by specialists using diving to (Crawl Key Canal), had very different as- collect animals (Van Name 1945; Monniot cidian communities. This was also true for C. 1972, 1983a, b, c; Monniot F. 1972, 1983a, b, mangrove communities and was appar- c, 1984; Monniot and Monniot 1984, Good- ently influenced by the age of the roots, body 2000, 2003). Most of the surveys made location relative to water currents, water before these depended on low tide or were TABLE 1. Distribution of ascidian species at the diving sites surveyed in Bocas del Toro, Panamá (for a description of the sites see Collin this issue and Fig. 1).

Diving site number # § 1 2 3 4 5 6 7 8 91011121314151617181920212223242526 Number of species 3 6 10 0 0 2 18 2 3 54234400914010051051 Polyclinum sp. M 1 X Euherdmania sp. 1 C 1 X Euherdmania sp. 2 C 1 X Didemnidae Didemnum conchyliatum ? (Sluiter, 1898) M 1 X D. granulatum ? Tokioka, 1954 C,T 3 X X X 603 BIODIVERSITY ASCIDIAN D. ligulum ? Monniot, 1983a C 1 X D. psammathodes (Sluiter, 1895) M 1 X D. speciosum ? (Herdman, 1886) C 1 X Trididemnum orbiculatum (Van Name, 1902) M 1 X T. savignyi (Herdman, 1886) C 1 X T. maragogi Rocha, 2002 T 1 X Lissoclinum verrilli (Van Name, 1902) C 1 X Diplosoma listerianum (Milne-Edwards, 1841) C 3 X X X Clavelinidae Clavelina sp. C 2 X X Holozoidae Distaplia bermudensis Van Name, 1902 1 X D. corolla Monniot, 1974 C 1 X Polycitoridae Cystodytes dellechiajei (Della Valle, 1877) C 3 X X X Cystodytes sp. 1 C 1 X Cystodytes cf. roseolus C1 X Eudistoma carolinense Van Name, 1945 C 2 X X E. olivaceum (Van Name, 1902) C,M 3 X XX E. obscuratum (Van Name, 1902) C 1 X Eudistoma clarum (Van Name, 1902) C 3 X X X Eudistoma sp. C 5 X X X X X 604

TABLE 1. Continued.

Diving site number # § 12 3 4567891011121314151617181920212223242526 0 Rhopalaea abdominalis (Sluiter, 1898) C 5 X X X X X Rhodosoma turcicum (Savigny, 1816) M 1 X multiclathrata (Sluiter, 1904) T 1 X Ecteinascidia styeloides (Traustedt, 1882) M 3 X X X Herdman, 1880 C 2 X X AL. ET ROCHA M. R. Ascidia curvata (Traustedt, 1882) M 4 X X X X Ascidia interrupta Heller, 1878 M 3 X XX A. sydneiensis Stimpson, 1855 C,M 2 XX Ascidia sp. 1 C,M 3 XX X Ascidia sp. 2 C,M 2 X X Ascidia sp. 3 M 1 X Phallusia nigra Savigny, 1816 C,M 5 X X X X X Botryllus planus (Van Name, 1902) C,T 2 X X B. tuberatus Ritter & Forsyth, 1917 T 1 X Symplegma brakenhielmi (Michaelsen, 1904) M 3 XX X S. rubra Monniot, 1972 M 1 X Styela canopus Savigny, 1816 C,M 4 X X X X ? S. plicata (Lesueur, 1823) C 1 X? X Polycarpa cartilaginea (Sluiter, 1898) C 1 X P. insulsa (Sluiter, 1898) C,M 4 X X X X P. spongiabilis (Traustedt, 1883) C 3 X X X Polycarpa sp. 1 C 1 X Polycarpa sp. 2 M 1 X Eusynstyela floridana ? (Van Name, 1921) M 1 X ASCIDIAN BIODIVERSITY 605

accomplished with dredging from ships. Direct visual samples made by either free or scuba diving in complex environments such as the mangrove roots and coral reefs is a more efficient way to collect animals and to adequately anesthetize them so that better identification is possible. Among the 15 species mostly wide- spread in the Caribbean (found in 10 or more different islands) only two, Polycli- X X num constellatum and Botrylloides nigrum,do not occur in Bocas del Toro. Unexpectedly, the families Polyclinidae and Molgulidae were poorly represented in this collection. Goodbody (1995) argued in favor of con- servation of Pelican Cays in Belize because of its rich ascidian community growing on mangrove roots. He mentioned “species richness is high: 30 species of ascidians along a 125 m stretch of the lagoon bank”. Diving site number After finding almost twice the number of species we have the impression that the Ar- chipelago Bocas del Toro should also be the object of intensive conservation effort be- cause of the varied types of habitats and the high diversity of the area.

