Common Sponges from Shallow Marine Habitats from Bocas Del Toro Region, Panama

Home , Clionaidae, Grantiidae, Mycale laevis, Phloeodictyidae

Common Sponges from Shallow Marine Habitats from Bocas del Toro Region, Panama

MARÍA CRISTINA DÍAZ1,2

1Department of Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0163. 2Museo Marino de Margarita, Boulevard El Paseo, Boca del Rio, Peninsula de Macanao, Nueva Esparta, Venezuela. [email protected]

ABSTRACT.—A survey of Porifera biodiversity and distribution was done in three islands of the Bocas del Toro region, Panama. Fourteen sites were explored using mask and snorkel, and occasionally scuba equip- ment, during ten days. Forty-one species are added to the species known for the region. Our current estimate of sponge diversity in shallow exposed marine habitats of Bocas del Toro is 120 species. Twenty-three species are added to the Panamanian sponge fauna, four of which are new species currently being described. A patchy distribution was observed for most of the sponge species, while few species were present at all sites visited. Approximately 23% (30 species) of the diversity encountered was widely distributed. The species composition of the most typical or common sponges from reef, seagrass beds, or mangrove habitats, agrees with that of other Caribbean sites. An appendix with a field guide to the species encountered in this study is presented.

KEYWORDS.—Sponges, biodiversity, mangrove, reefs, field guide.

INTRODUCTION Litter et al. 1985; Taylor 1986; Ellison and Farnsworth 1992; Rützler 1995; Rützler et Sponges are among the most diverse and al. 2000; Wulff 2000). Despite the ubiquity abundant inhabitants of benthic tropical of this group in Caribbean ecosystems, its communities such as coral reefs and mangrove systems (Rützler and Feller 1987; diversity and relative abundance remains Diaz and Rützler 2001). Coral reef sponges poorly understood for large parts of the re- in the Caribbean usually constitute the gion. Whenever unexplored geographic ar- most diverse benthic group, 100-200 spe- eas are studied or particular sponge taxa cies, while other conspicuous coral reef (genus, family, or order) are revised, a con- groups such as hexacorallians or octocoral- siderate number of new species is discov- lians are represented by 50-70 species (see ered. Examples of this trend are among re- Diaz and Rützler 2001 for a review). In sub- cent work carried out in Jamaica (Lehnert tidal mangrove habitats, sponges, along and Van Soest 1998, 1999), and Belize (cer- with algae, ascidians, anemones, and hy- tain taxa such as Chalinidae [de Weerdt et droids, are among the most conspicuous al. 1991; de Weerdt 2000]; or the Axinelli- epibionts (Litter et al. 1985; Rützler and dae [Alvarez et al. 1998; or Mycalidae Feller 1987, 1996; Calder 1991a; Ellison and Hajdu and Rützler 1998]), and on the sys- Farnsworth 1992; Rützler et al. 2000; Good- tematic revisions of the familiy Chalinidae, body 2000; Macintyre et al. 2000). Sponge Haplosclerida, by De Weerdt (2000), or the species richness from mangrove systems in family Axinellidae by Alvarez et al. (1998). the Caribbean ranges from a dozen to 150 The archipelago of Bocas del Toro is lo- species, depending apparently on a combi- cated on the western Caribbean shore of nation of physical factors, such as water Panama and comprises more than 68 is- turbidity, salinity, wave exposure, sub- lands and mangrove keys. Knowledge of strate availability, and tidal ranges, and its shallow water sponge fauna is limited to biological factors such as competition, lar- two major studies: Guzmán and Guevara val supply, and predation (Calder 1991b; (1998, 1999), and Guzmán (2003), who list 63 465 466 MARÍA CRISTINA DÍAZ sponge species from open reef habitats, and two sites were surveyed by scuba-diving in Nichols and Barnes (unpublished) who reg- August 2004 (15-60 ft depth). Table 1 lists istered 86 taxa from reefs, mangroves, and the coordinates as well as the type of ma- seagrass beds in the Bocas region, includ- rine community explored during each visit. ing 12 taxa identified only to family and 11 Surveys consisted of swimming for ap- identified only to genus. The results add up proximately one hour and registering the to approximately 79 species of sponge spe- occurrence of each sponge species. When a cies reported from the area. During August sponge was encountered it was photo- 2003 the first Smithsonian Marine Science graphed and a small fragment was col- Network workshop took place at the Bocas lected, labeled, and transported to the labo- del Toro Station-Smithsonian Tropical Re- ratory in a cooler. Once in the laboratory, search Institute (STRI). The goal of this the specimens were fixed in 10% formalin workshop was to increase our understand- (seawater) and within a week transferred ing of the diversity and distribution of into ethanol 70%. If a species was not rec- poorly-known marine taxa. The present ognized immediately, a taxonomic study study summarizes the results of a survey of was performed in the laboratory, using a the diversity and distribution of sponges compound microscope to examine skeleton from various shallow water habitats structure in dried hand sections cleared in around this region’s islands and presents a Permount medium . Spicule types were de- field guide to facilitate the identification of termined after dissolving a fragment (2-4 the most common sponges (see Photo- mm3) in concentrated household bleach graphic Indentification Guide on this vol- (5% sodium hypoclorite). The samples ume). taken are kept in the Bocas del Toro, STRI, marine station. MATERIALS AND METHODS A qualitative estimate of the abundance Twelve sites in the Bocas del Toro Archi- of each sponge species was carried out pelago were surveyed by mask and snorkel while surveying each site. Species were during August 3-12 2003 (0-20 ft deep), and classified in four categories: very common

