Caribbean Journal of Science, Vol. 41, No. 3, 465-475, 2005 Copyright 2005 College of Arts and Sciences University of Puerto Kico, Mayagiiez

Common Sponges from Shallow Marine Habitats from Bocas del ToroRegion, Panama

M a r í a C r is t i n a D í a z 1,2

■‘-Department o f Zoology, National M useum o f Natural H istory, Smithsonian Institu tion , W ashington, D C 20560-0163. 2M useo M arino de Margarita, Boulevard El Paseo, Boca del Rio, Peninsula de Macanao, Nueva Esparta, Venezuela.

crisdiaz@ix.netcom.com

ABSTRACT.—A survey of Porifera biodiversity and distribution w as done in three islands of the Bocas del Toro region, Panama. Fourteen sites were explored using mask and snorkel, and occasionally scuba equip­ment, 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 m ost of the sponge species, w h ile few species were present at all sites visited. Approximately 23% (30 species) of the diversity encountered was w idely distributed. The species com position of the m ost typical or common sponges from reef, seagrass beds, or mangrove habitats, agrees with that of other Caribbean sites. An appendix w ith a field guide to the species encountered in this study is presented.

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

In t r o d u c t i o n

Sponges are among the most diverse and abundant inhabitants of benthic tropical communities such as coral reefs and man­grove systems (Riitzler and Feller 1987; Diaz and Riitzler 2001). Coral reef sponges in the Caribbean usually constitute the most diverse benthic group, 100-200 spe­cies, while other conspicuous coral reef groups such as hexacorallians or octocoral- lians are represented by 50-70 species (see Diaz and Riitzler 2001 for a review). In sub- tidal mangrove habitats, sponges, along with algae, ascidians, anemones, and hy- droids, are among the most conspicuous epibionts (Litter et al. 1985; Riitzler and Feller 1987, 1996; Calder 1991a; Ellison and Farnsworth 1992; Riitzler et al. 2000; Good- body 2000; Macintyre et al. 2000). Sponge species richness from mangrove systems in the Caribbean ranges from a dozen to 150 species, depending apparently on a combi­nation of physical factors, such as water turbidity, salinity, wave exposure, sub­strate availability, and tidal ranges, and biological factors such as competition, lar­val supply, and predation (Calder 1991b;

Litter et al. 1985; Taylor 1986; Ellison and Farnsworth 1992; Riitzler 1995; Riitzler et al. 2000; Wulff 2000). Despite the ubiquity of this group in Caribbean ecosystems, its diversity and relative abundance remains poorly understood for large parts of the re­gion. Whenever unexplored geographic ar­eas are studied or particular sponge taxa (genus, family, or order) are revised, a con­siderate number of new species is discov­ered. Examples of this trend are among re­cent work carried out in Jamaica (Lehnert and Van Soest 1998, 1999), and Belize (cer­tain taxa such as Chalinidae [de Weerdt et al. 1991; de Weerdt 2000]; or the Axinelli­dae [Alvarez et al. 1998; or Mycalidae Hajdu and Riitzler 1998]), and on the sys­tematic revisions of the familiy Chalinidae, Haplosclerida, by De Weerdt (2000), or the family Axinellidae by Alvarez et al. (1998).

The archipelago of Bocas del Toro is lo­cated on the western Caribbean shore of Panama and comprises more than 68 is­lands and mangrove keys. Knowledge of its shallow water sponge fauna is limited to two major studies: Guzmán and Guevara (1998,1999), and Guzmán (2003), who list 63

465

466 MARÍA CRISTINA DÍAZ

sponge species from open reef habitats, and Nichols and Barnes (unpublished) who reg­istered 86 taxa from reefs, mangroves, and seagrass beds in the Bocas region, includ­ing 12 taxa identified only to family and 11 identified only to genus. The results add up to approximately 79 species of sponge spe­cies reported from the area. During August 2003 the first Smithsonian Marine Science Network workshop took place at the Bocas del Toro Station-Smithsonian Tropical Re­search Institute (STRI). The goal of this workshop was to increase our understand­ing of the diversity and distribution of poorly-known marine taxa. The present study summarizes the results of a survey of the diversity and distribution of sponges from vario u s shallow w ater hab ita ts around this region's islands and presents a field guide to facilitate the identification of the most common sponges (see Photo­graphic Indentification Guide on this vol­ume).

