The roof was covered by T~rl~nstraen nlrrecr, the walls mostly by Th)~r~oscv~ ... The floor is covered by shingle, with rubble and boulders. In the inner part of the ...
SHALLOW EPIBENTHIC COMMUNITIES OF ILHA DO SAL (CAPE VERDE ARCHIPELAGO, EASTERN ATLANTIC) CARLA MORRI, BANCHI
I
RICCARDO CATTAENO-VIETTI, CIANFRANCO SARTONI & C. NIKE
Molicar~lrs, in an overhanging portion of the cliff at about 3 In Mulloidiclztlz~~s riznr-tirricus, Sccrri~s hoefleri. to 8 m. Below 8 m, the slope became less steep Selcrre tlorsalis, Sirnill>nrrrinIlerriznni, Sl~arisoriza and the biotic community was composed by cretet~se.S. cf strigatrun, Ste/~hnr~ole/~is hisl~idus Milieporn sp, Megabalnr7us nzoricrrs, Dictyota and Tlznlassornn par'n. dichotornrr and the a~ooxanthellate coral The coral-dominated community nearly B n l a r ~ o p l ~ ~ ~ l lsi pn (Fig. 2c). Mcrrtllasterins disappeared after 15 m depth. the only hard coral glcrcinlis was seen feeding on Megnbalar~us.This still found at 2 0 m being P. nstreoides. It was community remained similar down to about 2 0 In, replaced by a colnlnunity dominated by the where P. corzcinrrcrt~zreplaced D. diclloton~nas the antipatharians Cirrilxrthes (Sticlzopathes) main erect alga. Pe).ssotznelin, crustose corallines leutker~i. Arltipnthes ~vollnstorli (Fig. 2a) and and a few colonies of Porites nstreoicfes were A. spinescer~s (Fig. 2b) and the gorgonians present on large boulders toward the base of the cliff. Au1ostor~zrr.s .stri,qosu.s. Ceplrnlol~l~oli~s E~rrzicella pc~llillifem,E. gr.anlrlat~~, Leptogorgia cal~verdensisand L. gnirri, the latter on the sandy tner~io/~s, Ah e~d~jZrlf Izoefleri, C1irorni.s lrrhbocki, floor below 30111. On the whole. the most Sirnill~~rrrncrhernlnrli, S~~urisotlrt~ creterzsc. and abundant species in this com~nunity were S. ruhr-illirzrle were the commonest fish. A large cave opened at the base of the L. cnl~verdensisand C. leutker~i.The sessile fauna also included encrusting sponges and the stalked underwater cliff, between 19 In and 27 n~ depth. colonial ascidian Distaplio corollcz. The spiny The roof was covered by T~rl~nstraen nlrrecr, the walls mostly by Th)~r~oscv~7l1us ~~z~rr,yi~zcrtr~~. On the lobster Pnr1u1irr.r.r regirrs and the sea-star Ophidinster ol~lzidinrr~r.~ were noticeable among floor, Lcptogorgin capverderisis and Arrtil~~lthes the motile species, Aulostor71rr.s str.igos~~s, Chrornis sl>irze.scerzs (Fig. 2b) were present amidst blocks lcrbbocki. El)iriephelu.v mnrgirzntr~.~,Fi.rtultrria covered by encrusting sponges. Several fish
species were observed at the entrance of the cave, Chnetodori rnarcellne (Fig. 2d), including Diplodus fnsciatus, Holacanthus africanus and Mycteroperca fusca.
The community of the supralittoral pool was distinctly zoned according to a spatial gradient perpendicular to the sea. On the seaward wall, encrusting corallines and the sea-urchin Arbacin l i x ~ l l awere the most conspicuous organisms. The bottom of the pool, reaching 4 m at the seaward side of the pool and less than 1.5 m at the landward side, contained blocks covered w ~ t ha lawn of filamentous green algae. A rich motile fauna was present among the blocks, such as the sea-star Coscinasterins tenuispina, the carnivorous gastropods Purpura nodosa, Thais hemastoma and Latirus triserialis, and many hermit crabs (probably Calcinus talisrnani) within Littoritla and Mot~odotitashells. Two species of Littorina occurred at the landward side of the pool L. striata above the water line, L. punctnta ,i few cm depth. Among the fish, Lipophys caboverdensis and Parabletiniirs ~mrvicorniswere also abundant on the bottom of the pool, whereas Abudefduf hoefleri and juveniles of Thalassoma pavo and Oblada melanura swam around in midwater. WESTCOAST. SHELTERED SITE A small inlet, named Jorge Fonseca, opens in the middle of the coast of the large Baia de Mordeira (Fig. 1). The inlet is nearly 60 m long and wide and less than 1 m deep for most of its extension. The floor is covered by shingle, with rubble and boulders. In the inner part of the inlet, the community was dominated by zooxanthellate corals. Siderastreu radians, Porites astreoides and especially P. porites were the most abundant species, but Millepora alcicornis and Favin f r a g ~ l m were also common. Another important species was the green alga Cnulerpa sertulariodes, especially in more sandy areas. Near the seaward opening, where waves break into the inlet, massive construction by Millepora sp has formed a miniature barrier reef. Leeward of this reef, in slightly deeper water ( I to 2 m),
S. rcldians has formed slabs up to I m wide. F. fragum and P. astreoides, but no Porites porites, were also present. The brown alga Dictyota covered the vertical side of the largest
blocks. 'ITE
The southern coast of Sal is sandy and a long beach borders the largely open bay called Baia de Santa Maria (Fig. 1). The sea floor is sand and gravel to about 4 m. At this depth, a line of beach rock was found nearly parallel to the coast. Another similar line occurs at about 9 m. At both depths, the beach rock was colonised by pinnacles (stout erect and branched growth forms) of Millepora up to 25 cm high and 50 cm wide. Porites astreoides and Siderastrea radians were also abundant. The former preferred the highest blocks, the latter formed large colonies at the base, at the boundary between rock and sand. The sandy floor was covered by a meadow of Udotea flabellurn, whereas the most conspicuous invertebrates on sand were the sea-urchin Kotuln deciesdigitata and the gastropod Stro~nbuslatus. Fish were mainly represented by Tueniura grabata, A~llostomus strigosus, Pseudupeneus prayetzsis, Holncnnthus cfricatius, Ahudefduf saxutilis, Sparisoma cretense, S. cf strigatu~nand Acarzthurus monroviae.
