A comparative survey of reef fISh es in and Pacific

Peter C. Phillips Escuela de Ciencias Biológicas, Universidad Nacional, Heredia, Costa Rica. Mikie 1. Perez-Cruet Associated Colleges of the Midwest (ACM), San José, Costa Rica. Present address: University of South Florida, Tam­ pa, Florida 33604, U.S.A.

(Received fOI publication November 11, 1983)

Abstraet: Fishes from five transects in Manuel Antonio National Park (pacific) and seven transects in Cahuita National Park (Caribbean), Costa Rica were censuscd using a rapíd visual technique from February to April 1982. Cahuita, with 49 species, had a higher overaU species diversity, whereas more individuals were usually seen on the Manuel Antonio transects, where 39 species were counted. Transects with similar habitat com­ plexity showed highest PS values (approximately 60%). Wrasses (Labridae) and damselfish (Pomacentridae) dominated in both the Pacific rock reer and the alribbean reef study areas. Individuals up to 7.5 cm usually dominated on transects, but an ¡ncrease in size was observed as the habitat complexity increased. Diversity was low at Cah�ita in relation to other Caribbean reer areas, probably due to the declining state of the coral reer. Thc specics diversity valucs at Manuel Antonio are within the range of values found in Cen­ tral American Pacific estuarine and tidepool studies.

Rock reefs and coral reefs morphologically tional Parks, Cahuita (Caribbean) and Manuel present very similar habitats of considerable Antonio (Pacific) during February, March and niche diversity harboring a great variety of fish April 1982. assemblages. Since the structure of a reef is very irregular and complex, most standard STUDY AREA methods of fish sampling (nets), are inadequate (Alevizon and Brooks, 1975). Other melhods Cahuita National Park is situated 35 km which involve lhe use of rotenone (Srnith, soulh of Limón on the Caribbean coast of 1973) or explosives (Talbott and Goldman, Costa Rica (Fig. 1). Its coral reef extends to 1973) are efficient for collections but are 300 m from lhe shore around Cahuita Point frequently incompatible with the goals of areas and is 4 km long. The nearshore currents in lhe set aside for conservation purposes. In order to area flow from north to south. Risk el al. overcome this problem, various visual census (1980) noted that the reef at Cahuita is in a techniques have been developed which require state of decline probably due to heavy sediment no direct collection of fishes (Brock, 1954; loading in the rivers to lhe north of lhe park Hobson, 1974; Jones and Thompson, 1978; during heavy rains aand contamination from Gladfelter el al. , 1980; Sale, 1980). One visual the port city of Limón. In a study by Wel­ method places transects over the study areas tington (1974), 34 species of coral were found and by repeatedly taking sightings of the on the Cahuita reer. abundance of fish species a10ng a transect, an Manuel Antonio National Park, on lhe accurate estimation of fish inhabiting the Pacific coast (Fig. 2), is characterized by rock transect area can be determined. ctiffs interspersed by sand beaches. Allhough The present study utllizes a rapid visual there is no extensive living coral reef at Manuel census technique to estimate the number of Antonio there are numerous rack reefs and fish species, their abundance and size distri­ occasional coral fo nnations. bution in relation to habitat complexity for The transects in bolh study areas were a series of transects at two Costa Rican Na- numbered according to degree of habitat 96 REVISTA DE BIOLOGIA TROPICAL

f)

Internol eresl PACIFIC OCEAN . �. .. ,. \ . .• .. ... • ......

CAR'8B�AN SEA

o 500 � m o 500 S � m ! Fig. 1. Transects in Cahuita National Park (Caribbean), Fig. 2. Transects in Manuel Antonio National Park Costa Rica. (pacific), Costa Rica.