TABLE 1. Sites surveyed in the Bocas del Toro Archipelago region in August 2003, and August 2004 (*).

Locality Latitude-N Longitude-W Habitat type Crawl Key 1 9°15Ј 82°9Ј030Љ Rhizophora mangle fringe (3 ft deep), contiguous to shallow reef Eagle Park-SW 9°19Ј 82°13Ј R. mangle fringe (6 ft deep) and seagrass Solarte Island Solarte 1 9°17Ј030Љ 82°10Ј020Љ R. mangle fringe (6 ft deep) on internal lagoon Solarte 2 9°17Ј987Љ 82°12Ј404Љ R. mangle fringe (6 ft deep) on canal between islands Solarte 3 9°17Ј208Љ 82°11Ј650Љ R. mangle fringe (3 ft deep), contiguous to shallow reef Crawl Key Canal 9°15Ј050Љ 82°07Ј631Љ Shallow reef (20-30 ft deep) Millepora and Porites dominant corals Colo´n Island Punta Caracol 9°22Ј627Љ 82°18Ј117Љ R. mangle fringe (3 ft deep), contiguous to shallow reef STRI Point 9°21Ј169Ј 82°15Ј528Љ R. mangle fringe (3-6 ft deep), contiguous to shallow reef ITEC, Colo´n Is. Shallow reef. (5-15 ft deep) Big Bight. 9°22Ј184Ј 82°16Ј586Љ R. mangle fringe (6-9 ft deep) along a lagoon Swan Key 9°27Ј198Љ 82°18Ј024Љ Shallow reef. (5-15 ft deep) Strong surge and currents Pastores Island 9°14Ј332Љ 82°19Ј.968Љ Shallow, extensive, Porites reef Adriana’s Key* 9°14Ј021Љ 82°10Ј023Љ Coral reef associated with nearby keys, (10-60 ft deep) High coverage of stony coral, algae and sponges Cristobal Island 9°15Ј379Љ 82°14Ј136Љ Fringing reef dominated by Agaricia, Porites and large south* clumps of red alga Amphiroa rigida var. antillana, specially below 20-30 ft deep. Large amount of fine sediments along the reef profile (10-55 ft deep). COMMON SPONGES SHALLOW MARINE HABITATS 467