1. Continued. Acknowledgments.—We thank Dr. Rachel Collin for having invited the senior author ABLE

T to participate in the Marine Invertebrate Taxonomy Workshop held in the Smithso- nian Tropical Research Station (STRI) at Bo- cas del Toro in August, 2003. We also thank Gabriel Jacome, Willie Pomaire, and Arca- 1234567891011121314151617181920212223242526 dio Castillo who aided with diving and § laboratory logistics during the Workshop. #

Thalassia This paper is the contribution 1494 of De- partamento de Zoologia, Universidade Federal do Paraná.

LITERATURE CITED Berrill, N. J. 1932. Ascidians of the Bermudas. Biol. Bull. 62(1):77-88.

Heller, 1878 C,M 8Goodbody, X X I. 1984a. The ascidian fauna of two X con- X X X X X trasting lagoons in the Netherlands Antilles: Pis-

Traustedt, 1883 Mcadera 2 Baai, Curacao, and the Lac of Bonaire. Stud. X X (Heller, 1878) MFauna 2 Curaç Xao Caribb. Isl. 67(202):21-61. X

(Van Name, 1902) C,M 2Goodbody, X I. 1984b. Ascidians from Caribbean shal- X Monniot, 1994 C 3low water localities. XStud. Fauna Curaçao Caribb. Isl. X X (Sluiter, 1898) M 1 X (Stimpson, 1852) C,M 367(203):62-76. X X X sp. 1sp. 2sp. 3Goodbody, M C,MI. 1 1993. 2 M 1 The ascidian fauna of a Jamaican X X number of sites with the species

# habitat: M =§ mangrove, C = coral reef, T = lagoon: Thirty years of change. Rev. Biol. Trop., supl.

Pyuridae Pyura munita P. torpida P. vittata P. vannamei Pyura Pyura Pyura Microcosmus exasperatus Herdmania pallida Molgulidae 41(10):35-38. 606 TABLE 2. Distribution of ascidian species in the Caribbean and adjacent areas, based on the following publications: Berrill (1932), Plough and Jones (1939), Van Name (1921, 1924, 1930, 1945), Millar (1962, 1978), Van der Sloot (1969), Millar and Goodbody (1974), Lafargue and Duclaux (1979), Monniot, C. (1972a, b, 1983a, b, c), Monniot, F. (1972, 1983a, b, c, 1984), Monniot and Monniot (1984), Goodbody (1984a, 1984b, 1993, 1995, 1996, 2000, 2003), Goodbody and Cole (1987), and Hernandez-Zanuy. ao ¸ Total Anguilla Antigua Aruba Aves island Bahamas Barbuda Bermuda Belize Bimini Bocas del Toro Bonaire Cuba Curac Florida Grenada Guadeloupe Guianas Grand Cayman Jamaica Little Cayman Los Frailes Margarita Martinique Puerto Rico Saba St. Croix St. Kitts St. Lucia St. Martin St. Thomas St. Vincent Trinidad

Total 2 2 13 1 8 2 49 70 2 58 133432451196155481115324272472031 APLOUSOBRANCHIA Polyclinidae