M a t e r ia l s a n d M e t h o d s

Twelve sites in the Bocas del Toro Archi­pelago were surveyed by mask and snorkel during August 3-12 2003 (0-20 ft deep), and

two sites were surveyed by scuba-diving in August 2004 (15-60 ft depth). Table 1 lists the coordinates as well as the type of ma­rine community explored during each visit. Surveys consisted of swimming for ap­proximately one hour and registering the occurrence of each sponge species. When a sponge was encountered it was photo­graphed and a small fragment was col­lected, labeled, and transported to the labo­ratory in a cooler. Once in the laboratory, the specimens were fixed in 10% formalin (seawater) and within a week transferred into ethanol 70%. If a species was not rec­ognized immediately, a taxonomic study was performed in the laboratory, using a compound microscope to examine skeleton structure in dried hand sections cleared in Permount medium . Spicule types were de­termined after dissolving a fragment (2-4 mm3) in concentrated household bleach (5% sodium hypoclorite). The samples taken are kept in the Bocas del Toro, STRI, marine station.

A qualitative estimate of the abundance of each sponge species was carried out while surveying each site. Species were classified in four categories: very common

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

Locality Latitude-N Longi tude-W H abitat type

Crawl Key 1 9°15' 82°9'030" Rhizophora mangle fringe (3 ft deep), contiguous to shallow reef

Eagle Park-SW 9°19' 82°13' R. mangle fringe (6 ft deep) and seagrassSolarte Island

Solarte 1 9°17'030" 82°10'020" R. mangle fringe (6 ft deep) on internal lagoonSolarte 2 9°17'987" 82°12'404" R. mangle fringe (6 ft deep) on canal betw een islandsSolarte 3 9°17'208" 82°11'650" R. mangle fringe (3 ft deep), contiguous to shallow reefCraw l Key Canal 9°15'050" 82°07'631" Shallow reef (20-30 ft deep)

Millepora and Porites dom inant coralsColón IslandPunta Caracol 9°22'627" 82°18'117" R. mangle fringe (3 ft deep), contiguous to shallow reefSTRI Point 9°21T69' 82°15'528" R. mangle fringe (3-6 ft deep), contiguous to shallow reefITEC, Colón Is. Shallow reef. (5-15 ft deep)Big Bight. 9°22'184' 82°16'586" R. mangle fringe (6-9 ft deep) along a lagoonSwan Key 9°27'198" 82°18'024" Shallow reef. (5-15 ft deep) Strong surge and currentsPastores Island 9°14'332" 82°19'.968" Shallow, extensive, Porites reefA driana 's Key* 9°14'021" 82°10'023" Coral reef associated w ith nearby keys, (10-60 ft deep)

H igh coverage of stony coral, algae and spongesCristobal Island 9°15'379" 82°14'136" Fringing reef dom inated b y Agaricia, Porites and large

south* clum ps of red alga Amphiroa rigida var. antillana, specially below 20-30 ft deep. Large am ount of fine sedim ents along the reef profile (10-55 ft deep).

COM MON SPONGES SHALLOW MARINE HABITATS 467

(more than ten specimens seen), abundant (6-10), few (2-5 specimens), and rare (one specimen).

R e s u l t s

Sponge b iod iversity

One hundred and four species were en­countered in this study, of which 41 are new for the area (Table 2). As a result, sponge diversity known from the Bocas del Toro region reached 120 species. Four of 12 species identified only to genus are consid­ered new species and are currently being described. These species include two from the order Haplosclerida (H aliclona sp. and ^.Xestospongia sp.), and two from the order Poecilosclerida (M y c a le (P aresperella ) sp., and Iotrochota sp.).