DISCUSSION Epibenthic communities at Sal exhibited a clear zonation pattern, in most cases consistent with that already described for the Cape Verde Archipelago by OTERO-SCHMITT (1993, 1995). Apart from minor differences in relation to exposure, the change of flora and fauna with depth was sharp, thus allowing recognition of at least three main bionomic zones (BOUDOURESQUE & FRESI 1976): 1) the littoral, with littorinids, chthamalids and encrusting corallines; 2) the shallow sublittoral, with corals and algae; 3) the sublittoral below 20 m depth, with gorgonians and antipatharians.
The littoral communities fitted in well with the Brook (=A.subpirlrrata Gray. not Ellis and scheme outlined for West Africa by L.AWSON Solander). (1966). but the eulittoral (or midlittoral) zone Many authors have already underlined the showed Atlantic-Mediterranean affinities. The composite biogeographic character of the Cape dominant organism was Chtlzarncilus stellatus, as Verde marine biota, and also expressed divergent in most western European coasts and throughout views on this topic in some instances. BRIGGS the Mediterranean, in the Canaries (HAROUN ( 1 974), who based his analysis rnostly on fish, TABIZ.I\UE et al. 19841, Madeira (BIANCHI et al. in maintained that the relationships of the Cape press) and the Azores (MORTONet trl. 1998). In Verde Archipelago are more tropical than warmtruly tropicill West Africa shores, C. stel1crtu.s is temperate. At Sal, we observed strictly tropical replaced by Chfhtlrna1u.s denfarus (LAWSON fishes, such as the butterflyfish CI7~retodonand the 1966). angelfish Holactznthus, swimming together with In the sublittoral, analogies can be found with warm-temperate breams, such as Ohlada and both warm-temperate and tropical zones. The Diplodus (Fig. 2d). Within groupers. biotic change observed at Sal around 20 m depth Ccphalopholis taer1iop.s has a clear tropical corresponds with the transitions from infralittoral affinity, whereas E~~irleplzelusriztrrgirznt~~.~ and to circalittoral in the Mediterranean (PERES1967) Mpctc.rol~ercc~ f~~sctr are two Atlanticand from reefal to subreefal in coral reef areas Mediterranean species. Algae and corals give the (DONE1983). Cape Verde Archipelago a Caribbean affinity, as The shallow sublittoral at Sal shares with the mentioned above, but infaunal communities Mediterranean the dominance by algae. with the (LONGHURST 1959), sponges (VAN SOEST 1093) tropics the abundance of zooxanthellate corals. and, in part, brachiopods (LOGAN1993) sngg~s: Both groups give the same biogeographic picture: an Atlantic-Mediterranean affinity. Cape Verde corals are Caribbean in origin VAN SOEST (1993) observed that these rival interpretations centre on the classical (LABOIZEL 1974) and algae have a distinct East disagreement between dispersal and vicariance. V A N REINE& American affinity (PIZUII'HOMME VAN DEN HOEK1988). Clearly, dispersal capabilities vary enormously Dominance by gorgonians below 20 m is a among individual taxa and this may well explain common feature in the Mediterranean Sea, but the different biogeographic patterns observed in antipatharians live deeper in the circalittoral the different groups. However. the zonation seen (PEIZES1967). Communities dominated by co- at Sal suggests that differences might also originate according to the depth zone. In the occurring gorgonians and antipatharians are usual in subreefal habitats in tropical seas. However, the inf~.alittoral zone we found more pronounced gorgonian species we found at Sal exhibit a tropical West Atlantic affinities, whereas in the midlittoral, and perhaps in the circalittoral, the eastern Atlantic affinity and closely related species belonging to the siune genera occur in the affinity was rather Atlantic-Mediterranean. As LONGHURST (1962) pointed out in his Atlantic-Mediterranean region (GRASSHOFF 1992). The biogeographic affinity of Cape Verde review of the oceanography of the Gulf of Guinea, tropical waters form only a thin surface layer in antipatharians is unclear. rnostly because of doubtful species identification. The two species the Eastern Atlantic, whereas deeper waters are that we call Ariti/~t~thes wollastorzi (Fig. 2a) and colder because of the influence of the Canary A. sl~irle.scerls(Fig. 2b) in the present paper, were Current. According to LABOREL( 1 974) this warm first tentatively referred to their west west- water layer is not deeper than 20 to 30 m and this seems in perfect accord with our findings of a Atlantic relatives A. Darbni1errsi.s and A . tcrrzncetLlrl7. respective]y (MORRI& BI.ANCHI sharp bionomic and biogeographic change below 1995). Several different species, however, may be 20 in depth. Midlittoral coinmunities, on the other included under the name Arlti11athe.s \vollastorzi hand, may be more directly influenced by the