(more than ten specimens seen), abundant calcareous sponge Clathrina primordialis (6-10), few (2-5 specimens), and rare (one that can be found either in reef or man- specimen). groves. Fifteen species are first records for Bocas del Toro region but were reported RESULTS previously for other localities in the Pana- manian Caribbean. Such is the case of Che- Sponge biodiversity lonaplysilla erecta, Verongula reiswigi One hundred and four species were en- (Guzmán 2003), Dysidea etheria, Haliclona countered in this study, of which 41 are manglaris, Cribochalina vasculum, Oceanapia new for the area (Table 2). As a result, peltata, Nipahtes caycedoci, Spirastrella mollis, sponge diversity known from the Bocas del Lissodendoryx colombiensis, Erylus formosus, Toro region reached 120 species. Four of 12 and Halichondria magniconulosa (Clifton et species identified only to genus are consid- al. 1997; Wulff 2000). ered new species and are currently being described. These species include two from the order Haplosclerida (Haliclona sp. and Sponge species distribution and ?Xestospongia sp.), and two from the order relative abundance Poecilosclerida (Mycale (Paresperella) sp., and Iotrochota sp.). Most species (58%) occur in one to four Haliclona sp., and ?Xestospongia sp. (see sites, while only one species (Niphates illustration in the Photographic Identifica- erecta) occurs at all fourteen sites surveyed tion Guide; this volume) are abundant on (Table 2). Thirty of the 126 species (23%) hard substrata of shallow reef habitats, and occur at five or more sites. Aplysina cauli- both posses endosymbiotic filamentous formis, A. fulva, Aiolochroia crassa, Verongula cyanobacteria. Iotrochota sp. is found in rigida, Ircinia felix, I. campana, Niphates muddy areas of certain deeper reefs (>40 erecta, Amphimedon compressa, A. erina, Xes- ft). Mycale (Paresperella) sp. is a common tospongia caycedoci, X. muta, X. proxima, X. species but only found on mangrove roots. carbonaria, Xestospongia sp., Iotrochota birotu- It represents the first find of a whole speci- lata, Mycale laevis, M. laxissima, and Cliona men of this sub-genus in the Caribbean. delitrix were common in reefs sites, while Hadju and Rützler (1998) describe the dis- Hyrtios proteus, Spongia pertusa, Haliclona tinctive spicules of this subgenus (serrated implexiformis, H. manglaris, H. curacaoensis, sigmas) found contaminating a specimen of H. tubifera, Chalinula molitba, Lissodendoryx Mycale (Aegropila) citrina, therefore assum- isodyctialis, Clathria schoenus, and Tedania ing the occurrence of a species of this sub- ignis were common in mangroves. Three genus in Belizean mangroves but lacking a species—Niphates erecta, Scopalina ruetzleri, complete specimen. and Chondrilla nucula—may be common in Twenty-four species are new record for both reef and mangrove habitats. The ma- the Panamanian sponge fauna. Half of jority of the species (74 of 126) were at least these species (13) are exclusively found in- in two of the three habitats surveyed, while habiting mangrove roots or peat (Haliclona 28 species occur exclusively in mangroves piscadaerensis, H. curacaoensis, H. mucifi- and 25 species in reefs (Table 2). brosa, H. vermeuleni, H. tubifera, Chalinula zeae, C. molitba, Terpios manglaris, Tethya aff. seychellensis, Mycale magnirhaphidiphera, M. DISCUSSION carmigropila, Mycale undescribed species, and Geodia papyracea). Eleven inhabit pre- dominantly reef environments (Amphim- After surveying the shallow water habi- edon viridis, Callyspongia fallax, C. pallida, tats of three islands in Bocas del Toro re- Haliclona vansoesti, Oceanapia oleracea, Spi- gion in a relatively short time, the known rastrella hartmani, Clathria echinata, plus sponge diversity increased 34%. This dem- three undescribed species of Haliclona Xes- onstrates the importance of this kind of ex- tospongia sp., and Iotrochota sp.), and the ploration to further our biodiversity knowl- 468 TABLE 2. Sponge species distribution and relative abundance (1 = very common, 2 = abundant, 3 = few, 4 = rare) in 14 studied sites in islands from the Bocas del Toro region, Panama. Column legends: Rf = References (1. Nichols and Barnes, unpublished data; 2. Guzman 2003; 3. Present study). Sites are organized according to the major habitat (s) surveyed: EP = Eagle Park, S1 = Solarte site 1, S2 = Solarte site 2; AK = Adriana’s Key, CK2 = Crawl Key Bank, SK = Swan Key, Ti = TINC, Cr 1 = Cristobal Island reef; CK 1 = Crawl Key 1, PI = Isla Pastores, SP = STRI Point, PC = Punta Caracol, BB = Big Bight, and S3 = Solarte site 3. (*found only on mangroves, **found only on reefs, ***found only on seagrass); D. & M. = Duchassaing and Michelotti; G. & W. = George and Wilson. The order (O), sub-order (S.O.), and family of each species is indicated.