...... _". ---. - . . F?====. -:i

Fig. 3. Overlap in the ichthyofaunal species compo­ Fig. 4. Overlap in the ichthyofaunal species composi­ sition for the seven transects in Cahuita. tion fOI the five transects in Manuel Antonio.

complexity, Transect I being tbe least complex. Transeet 3: Inside part of tbe intemal crest,

Here, habitat complexity refers to an area 'S composed of c1umps of coral up to 1 m in topographical irregularity. height covered by numerous small patches of Figure 1 indicates the seven different hab· vegetation and algae. The most corumon itats which were sampled at Cahuita: coral s were Diploria sp., Porites sp. and Sideras­ trea sp. Between the coral fo rmations were flat Transeet 1: Located inside the internal reef areas of sand and coral rubble. Deptbs ranged erest, this area was characterized by loose from 0.5 to 1.5 m. dead coral rubble free of vegetation. The bottom was flat witb a depth from 0.5 to 1.5 Transee! 4: Northem part of the reef erest m. loca!ed on tbe poin!. There were large coral fo rmations to 2 m in height. There was also Transect 2: Inside of the reef crest of much vegetation and small patches of the Cahuita point and surrounded by extensive ; Siderastrea ra dians, MilIepora sp., fields of turtle grass (Thalassia sp.), thls flat Agaricia sp. and Diploria clivosa. This area area was covered by coral formations up to aeted as a buttress against tbe strong incoming 0.75 m in height and small patches ofthe corals current and waves. Mon tastrea sp., Porites spp., and MilIepo­ ra sp. Transee! S: This small coral reef in tbe PHI LLIPS & PEREZ-CR UE T: Reef fishes of Caribbean and Pacific Costa Rica 97

shallowest part of the lagoon include d the of the island and the rack outcrapping, but the following corals� Acropora sp .. Siderastrea sp., rocks in the channel were generally of mediurn Po rites sp., Diploria sp., and Corgonia sp. The - size. This area was exposed to the waves surrounding lagoon was of coral rubble. sand entering the bay and Ihere was usual!y a strang beds of Thalassia sp. and large blocks of coral. current. The water depth was from 0.5 to 6 m. This small reef was from 1 to 2 m in depth. Transect s: lnside part of a rack point in Transect 6: On the outer side of the intemal Manuel Antonio Bay. There was one large crest and composed of fields of branching coral head which, along with Ihe racks, ranged o::: Acropora sp. surraunded by areas of MilIepora in height from 0.5 to 2 m. During low tide Ihis sp., Siderastrea radlans, Montastrea caver­ area was protected from the incoming waves by nosa, Pon' tes astreoides, Agarióa agaricites, the rock pojnt, which was uoderwater at high Diploria stn'gosa and Mo ntastrea annualus. tide. The water depth was fram 0.5 to 5 m. This area 's irregularity provided a very complex habitat for fishes. The water depth was fram 0.5 to 3 m. MATERIAL AND METHODS