atmospheric climate, which in the Cape Verde parrotfish apparently similar to 5. strigarum from Archipelago is relatively "temperate" (BAEZ& St. Helena and Ascension, which should not occur in Cape Verde waters. SANCHEZ-PINTO 1983). All the above discussion clearly highlights the GOLIKOV & SCAI~LATO (1968) related biogeographic differences within bottom need for further taxonomic studies on the Cape biocoenoses of Japan and Okhotsk seas to the heat Verde marine biota. Better taxonomic knowledge exchange between sea and atmosphere and the may allow biogeographic analyses to be done, not position of the layers of different water masses. only taking into account whole species llsts, as is Our observations at Sal might lead to similar usually done, but also according to depth zones. A conclusions, but the alternation of tropical and bionornic approach, relating species distribution warm-temperate affinities with depth derived to the characteristics of water masses. may help essentially from the analysis of the sessile elucidate the complex biogeographic pattern of the Cape Verde marine biota. component of the epibenthic con~munities. 'The motile Fauna gave contradictory indications: for instance, the most abundant starfish were ACKNOWLEDGEMENTS Marrhasterias glacialis, a temperate species, in the "tropical" infralittoral and Ophidiaster ophidial~us, a warm-water species (TORTONESE The Cabo Verde expedition was organised within 1985), in the colder circalittoral. For motile the research pro-ject "Gli Cnidari del benthos species, however, the zonal pattern of the marino: ecologia e biogeografia" (60% MURST). reproductive stages may probably be more Staff of "Dive Cape Verde" at Santa Maria (Sal) helped with logistic and diving assistance. important than the distribution of the adults. (Amsterdam), F. Gentil A major problem with both sessile and motile G. J. Boekschoten species is the persistent lack of sufficient (Roscoif), M. Grasshoff (Frankfurt), P. KnightJ. Laborel (Marseilles), knowledge on the Cape Verde marine Jones(Swansea), biodiversity. Even in a study like the present one, A. Logan (Saint John). F. Monniot (Paris), based only on the most conspicuous and common J. Otero-Schmitt (Santiago de Compostela), species, species recognition turned out to be a M. Pansini (Geneva), W. Prud'Homme van Reine (Leiden), F. Keiner (Cascais), P. Wirtz (Madeira) hard task. Most invertebrate groups require extensive study. Among Cnidaria, for instance, and H. Zibrowius (Marseille) gave information only gorgonians (GRASSHOFF 1992) and and advice. A. Logan and P. Wirtz also revised & BEST 1988) the manuscript. Special thanks are due to H. Rost hermatypic corals (BOEKSCHOTEN may be considered adequately known. The Martins (Horta) for her patient editing. Part of this species we called Balanophyllia sp (Fig. 2c) work was done under the aegis of the Research seems to correspond with what CHEVALIER (1966) Project "MACMED" (comparing marine benthic erroneously called Balnnophjllia italica and is communities and zonation in MACaronesia and probably new to science (H. Zl8ROWIUS ill the MEDiterranean) of the Marine Environment Research Centre of La Spezia. litteris). Milleporc~ sp. is possibly a different species from or simply an ecotype of M. alcicort~is(MORRIRc BIANCHI1995). Actit~ia REFERENCES sali was only recently distinguished from the A. equilln common Atlantic-Mediterranean 1983. Islus rle,filrgo y (MONTEIROet a / . 1997). and we have already BAEZ,M. & L. SANCHEZ-PINTO agurr. Carlurias, Azores, Madeira, Scrlvi~jes,Cabo mentioned difficulties with antipatharians. Fish Verrlr. Mric~~ror~esin. Edirca. Las Palmas de Gran are better known (SERETRc OPlC 1986; REINEI< Canaria. 183 pp. 1996; DEBELIUS1997). Nevertheless, we met BIANCHI. C. N.. S. COCITO,C. MORRI& S. SGORRINI problems with several families. We tentatively 199 1. Rilevamento bionomico subacqueo. Pp. used the name Spari.snma cf strigarum for a 67-83 in: ABBIATI, M. (Ed.) Lezior~i (/el corso
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