Reef Mangrove and reef Mangrove Sample CK C CK Species Rf ն EP S1 S2 AK 2SKTi1 1 PISPPCBBS3 Class Demospongiae O. Verongida, Aplysinidae Aiolochroia crassa (Hyatt, 1875) 1-3 NS 2 4 2 2 3 3 2 Aplysina cauliformis (Carter, 1882) 1-3 BT-048 2 3 3 3 3 2 4 3 A. fulva (Pallas, 1776) 1-3 BT-049 3 3 1 3 1 1 3 2 3 3 2 MAR A. insulares D. & M., 1864 1-3 BT-061 3 3 3 3 Í A. lacunosa (Lamarck) 1-3 NS 3 3 D CRISTINA A Verongula reiswigi (Alcolado, 1984) 3 BT-051 2 2 2 V. rigida (Esper, 1794) 1-3 BT-050 2 3 2 3 3 3 O. Dyctioceratida, Spongiidae Spongia pertusa Hyatt, 1877 1,3 BT-005 1 3 2 4 2 2 S. tubulifera Hyatt, 1877 3 NS 4 Í Hyatella cavernosa (Pallas, 1766) 1-3 NS AZ O. Dyctioceratida, Thorectidae Hyrtios proteus (D. & M., 1864) 1,3 BT-006 1 3 4 2 3 3 3 O. Dyctioceratida, Irciniidae Ircinia campana (Lamarck, 1816) 1-3 NS 2 2 2 2 2 2 2 2 I. cf.felix (ramose form) 3 BT-093 2 3 2 I. felix (D & M, 1864) 1-3 BT-052 2 3 2 2 2 3 2 I. strobilina (Lamarck, 1816) 1-3 BT-075 12 2 I. sp. A* 3 BT-032 4 I. sp. B* 3 BT-100 4 Dysidea etheria de Laub., 1936* 3 BT-026 3 1 3 2 2 2 O. Halisarcida, Halisarcidae Halisarca caerulea Vacelet and Donadey, 1989 3 NS Halisarca sp.* 1-3 223 O. Dendroceratida, Darwinellidae Aplysilla gracilis (Merejkowsky, 1878) 2,3 NS 3 TABLE 2. Continued.