H aliclona sp., and ?X estospong ia sp. (see illustration in the Photographic Identifica­tion Guide; this volume) are abundant on hard substrata of shallow reef habitats, and both posses endosymbiotic filamentous cyanobacteria. Io trocho ta sp. is found in m uddy areas of certain deeper reefs (>40 ft). M yca le (Paresperella) sp. is a common species but only found on mangrove roots. It represents the first find of a whole speci­men of this sub-genus in the Caribbean. Hadju and Riitzler (1998) describe the dis­tinctive spicules of this subgenus (serrated sigmas) found contaminating a specimen of M yca le (A egrop ila ) c itrina , therefore assum­ing the occurrence of a species of this sub­genus in Belizean mangroves but lacking a complete specimen.

Twenty-four species are new record for the Panam anian sponge fauna. Half of these species (13) are exclusively found in­habiting mangrove roots or peat (H aliclona p isca d a eren sis , H . cu racaoensis, H . m u c ifi­brosa, H . verm eu len i, H . tubifera , C halinu la zeae, C. m olitba, Terpios m ang laris, T ethya aft. seychellensis, M yca le m agnirhaphid iphera , M . carm igrop ila , M y c a le undescribed species, and Geodia papyracea). Eleven inhabit pre­dominantly reef environments (A m p h im ­edon v ir id is , C a llyspong ia fa lla x , C. pallida, H aliclona vansoesti, O ceanapia oleracea, S p i­ra s tre lla h a r tm a n i, C la th ria ech in a ta , plus three undescribed species of H aliclona X e s­tospongia sp., and Iotrochota sp.), and the

calcareous sponge C la th r in a p r im o rd ia lis that can be found either in reef or man­groves. Fifteen species are first records for Bocas del Toro region but were reported previously for other localities in the Pana­manian Caribbean. Such is the case of C he­lo n a p l y s i l l a e r e c ta , V e r o n g u la r e i s w ig i (Guzmán 2003), D ysidea etheria, H aliclona m anglaris, C ribochalina va scu lu m , O ceanapia pelta ta , N ip a h tes caycedoci, Sp irastrella m ollis, L issodendoryx colom biensis, E ry lu s fo rm o su s , and H alichondria m agniconulosa (Clifton et al. 1997; Wulff 2000).

Sponge species d is tr ib u tio n and rela tive abundance

Most species (58%) occur in one to four sites, while only one species (N ip h a te s erecta) occurs at all fourteen sites surveyed (Table 2). Thirty of the 126 species (23%) occur at five or more sites. A p ly s in a cauli­

fo rm is , A. fu lv a , A iolochroia crassa, V erongula rig id a , Irc in ia f e l i x , I. ca m p a n a , N ip h a te s erecta, A m p h im e d o n com pressa, A. erina, X es­tospongia caycedoci, X . m u ta , X . p roxim a , X . carbonaria, X estospong ia sp ., Iotrochota b iro tu ­lata, M yca le laevis, M . laxissim a, and Cliona d elitr ix were common in reefs sites, while H y r tio s p ro teu s, Spong ia p ertu sa , H aliclona im p lexiform is, H . m ang laris, H . curacaoensis, H . tubifera , C halinu la m olitba , L issodendoryx isodyctia lis , C lathria schoenus, and Tedania ign is were common in mangroves. Three species—N ip h a tes erecta, Scopalina ruetzleri, and C hondrilla n u c id a—may be common in both reef and mangrove habitats. The ma­jority of the species (74 of 126) were at least in two of the three habitats surveyed, while 28 species occur exclusively in mangroves and 25 species in reefs (Table 2).

D is c u s s io n

After surveying the shallow water habi­tats of three islands in Bocas del Toro re­gion in a relatively short time, the known sponge diversity increased 34%. This dem­onstrates the importance of this kind of ex­ploration to further our biodiversity knowl-

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edge. Three m ain fac to rs m ay 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. Second, the taxonomic expertise of the ob­server might have aided in the discrimina­tion of species in this difficult group. Third, the general patchiness of sponge distribu­tion in mangrove systems and the rarity of most species (Diaz et al. 2004) could also contribute to the increase of species num ­bers as new areas were surveyed. Another consideration is that more species are de­tected if large areas are surveyed by unre­stricted swimming along suitable substrata, rather than being restricted to line transects placed along the mangrove fringe (Diaz et al. 2004).