Polyclinum constellatum Savigny, 1816 14 X X X X X X ? X X X X X X X AL. ET ROCHA M. R. Polyclinum sp. 1 X Euherdmania areolata Millar, 1978 1 X E. morgani Millar & Goodbody, 1974 1 X E. fasciculata Monniot, 1983 2 X X Euherdmania sp. 1 1 X Euherdmania sp. 2 1 X Aplidium antillense (Gravier, 1955) 4 X ? X X A. bermudae (Van Name, 1902) 5 X X X X X A. constellatum (Verrill, 1871) 3 X X X A. exile (Van Name, 1902) 5 X X X X X A. lobatum Savigny, 1816 4 X X X X A. pentatrema (Monniot, 1972) 2 X X Aplidiopsis stellatus Monniot & Monniot, 1984 3 X X X Pseudodistoma saxicavum Gaill, 1972 1 X Didemnidae Didemnum calliginosum Monniot, 1984 1 X D. cineraceum (Sluiter, 1898) 3 X X X D. conchyliatum (Sluiter, 1898) 8 X X XXXX X X D. duplicatum Monniot, 1983a 5 X X X X X D. granulatum Tokioka, 1954 1 X D. halimedae Monniot, 1983a 2 X X D. inauratum Monniot, 1983a 2 X X D. ligulum Monniot, 1983a 2 X X TABLE 2. Continued. ao ¸ Total Anguilla Antigua Aruba Aves island Bahamas Barbuda Bermuda Belize Bimini Bocas del Toro Bonaire Cuba Curac Florida Grenada Guadeloupe Guianas Grand Cayman Jamaica Little Cayman Los Frailes Margarita Martinique Puerto Rico Saba St. Croix St. Kitts St. Lucia St. Martin St. Thomas St. Vincent Trinidad

Total 2 2 13 1 8 2 49 70 2 58 133432451196155481115324272472031 D. perlucidum Monniot, 1983a 2 X X D. psammathodes (Sluiter, 1895) 8 X X X XX X X X D. speciosum (Herdman, 1886) 1 X D. vanderhorsti Van Name, 1924 2 X X

Polysyncraton amethysteum (Van Name, 1902) 3 X X X 607 BIODIVERSITY ASCIDIAN P. louminae Monniot, 1984 1 X Trididemnum cyanophorum Lafargue & Duclaux, 1979 2 X X T. hians Monniot, 1983a 4 X X X X T. maragogi Rocha, 2002 1 X T. orbiculatum (Van Name, 1902) 5 X X X XX T. palmae (Monniot, 1984) 1 X T. savignii Herdman, 1886 6 X X XXX X X T. solidum (Van Name, 1902) 5 X X X X x X Lissoclinum abdominale Monniot, 1983a 2 X X L. fragile (Van Name, 1902) 6 X X X X X X L. fragile (Van name, 1902) form hospes Monniot, 1984 1 X L. perforatum (Giard, 1872) 1 X L. verrilli (Van Name, 1902) 5 X X X XX Diplosoma glandulosum Monniot, 1983 2 X X D. listerianum (Milne-Edwards, 1841) 11 X X X X XXXXXX X D. macdonaldi Herdman,1886 6 X X X X X X D. tritestis Monniot, 1984 1 X D. virens (Hartmeyer, 1909) 1 X Leptoclinides latus Monniot, 1983a 1 X L. torosus Monniot, 1983a 1 X Clavelinidae Clavelina oblonga Herman, 1880 9 X X X X X X X X X C. picta (Verrill, 1900) 3 X X X C. puertosecensis Millar & Goodbody, 1974 3 X X X 608 TABLE 2. Continued. ao ¸ Total Anguilla Antigua Aruba Aves island Bahamas Barbuda Bermuda Belize Bimini Bocas del Toro Bonaire Cuba Curac Florida Grenada Guadeloupe Guianas Grand Cayman Jamaica Little Cayman Los Frailes Margarita Martinique Puerto Rico Saba St. Croix St. Kitts St. Lucia St. Martin St. Thomas St. Vincent Trinidad

Total 2 2 13 1 8 2 49 70 2 58 133432451196155481115324272472031 Clavelina sp. 1 X belizeana Goodbody, 1996 1 X Podoclavela brasiliensis Millar, 1977 1 X Polycitoridae Cystodytes dellechiajei (Della Valle, 1877) 7 X X X XXXX