Reef Mangrove and reef Mangrove Sample CK C CK Species Rf ն EP S1 S2 AK 2SKTi1 1 PISPPCBBS3 Chellanoplysilla erecta (Row, 1911) 2,3 BT-157 3 3 2 3 O. Haplosclerida, Chalinidae H. (Reniera.) curacaoensis (Van Soest, 1980)* 3 BT-017 1 2 3 2 2

Haliclona (R.) implexiformis (Hechtel, 1965)* 3 BT-015 2 3 2 1 2 469 3 3 HABITATS MARINE SHALLOW SPONGES COMMON H. (R.) manglaris Alcolado 1984* 3 BT-014 1 1 1 1 2 1 1 H. (R.) mucifibrosa de Weerdt et al., 1991* 1 BT-088 23 H.(R.) tubifera (G. & W., 1919)* 1-3 BT-12 2 2 3 2 2 3 H. (Soestella) caerulea (Hechtel, 1965) 1,3 NS 3 H.(S.) picadaerensis van Soest 1980* 3 BT-121 3 3 2 2 H. (S.) twincayensis de Weerdt et al., 1991* 1,3 BT-062 3 3 H. (S.) vermeuleni de Weerdt, 2000* 1,3 NS 1 Haliclona sp.1 (new species)** 3 BT-045 1 Chalinula. molibta (de Laub., 1949)* 1,3 BT-016 2 3 2 1 2 2 C. zeae de Weerdt, 2000* 3 BT-129 4 O. Haplosclerida, Niphatiidae Amphimedon compressa D. & M., 1864 1-3 BT-043 1 2 2 2 3 2 A. erina (de Laub., 1936) 1-3 BT-058 1 2 1 2 4 A. viridis D. & M., 1864 3 BT-064 1 N. caycedoci Zea & Van Soest 1986 3 BT-11 2 2 2 3 Niphates erecta D. & M., 1864 1-3 BT-07 4 3 3 1 2 2 2 1 2 1 2 1 3 2 Aka cf. brevitubulata (Pang, 1973)** 1 A. coralliophagon (Rützler, 1971)** 1-3 NS 3 3 O. Haplosclerida, Callyspongiidae Callyspongia. armigera D. & M., 1864 2,3 BT-142 @ C. fallax (D. & M., 1864) 3 NS 3 4 3 2 C. pallida (Hechtel, 1965) 3 BT-079 4 C. vaginalis (D. & M., 1864) 1-3 NS 3 2 2 2 O. Haplosclerida, Phloeodictyidae Oceanapia bartschi (De Laub., 1934) 2,3 BT-071 1 O. nodosa (G. & W., 1919) 1,3 NS 44 O. oleracea (Schmidt, 1870) 3 BT-149 3 O. peltata (Schmidt, 1870) 3 BT-148 22 Calyx podatypa (de Laub., 1934)** 2,3 BT-057 3 3 2 470 TABLE 2. Continued.

Reef Mangrove and reef Mangrove Sample CK C CK Species Rf ն EP S1 S2 AK 2SKTi1 1 PISPPCBBS3 O. Haplosclerida, Petrosiidae Xestospongia carbonaria (Lamark, 1814) 1-3 BT-010 2 2 2 3 2 2 2 3 X. muta (Schmidt, 1870)** 1-3 NS 3 4 2 3 3 2 X. proxima (D. & M., 1864) 1-3 BT-091 2 3 3 1 3 2 X. rosariensis Zea&Rützler, 1983** 1-3 BT-086 1 3 3 3 X. subtriangularis (Duch., 1850) 1,3 BT-060 3 1 2 2 Xestospongia sp. (undescribed) 3 BT-019 4 3 2 2 3 4 Petrosia pellasarca (de Laub., 1934)** 2,3 BT-186 4 P. weinbergi van Soest, 1980** 3 BT-056 4 MAR O. Agelasida, Agelasidae A. clathrodes (Schmidt, 1870)** 2 Í Agelas conifera (Schmidt, 1870)** 2 D CRISTINA A A. dispar (D. & M., 1864)** 2 O. Halichondrida, Halichondriidae Halichondria lutea Alcolado, 1984** 3 BT-172 3 H. magniconulosa, Hechtel 1965* 3 BT-018 3 3 3