Transect 7: This most complex habitat was a Dne huodred foot transects were perma­ reef located between the internal and externa! nentIy identified with plastic jugs secured by crests. It was made up of large branching eord to the bottom. Fishes were sighted by the Acropora palmata that extended to heights of 4 junior author equipped with snorkeling gear. m. Also there were large heads of Diploria Eight sightings of each lOO foot transect stngosa, Siderastrea radians and Montastrea were taken irrespective of tidal stage and cavernosa and large patches of MilIepora sp., recorded 00 an underwater writing pad. Each Porites astreoides, Dichocoenia stokesii, Col­ transect included a field 1 m to either side pophyllia sp., Porites porites and Agarida and from Ihe bollom to Ihe surfaee. Photo­ agaricites. The water depth ranged fram O to 6 graphs of fish species that could not be im­ m. mediately identified were taken with a Nikonos Figure 2 indica tes the five different habitats 4A underwater camera. An effort was also which were sampled at Manuel Antonio: made to record the fish species that hid in caves or under coral during the day , sinee the visual Transect 1: This plain area was located oear technique is only accurate for diurnal species the beach in Manuel Antonio Bay and was (Broek, 1982). Therefore, the transect data characterized by smal! racks, pebbles and coral reflect relative rather than absolute abundance. rubble which were covered in sorne places by The use of snorkeling gear tends to reduce Ihe algae. The water depth was approximatley 4 m. amount of noise assoeiated wilh SCUBA and also facilitates the fish census in shallower Transect 2: Rock reer also in Manuel Anto­ waters (Chapman el al., 1974). It was noted nio Bay. The bOllom was of irregular racks that that smaller species seemed to be attracted to measured from a few contimeters to 0.3 m in Ihe bOllom sediment Ihat was raised during height. In between the racks there were sandy sampling. This allraetion of smaller fishes may areas and areas that were sparsely covered by have led to a bias of higher eounts of these low vegetation. smaller individuals. Upon completion of all sightings, eaeh N Transect 3: Roek reef on the calm east side (number of individual sightings per species of a small island in Puerto Escondido Bay. per transect area) was divided by 8 and Ihe Characteristic of this area were the medium­ respective standard deviation was calculated. sized raeks that ranged in height fram 0.2 Individual fish in each sighting were grouped to 1 m fo rrning nurnerous small caves. The into the following size categories; Graup water depth ranged from 0.5 to 2 m. 1, to 7.5 cm; Group 2, 7.5-20 cm; Group 3, 20.5-28 cm; Group 4, 28.546 cm; and Graup Transect 4: West side of the small island in 5>46 cm. Puerto Escondido Bay. This transect ran For each trans.ect. data on the fish species fram the island to a rack outcrapping in frant was analyzed as follows. Species diversity 98 REVISTA DE BIOLOGIA TROPICAL was calculated wilh an approximate equivalent individual sightings were made represente to the Shannon-Weiner Index: by 49 speeies and 21 different families (Tabl 1). The most speciose families were th n , butterflyfishes, Chaetodontidae spp. H' = C/N (NlogION. �nilogIO i) (6 ), th grunts, Pomadasyidae (7 spp.) and the dan where C equals 2.302585, N is lhe total number selfishes, Pomaeentridae ( 5 spp.). The bluehea of individuals at a transect area and Di is the wrasse, Thalassoma bifasciatum (1148) and th number of individuals in lhe ith species (Lloyd lhree spot damselfish, SIegas tes planifro>; el al. , 1968). High species diversity indicates (1156) were the two most abundant speci, the presence of a great number of species with and were observed on all transects. individuals evenly distributed among the species. The species diversity i5 low when few TABLE 1 individuals are concentrated among a few species. List offa milies and species and total number of The evenness component of diversity (J') is invididual sightings at Cahuita National Park calculated using lhe equation: (Caribbean), Costa Rica Acanthuridae J' = H'/lnS, Acanthurns bahianus 278 Acanthurns coernleus 397 where H' is the Shannon-Weiner Index calcu­ Apogonidae I Apagon maculatus 20 lated above, S is lhe number of speeies, and nS Blennüdae is lhe maximum possible diversity as defined by Ophioblennius atlanticus 75 Pielou (1966). An even distribulion of indio Carangidae viduals among speeies implies high diversity. A Caranx bartholomaei 41 Ca ranx fusus I maximum evenness value of 1 fOI an area Chaetodontidae occurs when a11 species are represented by equal Chaetodon capistratus 4 numbers of individuals. Chaetodon ocellatus 63 Finally, in arder to extraet more informa­ Chaetodon striatus 26 lion from the trends of the two indiees, the Holocanthus ciliaris 6 Holocanthus tricolor 1 pereent eontribution of lhe two most abundant Pa macanthus paru 3 speeies (D) at eaeh transeet area was summed. Clinidae In arder to measure the species composition Acanthemblemaria sp. 4 similarity between the transects, a percent Diodontidae similarity value was ealeulated by the formula: Diodon histrix Holocentridae Holocentrns rnfus 119 P P , PS = 100 (1.0·0.5 �i ia - ibl) My ripristis jacobus 51 �yphosidae Kyphosus incisor 144 where Pia is the number of individual s in the ith speeies in Transeet a divided by lhe total Labridae Bodianus rnfus 79 number of individuals in Transect a, and Pib i8 Halichoeres bivittatus 76 the same fOI sample b. Percent similarity Thalassoma bifasciatum 1148 ranges from O, when two areas being compared Lutjanidae have no species in common, to 100, when Lutjanus apodus 54 Lu tjanus griseus 1 the two areas have identical species and Lutjanus mahogani 52 amounts of individuals (Haedrieh, 1975). Ocyurns chrysurns 5 Dendrograms of the transeets in Cahuita and Mullidae Manuel Antonio were constructed using meth· Mulloidichthys martinicus 1 ods developed by Sokal and Miehener (1958) Pseudupeneus maculatus 17 Pepheridae for a better visual presentation of the similarity Pempheris schamburgki 28 in species composition between the transects. Pomacentridae Abudefduf saxatilis 647 RESULTS AND D1SCUSSION Chromis multilineata 1 Micropathodon chrysurns 372 Stegastes partitus 6 At Cahuita Nalional Park, a total of 7,582 Stegastes planifrons 1156 PHIL LIPS & PEREZ-CRUET: Reef fishes of Caribbean and Pacific Costa Rica 99