Cystodytes sp. 1 1 X AL. ET ROCHA M. R. Cystodytes cf. roseolus 1 X Eudistoma capsulatum (Van Name, 1902) 6 X X X X X X E. carolinense Van Name, 1945 2 X X E. clarum (Van Name, 1902) 5 X X X ?X E. hepaticum (Van Name, 1921) 5 X X X X X E. obscuratum (Van name, 1902) 4 X X X X E. olivaceum (Van Name, 1902) 20 X X X X X X XXXXXX XXX X X X XX E. saldanhai Millar, 1977 1 X E. surinamense Millar, 1978 1 X Eudistoma sp. 1 X Stomozoa gigantea (Van Name, 1921) 2 X X Holozoidae Distaplia bermudensis Van Name, 1902 11 X X X XX XX X X X X D. bursata Van Name, 1921 2 X X D. corolla Monniot, 1974 3 X X X D. crassa Monniot, 1983c 1 X D. stylifera (Kowalevsky, 1874) 1 X PHLEBOBRANCHIA Diazonidae Rhopalaea abdominalis (Sluiter, 1898) 4 X X XX Corellidae minuta Traustedt, 1882 6 X X X X X X Rhodosoma turcicum (Savigny, 1816) 8 X XX X X X X X TABLE 2. Continued. ao ¸ Total Anguilla Antigua Aruba Aves island Bahamas Barbuda Bermuda Belize Bimini Bocas del Toro Bonaire Cuba Curac Florida Grenada Guadeloupe Guianas Grand Cayman Jamaica Little Cayman Los Frailes Margarita Martinique Puerto Rico Saba St. Croix St. Kitts St. Lucia St. Martin St. Thomas St. Vincent Trinidad

Total 2 2 13 1 8 2 49 70 2 58 133432451196155481115324272472031 Perophoridae Perophora bermudensis Berrill, 1932 12 ? X X ? X X X X X X X X P. carpenteria Goodbody 1994 2 X X P. formosana (Oka, 1931) 2 X X

P. multiclathrata (Sluiter, 1904) 3 X X X 609 BIODIVERSITY ASCIDIAN P. regina Goodbody & Cole, 1987 1 X P. viridis Verrill, 1871 7 X X X X X X X Ecteinascidia conklini Berrill, 1932 8 X X X X X X X X E. minuta (Berrill, 1932) 10 X X X X X X X X X X E. styeloides (Traustedt, 1882) 5 X X XXX E. turbinata Herdman, 1880 13 X X X X X XXXXX X XX Ascidiidae Ascidia corelloides (Van Name, 1924) 3 X X X A. curvata (Traustedt, 1882) 12 X X X X XXXX X X X X A. interrupta Heller, 1878 12 X X X X XXX X X X X X A. sydneiensis Stimpson, 1855 9 X X X XX XX X X A. tenue Monniot, 1983 2 X X A. xamaycana Millar & Goodbody, 1974 1 X Ascidia sp. 1 1 X Ascidia sp. 2 1 X Ascidia sp. 3 1 X Phallusia nigra Savigny, 1816 16 X X X XXXXXX X X X X XXX P. caguayensis Millar & Goodbody, 1974 1 X Agnesiidae Adagnesia charcotti Monniot & Monniot, 1973 1 X STOLIDOBRANCHIA Styelidae Botrylloides magnicoecum (Hartmeyer, 1912) 2 X X B. nigrum Herdman, 1886 13 X X X X X X X X X X X X X 610 TABLE 2. Continued. ao ¸ Total Anguilla Antigua Aruba Aves island Bahamas Barbuda Bermuda Belize Bimini Bocas del Toro Bonaire Cuba Curac Florida Grenada Guadeloupe Guianas Grand Cayman Jamaica Little Cayman Los Frailes Margarita Martinique Puerto Rico Saba St. Croix St. Kitts St. Lucia St. Martin St. Thomas St. Vincent Trinidad

Total 2 2 13 1 8 2 49 70 2 58 133432451196155481115324272472031 Botryllus planus (Van Name, 1902) 5 X X X XX B. perspicuum Herdman, 1886 1 X B. schlosseri (Pallas, 1766) 3 X X X B. tuberatus Ritter & Forsyth, 1917 3 X X X Symplegma brakenhielmi (Michaelsen, 1904) 6 X X XX X X

