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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.
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-
TABL
E 2.
Spon
ge
spec
ies
dist
ribut
ion
and
rela
tive
abun
danc
e (1
= ve
ry
com
mon
, 2
= ab
unda
nt,
3 =
few
, 4
= ra
re)
in 14
studi
ed
sites
in
isla
nds
from
the
Boca
s de
l To
ro
regi
on,
Pana
ma.
Col
umn
lege
nds:
Rf
= R
efer
ence
s (1
. N
icho
ls
and
Bar
nes,
unpu
blis
hed
data
; 2.
Guz
man
20
03;
3. Pr
esen
t st
udy)
. Si
tes
are
orga
nize
d ac
cord
ing
to the
m
ajor
ha
bita
t (s)
su
rvey
ed:
EP =
Eagl
e Pa
rk,
SI =
Sola
rte
site
1, S2
= So
larte
sit
e 2;
AK
= A
dria
na's
Key
, CK
2 =
Cra
wl
Key
Bank
, SK
= Sw
an
Key
, Ti
= T
INC
, Cr
1
= C
risto
bal
Islan
d re
ef;
CK
1 =
Cra
wl
Key
1, PI
= Isl
a Pa
stor
es,
SP =
STRI
Poi
nt,
PC =
Punt
a C
arac
ol,
BB =
Big
Bigh
t, an
d S3
= So
larte
sit
e 3.
(*fo
und
only
on
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COM MON SPONGES SHALLOW MARINE HABITATS 469
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COM M ON SPONGES SHALLOW MARINE HABITATS 471
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COM M ON SPONGES SHALLOW MARINE HABITATS 473
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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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