Pomadasyidae Mugilidae Anisotremus virginieus 9 Mugil curema 88 Ha emulon flavolineatum 394 Pomacanthidae Haemulon macrostomum 860 Holocan thus passer 6 Haemulon parrai 114 Pomacanthus zonipectus 6 Haemulon p/umieri 80 Pomacentridae Ha emulon sciurus 15 A bu defduf conc% r 57 Ha emulon sp. 334 Abudefdulf saxatilis 1159 Priacanthidae Microspathodon hairdii 37 Priacanthus cruentatus 63 .'t1icrospathodon dorsalis 610 Scaridae Stflgastes acapulcoensis 474 Scarus croicensis 460 Stegastes sp. 1160 Scarus taeniopterus 53 Pomadasyidae Spar;soma rubripinne 172 Anisotremus taeniatus 5 Sparisoma viride 97 Ha emulon sexfasciatum 61 Sciaenidae Haemu/on steindachneri 329 Odontoscion dentex 25 Scaridae Scorpaenidae Scarus perrico 62 Scorpaena plumien· 2 Serranidae Serranidae Ep inephelus labriformis 49 Epinephelus cruentatus 1 Epinephelus panamensis 25 Epinephelus fu lvus 18 Sparidae Sphyraenidae Ca lamus brachysomus Shyraena barracuda 8 Teatraodontidae Arothron meleagris 4 Sphoeroides annulatus 2 TABLE 2