S. viride Herdman, 1886 8 X XXXXX XX AL. ET ROCHA M. R. S. rubra Monniot, 1972 5 X X X XX Metandocarpa sterreri Monniot, 1972 2 X X Styela canopus Savigny, 1816 15 X X X X XXXXXX X XXX X S. plicata (Lesueur, 1823) 11 X X XXX X XXX XX Polyandrocarpa anguinea (Sluiter, 1898) 1 X P. sabanillae Van Name 1 X Polycarpa appropinquata (Sluiter, 1898) 1 X P. arnoldi (Michaelsen, 1914) 4 X X X X P. aurita (Sluiter, 1890) 3 X X X P. cartilaginea (Sluiter, 1898) 4 X X XX P. crossogonima Millar, 1962 2 X X P. insulsa (Sluiter, 1898) 3 X XX P. multiplicata Monniot, 1983b 1 X P. nivosa (Sluiter, 1898) 1 ? P. spongiabilis (Traustedt, 1883) 13 X X X X X XXX XX X X X P. tumida Heller, 1878 4 X X X X Polycarpa sp. 1 1 X Polycarpa sp. 2 1 X Eusynstyela floridana (Van Name, 1921) 3 X XX E. ordinata Monniot, 1983b 1 X E. tincta (Van Name, 1902) 6 X X X X X X Amphicarpa paucigonas Monniot & Monniot, 1984 1 X Stolonica sabulosa Monniot, 1972 2 X X TABLE 2. Continued. ao ¸ Total Anguilla Antigua Aruba Aves island Bahamas Barbuda Bermuda Belize Bimini Bocas del Toro Bonaire Cuba Curac Florida Grenada Guadeloupe Guianas Grand Cayman Jamaica Little Cayman Los Frailes Margarita Martinique Puerto Rico Saba St. Croix St. Kitts St. Lucia St. Martin St. Thomas St. Vincent Trinidad

Total 2 2 13 1 8 2 49 70 2 58 133432451196155481115324272472031 Tibitin halimedae Monniot, 1983b 2 X X Cnemidocarpa valborg Hartmeyer, 1919 1 X Pyuridae

Bathypera goreaui Millar & Goodbody, 1974 2 X X 611 BIODIVERSITY ASCIDIAN Pyura discrepans (Suiter, 1898) 1 X P. lignosa Michalsen, 1908 2 X X P. munita (Van Name, 1902) 5 X X X XX P. torpida (Sluiter, 1898) 3 X XX P. vannamei Monniot, 1994 1 X P. vittata (Stimpson, 1852) 14 X X X X XXXXXXX X X X Pyura sp. 1 1 X Pyura sp. 2 1 X Pyura sp. 3 1 X Microcosmus anchylodeirus Traustedt, 1883 2 X X M. exasperatus Heller, 1878 14 X X X X XXXXX X X X X X M. helleri Herdman, 1881 8 X X X X X X X X Herdmania pallida (Heller, 1878) 11 X X XXXXX X X X X Halocynthia microspinosa (Van Name, 1921) 4 X X X X Molgulidae Molgula contorta Sluiter, 1898 1 X M. davidi Monniot, 1972 1 X M. fortuita Monniot & Monniot, 1984 1 X M. habanensis Van Name, 1945 1 X M. manhattensis (De Kay, 1843) 1 X M. occidentalis Traustedt, 1883 9 X X XXXX X X X Bostrichobranchus pilularis Verrill, 1871 1 X B. digonas Abbott, 1951 1 X 612 R. M. ROCHA ET AL.