H. melanadocia de Laub., 1936* 1,3 BT-189 2 Í Petromyca ciocalyptoides (van Soest & Zea, 1986)** 3 NS 2 AZ O. Halichondrida, Axinellidae Dragmacidon reticulata (Ridley & Dendy, 1886)** 1-3 BT-066, 69 4 1 4 3 Ptilocaulis walpersi (D. & M. 1864)** 1,3 BT-77 3 4 Ectyoplasia ferox (D. & M. 1864)** 1 O. Halichondrida, Dictyonellidae Scopalina ruetzleri (Wie., 1977) 1-3 BT-01 4 3 2 3 Svenzea zeai (Alvarez et al., 1998)** 1 O. Spirophorida, Tetillidae Cinachyrella alloclada (Uliczka, 1929) 2,3 3112 C. apion (Uliczka, 1929)* 3 BT-130 3 3 4 3 O. Hadromerida, Placospongiidae Placospongia intermedia Sollas, 1888 1,3 BT-029 2 2 2 2 O. Hadromerida, Spirastrellidae Spirastrella coccinea Schmidt, 1868 2,3 4 S. hartmani Boury-Esnault et al., 1999 1,3 BT-154 3 3 3 TABLE 2. Continued.

Reef Mangrove and reef Mangrove Sample CK C CK Species Rf ն EP S1 S2 AK 2SKTi1 1 PISPPCBBS3 S. cf. mollis Verill, 1907 sensu Rützler, 1986 3 NS 4 Diplastrella megastelata Hechtel, 1965 1 O. Hadromerida, Suberitidae

Terpios sp. 1 471 HABITATS MARINE SHALLOW SPONGES COMMON Terpios manglaris Rützler y Smith, 1993 3 NS 32 Prosuberites laughlini (Diaz et al., 1987) 1-3 BT-190 3 Suberites aurantiaca (De Laub., 1936)* 3 BT-107 2 3 3 O. Hadromerida, Tethyidae Tethya actinea de Laub., 1950* 2,3 BT-131 4 4 T. aff. seychellensis (Wrighy, 1881)* 3 BT-094 4 4 Discodermia dissoluta Schmit, 1870** 2 23 O. Hadromerida, Clionaidae Cliona varians (D. & M., 1864) 1-3 NS 3 3 1 2 Cliona delitrix Pang, 1973** 1-3 NS 2 3 2 3 2 2 C. aprica Pang, 1973** 2,3 BT-072 3 1 3 3 C. tenuis Zea & Weil, 2003** 3 BT-073 4 1 3 C. caribbaea Carter 1882** 2,3 NS Spheciospongia vesparium (Lamarck) 1-3 BT-140 4 Cervicornia cuspidifera (Lamarck) 1 O. Hadromerida, Chondrillidae Chondrilla nucula Schmidt, 1862 1-3 BT-025 2 1 1 3 2 1 2 1 Chondrosia collectrix Schmidt, 1862 1-3 BT-188 3 2 3 2 O. Poecilosclerida S.O. Microcionina, Microcionidae Clathria echinata (Alcolado, 1984) 3 BT-181 C. (Thalysias) venosa (Alcolado, 1984)* 1-3 BT-003 4 2 4 3 C. (T) microchela(Hechtel, 1965) 3 NS C. (T.) schoenus (de Laub., 1936) 1-3 BT-002 3 2 3 3 3 2 1 3 3 2 C. cf. ferrea (de Laub., 1936) 3 BT-074 3 Holopsama helwigi (de Laub., 1936) 1 3 S.O. Microcionina, Acarnidae Acarnus nicoleae van Soest, et al.91** 1 S.O. Microcionina, Raspailiidae 472 TABLE 2. Continued.