List of fa milies and species and total number of individual sigh tings at Ma nuel Antonio National TABLE 3 Park (Pa cific), Costa Rica Percent of each size group at each transect Acanthuridae Acanthurus xanthopterus 22 Cahuita Balistidae Pseudobalistes naufragium 8 Transect 1 2 3 4 5 Blennüdae Ophioblennius 38 1 100 O Plagiotremus azaleus 48 2 54 33 13 Carangidae 3 76 19 2 Ca ranx hippos 18 4 67 12 9 8 3 Ca ranx stellatus 1 Chaetodontidae 5 61 24 13 2 Chaetodon humeralis 19 6 45 45 8 2 Johnrandallia nigrorostris 8 7 38 38 21 7 2 Chromidae Chromis atrilobata 12 Manuel António Cirrhitidae Cirrhitus rivulatus 37 99 Grammistidae 2 89 4 6 Rypticus bicolor Holocentridae 3 59 20 20 1 Adioryx suborbitalis 59 4 19 36 39 6 My ripristis leiognathos 47 5 46 42 11 1 Kyphosidae Kyphosus sp. 101 At Manuel Antonio Nalional Park, a some­ Labridae what lesser number of individual, were sighted Bodianus diplotaenia 37 Halichoeres dispilus 431 (6427), undoubtedly due to fewer transects_ Halichoeres nicho/si 40 Thirty-nine species were recorded and di,tribu­ Pseudojulis notospilus 816 ted among 21 famille, (Table 2)_ The domi­ Thalassoma lucasanum 480 nant families were the wrasses, Labridae Lutjanidae Lutjanus argentiventris 56 (5 ,pp.) and again the promacentrid, (6 ,pp.). Monocanthidae The two most abundant 'pecies were nearly Suffla men verres 13 identical in total number of sightings as !hose 100 REVISTA DE BIOLOGIA TROPICAL of Cahuita; Ihe sergeant major, Abudefd uf saxatilis (1159) and the damselfish, SIegas tes TABLE 4 sp. (1 160), both of Ihe family Pomaeentridae . Meal/ firh abundance (N/8) and index ca/cu/a/ions each Table 3 shows Ihe pereent of individual for trol/sect sightings wilhin each of Ihe previously detailed Cahuita size categories. The transects are ordered from anse t Mean S.D. SR Ihe least complex habitat (1) on, in increasing Tr c H' J' O eomplexity. In both Cahuita and Manuel 6.1 7.60 1.424 6 0.79 71.4 1 , 2 59.3 25.30 2.490 21 0.82 32.9 Antonio, the least complex habitat was exclu­ t 142.5 63.94 2.358 25 0.73 34.4 t sively inhabited by small individuals (to 7.5 4 159.5 76.15 2.367 27 0.72 43.7 t cm), mainly wrasses (Labridae) and damselfish 5 105.7 44.37 2.591 25 0.81 35.9 1: (pomacentridae). In all olher transect areas, 6 361.6 .t:l41.55 2.535 J4 0.72 34.8 113.3 39.54 2.713 30 0.80 29.7 there was a similar distribution of fishes 1: Ihroughout the different size categories, but Manuel Antonio with the greatest concentration usually in the 1 157.3 92.05 2.110 18 0.73 44.2 t smallest size category l. 2 159.6 90.05 2.145 19 0.73 36.9 1: 213.1 92.34 2.487 30 0.73 38.9 On the basis of mean number of individuals t 4 119.6 t 68.18 2.140 27 0.65 49.6 and species at Cahuita (Table 4), Transeet 5 179.5 ± 85.75 2.396 29 0.71 41.1 6, whieh has the seeond most eomplex habitat, eontained the most speeies (34) and mean Shannon-Weiner diversity index (bascd on N and SR). H' = number of individuals (361.6). The species SR numbcr spccies observcd on transcet. = of evenness componen! divcrsity (bascd on and SR). diversity and evenness fo! Transect 6 was l' = of H' perccnl dominance of the two most common spl'Cics. lower Ihan expected for Ihe complexity of Ihis O = area's habitat. But the two most dominant utable to the overall deterioration of Ihe reef as species, Ha emulon macrostomum and S. planifro ns, constituted 35% of all Ihe fishes cited in the literature_ seen on this transect, therefore resulting in At Manuel Antonio, Transeet 3 (Table 4) lower species diversity and evenness values. had the greatest mean number of individuals Transeet 1 had the smallest number of species (2 13.1), number of species (30) and species (6) and individuals (N= 6.1). This may be diversity (2.487), while Transect 1 had the directly related to the low habitat complexity lowest species diversity due undoubtedly, in whieh provided Httle eover for fish . The high part, to the low mean number of species dominance value (71 .4) indica tes Ihe suceess of (18), The two most dominant species on two dominant species (S. planifrons and Scarus Transect 4, Stegastes sp. and Microspathodon croicensis) in exploiting this area. Transects 5 dorsalis, contributed to the highest D value (50%). and 7 had similar mean numbers of fishes and Transects 3 and 4 are both loeated in Puerto species which eould be eorrelated with Iheir Escondido and around the same island exeept similar species diversity and evenness values. Ihat Transect 3 is located on Ihe calm proteeted 80th Transects S and 7 have numerous coral side of the island and Transect 4 is locáted on species, are surrounded by deeper water, and Ihe rough unprotected side. Therefore, Ihe are not located on a reef crest. An overall trend calmer water apparently allows for Ihe devel­ in increasing or decreasing species diversity , opment of a richer fish eommunity. The me1n species richness. mean rrumber of individuals species diversity of 2.627 calculated for the or evenness cannot be seen as the transects transects in Manuel Antonio is comparable with increase in habitat complexity. The mean olher Central American Pacifie fish eommunity speeies diversity value of 2.861 calculated for studies. In El Salvador, Phillips (i 981) reported Cahuita transeets can be eompared wilh Ihose Ihat H'diversity ranged from 0.60 to 2.46 in a computed for fish on a coral patch reef in Ihe coastal mangrove embayment. In an estuarine Virgin Islands where H' was 3.3 (Smilh and environment in Costa Rica, León (1973) found Tyler, 1972) and 3.4 n Ihe entire Caribbean H' values from 2.295 to 2.771 in Ihe Gulf of (Baekus et al., 1970). For Florida reef fish Nicoya, while at a later date, Bartels et al. assemblages H' was 4.290 to 4.529 (Jones and (1983) gave values from 0.671 to 3.163 for Ihe Thompson, 1978). The lower species diversity same estuary. Also, in a Costa Rican Pacific value calculated for Cahuita is probably attrib- eoast tidepool study, Weaver (1970) found a PHIL LIPS & PEREZ-CR UE T: Reef fishes of Caribbean and Pacific Costa Rica 101