Goodbody, I. 1994. The tropical western atlantic Pero- Monniot, C. 1983a. Ascidies littorales de Guadeloupe phoridae (): 1. The Perophora. Bull. 2. Phlebobranches. Bull. Mus. Natn. Hist. Nat., Ser. Mar. Sci. 55(1):176-192. 4, A, 5(1):51-71. Goodbody, I. 1995. Ascidian communities in Southern Monniot, C. 1983b. Ascidies littorales de Guadeloupe. Belize – a problem in diversity and conservation. 4. Styelidae. Bull. Mus. Natn. Hist. Nat., Ser. 4, A, Aquat. Conserv. 5:355-358. 5(2):423-456. Goodbody, I. 1996. Pycnoclavella belizeana, a new spe- Monniot, C. 1983c. Ascidies littorales de Guadeloupe cies of ascidian from the Caribbean. Bull. Mar. Sci. 6. Pyuridae et Molgulidae. Bull. Mus. Natn. Hist. 58(2):590-597. Nat., Ser. 4, A, 5(4):1021-1044. Goodbody, I. 2000. Diversity and distribution of ascid- Monniot, C. and F. Monniot. 1984. Ascidies littorales ians (Tunicata) in the Pelican Cays, Belize. Atoll de Guadeloupe. 7. Espèces nouvelles et comple- Res. Bull. 480: 302-326. mentaries a l’inventaire. Bull. Mus. Natn. Hist. Nat., Goodbody, I. 2003. The ascidian fauna of Port Royal, Ser. 4, A, 6(3):567-582. Jamaica.—I. Harbor and mangrove dwelling spe- Monniot, F. 1972. Ascidies Aplousobranches des Ber- cies. Bull. Mar. Sci. 73: 457-476. mudes. Polyclinidae et Polycitoridae. Bull. Mus. Goodbody, I., and L. Cole. 1987. A new species of Natn. Hist. Nat., Ser. 3, 82:949-962. Perophora (Ascidiacea) from western Atlantic, in- cluding observations on muscle action in related Monniot, F. 1983a. Ascidies littorales de Guadeloupe. species. Bull. Mar. Sci. 40(2):246-254. 1. Didemnidae. Bull. Mus. Natn. Hist. Nat., Ser. 4, A, Hernandez-Zanuy, A. 1990. Lista de ascidias cubanas. 5(1):3-49. Poeyana 388:1-7. Monniot, F. 1983b. Ascidies littorales de Guadeloupe. Kott, P. 1985. The Australian Ascidiacea. part 1, Phle- 3. Polyclinidae. Bull. Mus. Natn. Hist. Nat., Ser. 4, A, bobranchia and Stolidobranchia. Mem. Qd. Mus. 23: 5 (2):413-422. 1-440. Monniot, F. 1983c. Ascidies littorales de Guadeloupe Kott, P. 1990. The Australian Ascidiacea, part 2, 5. Polycitoridae. Bull. Mus. Natn. Hist. Nat., Ser. 4, Aplousobranchia (1). Mem. Qd. Mus. 29(1):1-226. A, 5(4):999-1019. Kott, P. 1992. The Australian Ascidiacea, part 3, Monniot, F. 1984. Ascidies littorales de Guadeloupe 8. Aplousobranchia (2). Mem. Qd. Mus. 32(2):37-620. Questions de systematique evolutive posées par les Kott, P. 2001. The Australian Ascidiacea. Part 4, Didemnidae. Bull. Mus. Natn. Hist. Nat., Ser. 4,A, Aplousobranchia (3), Didemnidae. Mem. Qd. Mus. 6(4):885-905. 47(1):1-407. Plough, H. H., and N. Jones. 1939. Ecteinascidia tortu- Lafargue, F., and G. Duclaux. 1979. Premier example gensis, species nova. With a review of the Pero- en Atlantique tropical d’une association symbio- phoridae (Ascidiacea) of the Tortugas. Pap. Tortu- tique entre une ascidie Didemnidae et une cyano- gas Lab. 32:47-60. phyce chroococcale: Trididemnum cyanophorum Ramos Esplá, A. A. 1988. Ascidias litorales del Medi- nov. sp. et Synechocystis trididemni nov. sp. Ann. terráneo Ibérico. Faunística, ecología y biogeo- Inst. Oceanogr. 55:163-184. grafía. Doctoral Thesis, Universidad de Barcelona. Millar, R. H. 1962. Some Ascidians from the Car- Van der Sloot, C. J. 1969. Ascidians of the family Sty- ibbean. Stud. Fauna Curaçao Caribb. Isl. 13(59):61-77. elidae from the Caribbean. Stud. Fauna Curaçao Car- Millar, R. H. 1978. Ascidians from the Guyana shelf. ibb. Isl. 110:1-57. Neth. J. Sea Res. 12(1):99-106. Millar, R. H. and Goodbody, I. 1974. New species of Van Name, W. G. 1921. Ascidians of the West Indian Bull. Am. Ascidians from the West Indies. Stud. Fauna region and southeastern United States. Mus. Nat. Hist Curaçao Caribb. Isl. 45:142-161. . 44:283-494. Monniot, C. 1972a. Ascidies Stolidobranches des Ber- Van Name, W. G. 1924. Ascidians from Curacao. Bijdr. mudes. Bull. Mus. Natn. Hist. Nat., Ser. 3, 43(57): Dierkd. 23:23-32. 617-643. Van Name, W. G. 1930. The Ascidians of Porto Rico Monniot, C. 1972b. Ascidies Phlebobranches des Ber- and Virgin Islands. N. Y. Acad. Sci. 10:401-512. mudes. Bull. Mus. Natn. Hist. Nat., Ser. 3, 61(82): Van Name, W. G. 1931. New North and South Ameri- 938-948. can ascidians. Bull. Am. Mus. Nat. Hist. 61:207-225.