Reef Mangrove and reef Mangrove Sample CK C CK Species Rf ն EP S1 S2 AK 2SKTi1 1PISPPCBBS3 Ectyoplasia ferox (D. & M., 1864)** 2 S.O. Myxillina, Coelosphaeridae Lissodendoryx isodictyalis (C.,1882)* 2,3 BT-24 2 3 1 3 2 L. colombiensis Zea & van Soest, 1986 3 BT-042 1 1 4 S.O. Myxillina, Family Myxillidae Desmasapma anchorata (Carter, 1882) 1-3 BT-122 4 3 S.O. Myxillina, Crambiidae Monanchora arbuscula (D & M, 1864) 2,3 BT-040 3 2 2 3 S.O. Myxillina, Family Tedaniidae MAR Tedania ignis (D & M, 1864) 1,3 BT-004 3 1 4 4 3 1 3 1 S.O. Myxillina, Family Myxillidae Í Iotrochota birotulata (Higgin, 1877) 1,3 NS 2 1 2 2 1 2 1 3 D CRISTINA A Iotrochota sp. 3 BT-174 4 S-O. Mycalina, Desmacellidae Biemna caribaea Pullitzer-Finali, 1986* 3 BT-022 3 3 Biemna sp.* 3 BT-023 3

Neofibularia notilangere (D & M, 1864) 2,3 BT-081 3 DR 2 Í S-O. Mycalina, Mycalidae AZ Mycale arndti Van Soest, 1984** 2 M. carmigropila Hadju & Rützler, 1998* 3 BT-119 3 4 4 M. citrina Hadju & Rützler, 1998* 3 NS 4 M. cf. americana Van Soest, 1984 2 M. laevis (Carter, 1882)* 1-3 BT-053 3 1 1 3 1 2 2 2 2 2 2 M. laxissima (D & M, 1864) 1-3 NS 3 2 2 2 2 4 M. magniraphidiphera Van Soest, 1984* 3 NS 3 M. microsigmatosa Arndt, 1927 1,3 BT-030 2 1 2 1 M. angulosa (D & M, 1864) 3 NS 4 Mycale sp. (thin yellow)* 3 BT-028 4 Mycale (Paresperella) new species 3 BT-169 3 2 O. Astrophorida, Geodiiae Geodia papyracea Hechtel, 1965* 3 NS 4 4 Erylus formosus Sollas 1886** 3 BT-103 2 4 O. Homoscleromorpha, Plakinidae COMMON SPONGES SHALLOW MARINE HABITATS 473

edge. Three main factors may have contributed to this large increase on species number. First, mangrove habitats were not included in earlier surveys (Guzmán and Guevara 1998, 1999; Guzmán 2003): forty- two percent (18 species) of the species added herein are restricted to this habitat.

4 Second, the taxonomic expertise of the ob- server might have aided in the discrimina- tion of species in this difficult group. Third, the general patchiness of sponge distribution in mangrove systems and the rarity of 1 PISPPCBBS3

CK most species (Diaz et al. 2004) could also contribute to the increase of species num-

1 bers as new areas were surveyed. Another C consideration is that more species are de- tected if large areas are surveyed by unre- stricted swimming along suitable substrata, rather than being restricted to line transects placed along the mangrove fringe (Diaz et Reef Mangrove and reef al. 2004). It is quite possible that estimates of 2SKTi CK sponge diversity in Bocas del Toro region will double once there is data for deeper

AK coral reefs (>20 ft), mangroves surrounding other islands (e.g., Cayo de Agua and Bas- timentos), the bordering mainland section, and new habitats (cryptic reef areas, muddy bottoms, and coral rubble). Fair comparison of this region’s diversity with

Mangrove that of others in the Caribbean realm must await the results of further exploration. EP S1 S2 With the collaboration of colleagues, four species, that seem new to science, are being studied to determine their generic and spe- ն NS BT-034 4 1 BT-184 BT-037 2 3 4 Sample cific status. It is expected that more undescribed species will be found as the region is examined in more detail. 1 3 3 3 2 2,3 The most species-rich sponge genera in Bocas el Toro reef habitats are Aplysina (Aplysinidae, Verongida) and Xestospongia (Petrosiidae, Haplosclerida), with four and five species, respectively. In the mangrove, the most diverse genera are Haliclona

was collected in Pta. Bastimentos (Chalinidae, Haplosclerida), and Mycale (Wil., 1902) (Haeckel, 1872)