fairly low species diversity (mean H' = 1.1-1.4 través de cinco transectos en el Pacífico y siete for Ihree sites)_ en el Caribe. The dendrograms of Ihe overlap in ich­ Ca}lUita, con 49 especies, mostró una diver­ thyofaunal composition for the seven transects sidad media de especies más alta mientras in Cahuita National Park (Figure 3) visually que en Manuel Antonio, con un menor número demonstrate Ihat Transects 2, 3, 4 and 5 have de especies (39), registró un mayor núme­ similar fish compositions due to their tighter ro de individuos. Los transectos con una clustering_ Transects 6 and 7 may represent complejidad de habitat semejante mostraron a loose cluster and l is Ihe least similar to aH valores más altos de PS (aproximadamente 60% ) others_ Figure 4 illustrates the same for Ihe Los lábridos Y los promacéntridos dominaron five transects in Manuel Antonio. Again from en ambas áreas de arrecife rocoso del Pacífico y the dendrogram one can see that transects de coralino del Caribe. Generalmente domina­ that have similar habitats have similar species ron en los transectos los individuos hasta de 7.5 compositions_ Transects 3, 4 and 5 which have cm. Conforme aumentó la complejidad de un · more complex habitats are loosely grouped transecto aumentó el número de peces grandes. while Transects I and 2 which do not have such La diversidad del arrecife de Cahuita fue baja en complex habitats are associated at a higher level relación con otros arrecifes del Caribe debido of similarity. probablemente al estado deteriorado en que se In summary, the traosects io the cora! reef at encuentra éste. Los valores de diversidad Cahuila had higher species diversity and de especies en Manuel Antonio están dentro del evenness values Ihan those calculated for the ámbito de valores encontrado en estudios reali­ rock reefs in Manuel Antonio. However, more zados en estuarios y en charcos de la zona entre fish were seeo io the transects of Manuel mareas del litoral Pacífico de Centroamérica. Antonio. The greater topographical variability of Ihe reef in Cahuila could be Ihe reason why Cahuita had a higher species diversity value. LITERATURE CITED The percent similarity values demonstrated Ihat similar habitats tend to have similar species Alevison, W.S., & M. G. Brooks. 1975. The compar­ compositions. The most outstanding conclusion ative structure of two Western Atlantic reef-fish assemblages. Bull. Mar. ScL, of this fish census is Ihe relatively low species 25 : 482490. diversity encountered at Cahuita which sup­ Backus, R.H., LE. Craddock, R.L. Haedrick, & D. E. ports evidence indicating the declining state of Shores. 1970. The distribution of mesopelagic Cahuita's coral reeL fishes in the western tropical North Atlantic. Ma:r. Re&, 28, 179-201.

ACKNOWLEDGEMENTS Bartels, C.E., K.S. Price, M.I. López, & W. A. Bussing 1983. Occurrence, distribution, abundance and We thank the Associated Colleges of Ihe diversity of fishes in the Gulf of Nicoya, Costa Midwest (ACM), San José, Costa Rica for Rica. Rev. Biol. Trop., 31: 75-101. Ihe logistic support of this study. The Servicio Brock, R.E. 1982. A critique of the visual census de Parques Nacionales, Costa Rica, extended method for assessing coral reef fish populations. permission to carry out the fish census at Bull. Mar. Sci., 32, 269-276. Cahuita and Manuel Antonio National Parks. Special Ihanks go to William Bussing at Ihe Brock, V.E. 1954. A preliminary report on a method of estimating reef fjsh populations. Wildlife Mgmt., University of Costa Rica for invaluable help in 18, 297-308. the identification and confirrnatioo of many fish species. Chapman, C.J., A.D.F. Johnstone, J.R. Dune, & D. J. Creasy. 1974. Reaction to fish of sound generated by diver's open circuit underwater apparatus. RESUMEN Mar. Biol., 27, 357-366.