It is quite possible that estimates of sponge diversity in Bocas del Toro region will double once there is data for deeper coral reefs (>20 ft), mangroves surrounding other islands (e.g., Cayo de Agua and Bas­timentos), the bordering mainland section, and new hab ita ts (cryptic reef areas, m uddy bottoms, and coral rubble). Fair comparison of this region's diversity with that of others in the Caribbean realm must await the results of further exploration. With the collaboration of colleagues, four species, that seem new to science, are being studied to determine their generic and spe­cific status. It is expected that more unde­scribed species will be found as the region is examined in more detail.

The most species-rich sponge genera in Bocas el Toro reef habitats are A p ly s in a (Aplysinidae, Verongida) and X estospongia (Petrosiidae, Haplosclerida), with four and five species, respectively. In the m an­grove, the most diverse genera are H aliclona (Chalinidae, Haplosclerida), and M y c a le (Mycalidae, Poecilosclerida), with six and nine species, respectively. These genera are also very diverse in other Caribbean loca­tions. In Bocas del Toro, the petrosiids, in particular X estospong ia , are highly diverse and abundant; it harbors typical Caribbean species (X. m u ta , X . carbonaria, and X. su b ­tr ia n g u la r is ) as well as species described

474 MARÍA CRISTINA DÍAZ

from Colombia, and observed mostly in the West-Central Caribbean (X. rosariensis, and X. proxim a).

Evaluation of species distribution at Bo­cas, although of qualitative nature, con­firms certain trends reported previously. The patchiness of species d istribution, where a large part of the species are seen only once or twice in a survey, was re­corded by Riitzler et al. (2001), Diaz et al. (2004), and Alcolado (unpublished data). The species spectrum of the most typical or common sponges from reefs, seagrass beds, or mangrove habitats conforms well with that of other Caribbean sites such as Cuba, Jamaica, Bahamas, Belize, Panama, Colom­bia, and Venezuela (Riitzler 1969; Wieden- mayer 1977; Zea 1984; Farnsworth and El­lison 1996; Riitzler et al. 2000; Wulff 2000; Diaz et al. 2004; Alcolado unpublished data).

The most common species on reefs habi­tats are A p ly s in a fu lv a (11 sites), A. caulifor­m is (8 sites), Aiolochroia crassa (7 sites), Ir­cinia cam pana and I. fe l ix (seven sites each), X carbonaria (eight sites), Iotrochota b iro tu ­lata (8 sites) and N ip h a tes erecta (14 sites), which are common throughout the Carib­bean and the Gulf of Mexico. Similarly, mangrove sponges such as H aliclona tu b i­fera (6 sites), H y r tio s p ro teus, H aliclona im ­p lex ifo rm is, and H aliclona m ang laris (7 sites), T edania ig n is , and Io trochota b iro tu la ta (8 sites), and C lathria schoenus (10 sites) are the most widely spread in Bocas del Toro, and in other Caribbean mangroves as well.

In this area one surprising pattern of spe­cies distribution is the high incidence of typical reef species, such as M yca le laevis, Irc in ia fe l ix , Irc in ia cam pana , A m p h im e d o n com pressa, and N ip h a tes erecta, on roots of the red m angrove (R h izo p h o ra m a n g le ). This situation might be a consequence of the proximity of reef and mangrove habitats. A similar pattern is found in mangrove ponds in the Pelican Cays, central Belize, where an important portion of sponges growing on the roots are typical reef species (Riitzler et al. 2000; Wulff 2000). According to Wulff (2000) in Pelican Cays, the closeness of the reef and the presence of spongivorous reef species on the mangrove fringe contribute to this phenomenon. Overall, sponges in

the Bocas del Toro region are similar to those at other Caribbean sites, sharing the identity of the most common and wide­spread species, as well as the patchy distri­bution of most species at local scales.

A c k n o w le d g m e n ts .—The author thanks the Smithsonian Institution-Marine Science Network for support to attend the first Bo­cas del Toro Biodiversity workshop held in August 2003. Dr. Rachel Collin (director) and Dr. Gabriel Jacóme (scientific coordina­tor), and support staff of the STR-Bocas del Toro Marine station are acknowledged for helping to accomplish this work. Dr. Klaus Riitzler assisted with taxonomic advice, and Drs. Scott Nichols, Penny Barnes, and Pedro Alcolado provided unpublished data.

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