Ulizka, 1929** (Mycalidae, Poecilosclerida), with six and

(Carter, 1882) nine species, respectively. These genera are (Schmidt) also very diverse in other Caribbean loca- tions. In Bocas del Toro, the petrosiids, in sp. 2. Continued. particular Xestospongia, are highly diverse and abundant; it harbors typical Caribbean ABLE Callyspongia armigera species (X. muta, X. carbonaria, and X. sub- T @ O. Leucosoleniida, Grantiidae Leucandra aspera Calcarea, O. Clathrinida, Clathrinidae Clathrina primordialis Oscarella Plakortis halichondroides Plakinastrella onkodes SpeciesP. angulospiculatus Rf triangularis) as well as species described 474 MARÍA CRISTINA DÍAZ from Colombia, and observed mostly in the the Bocas del Toro region are similar to West-Central Caribbean (X. rosariensis, and those at other Caribbean sites, sharing the X. proxima). identity of the most common and wide- Evaluation of species distribution at Bo- spread species, as well as the patchy distri- cas, although of qualitative nature, con- bution of most species at local scales. firms certain trends reported previously. The patchiness of species distribution, Acknowledgments.—The author thanks where a large part of the species are seen the Smithsonian Institution-Marine Science only once or twice in a survey, was re- Network for support to attend the first Bo- corded by Rützler et al. (2001), Diaz et al. cas del Toro Biodiversity workshop held in (2004), and Alcolado (unpublished data). August 2003. Dr. Rachel Collin (director) The species spectrum of the most typical or and Dr. Gabriel Jacóme (scientific coordina- common sponges from reefs, seagrass beds, tor), and support staff of the STR-Bocas del or mangrove habitats conforms well with Toro Marine station are acknowledged for that of other Caribbean sites such as Cuba, helping to accomplish this work. Dr. Klaus Jamaica, Bahamas, Belize, Panama, Colom- Rützler assisted with taxonomic advice, bia, and Venezuela (Rützler 1969; Wieden- and Drs. Scott Nichols, Penny Barnes, and mayer 1977; Zea 1984; Farnsworth and El- Pedro Alcolado provided unpublished lison 1996; Rützler et al. 2000; Wulff 2000; data. Diaz et al. 2004; Alcolado unpublished data). The most common species on reefs habi- LITERATURE CITED tats are Aplysina fulva (11 sites), A. cauliformis Aiolochroia crassa Ir- Alcolado, P. M. 1984. Nuevas especies de esponjas en- (8 sites), (7 sites), contradas en Cuba. Poeyana 27:1-22. cinia campana and I. felix (seven sites each), Alvarez , B., R. W. M. van Soest, and K. Rützler. 1998. X carbonaria (eight sites), Iotrochota birotu- A revision of Axinellidae (Porifera: Demospon- lata (8 sites) and Niphates erecta (14 sites), giae) of the Central West Atlantic region. Smiths. which are common throughout the Carib- Contrib. Zool. 598:1-45. bean and the Gulf of Mexico. Similarly, Calder, D. R. 1991a. Abundance and distribution of mangrove sponges such as Haliclona tubi- hydroids in a mangrove ecosystem at Twin Cays, fera (6 sites), Hyrtios proteus, Haliclona im- Belize, Central America. Hydrobiologia 216/217:221- 228. plexiformis, and Haliclona manglaris (7 sites), Calder, D. R. 1991b. Associations between hydroid Tedania ignis, and Iotrochota birotulata (8 species assemblages and substrate types in the sites), and Clathria schoenus (10 sites) are the mangal Twin Cays, Belize. 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