De febrero a mayo de 1982 se hizo un censo Gladfelter, W.B., J.C. Odgen, & E.H. Gladfelter. 1980. de los peces por medio de una técnic.a rápida Similarity and diversity among coral reef nsh visual en el Parque Nacional Manuel Antonio communities: a comparison between tropical western Atlantic (Virgin Islands) and tropical (Pacífico) y en el Parque Nacional de Cahuita central Pacific (Marshall lslands) patch reefs. (Caribe), Costa Rica. Los peces se obtuvieron a Ecology, 61. 1156-1168.

r, as , {m 60¡ 011"s,/c,/ Htf¿,jt.J I I 102 REVTSTA DE BIOLOGIA TROPICAL

Haedrich, R .L. 1975. Diversity am! averlap as meas­ Sale, P.r:. 1980. Assemblages of fish on patch reefs: ures 01" envÍIonmcntal quality, Water Res., 9: predictable or unpredictable. Environ. Biol. Fish., 945-952. 5, 243-249.

Hobson, ES 1974. ¡"ceding relationships of te1eostean Smith, e.L. 1973. Small rotcnone stations: a too1 for fishcs on coral rccfs in Kona. Fish. 8ull. U.S., 72: studying coral fish communities. Amer. Mus. 915-1031. Novit., 2521: 1-21;

Jones, R.S., & J.A. Cbase. 1975. Community structurc Smith, c.L., & J.e. Tylcr. 1972. Spacc resources and distribution of fishcs in an cncloscd island sharing in a coral red fish community, p. 125-170, lagoon in Guam. Micronesia, 11: 127-148. In B.B. Collctte & S.A. Earlc (eds.). Results of tite Tcktite Program: Ecology 01' coral rccf fishes. l3ul1. Jones, R,S., & M.J. Thompson. 1978. Comparison of Nat. Bist. Mus. Los Angeles. No. 14. Florida recr fish asscmblages using a rapid visual technique. Bul!. Mar. SeL, 28: 269·281. Sokal, R.R., & C.D. Michencr. 1958. A statistical method for evaluating systematie relationships. León, P.E. 1973. Ecología de la ictiol'auna del Golfo Kansas Univ. Sci. Bull., 38:1409-1439. de Nicoya, Costa Rica, un estuario tropical. Rev. Biol. Trop., 21: 5-30. Talbott, F.B., & B. Coldman. 1973. A prctiminary report on thc diversity and fceding rclationships Lloyd, M., 1. H. Zar, & LR. Kan. 1968. On the calcu­ of recf fisiles of Onc Tree Island, Creat Barricr latian of information-theorctical mensures of Recf System. p. 425-442, In Symposium on corals diversity. Amer. MidJ. NaL, 79: 257-272. and coral reefs. Mandoporan Camp, India. (12-16 January 1969). Marine Biological Association 01' Phillips, P.e. 1981. Diversity and fish community India. structure in a Central American mangrove cm­ baymcnt. Rev. Biol. Trop., 29: 227-236. Weaver, P.L. 1970. Specics diversity and ccology of tidepool fishes in thrce Pacific coastal areas of Pielou, E.C. 1966. Thc mcasurcmcnt of divcrsity in Costa RiCa. Rev. Biol. Trop., 17: 165-185. different types of biological collcctions. J. Theor. RioJ.,U, 131-144. Wellington, G.M. 1974. An ecological dcscription of the marine and associatcd cnvironments at Monu­ Risk. M.J., M.M. Murillo, & J. Cortés. 1980; Observa­ mento Nacional Cahuita. San José, Costa Rica. ciones biológicas preliminares sobre el arrecife Subdirección de Parques Nacionales. 81 p. (meca­ coralino en cl Parque Nacional de C .. huita, Costa nografiado). Rica. Rev. Biol. Trop., 28: 361-382.