articulos octocorales white papers cientificos investigaciones fauna arrecifal

BULLETIN OF MARINE SCIENCEVOLUME 23 1973 NUMBER 3

THE ECOLOGY OF THE CORAL-OCTOCORAL COM-MUNITIES OFF THE SOUTHEAST FLORIDA COAST

GEOMORPHOLOGY SPECIES COMPOSITIONAND ZONATION1

WALTER M GOLDBERG2University of Miami Rosenstiel School of Marine

and Atmospheric Science

ABSTRACT

Three parallel submarine terraces found along the southeast coast ofFlorida stretching from Miami through Palm Beach County are de-scribed The central portion of this area near southern Palm BeachCounty was analyzed with respect to geomorphology community com-position and zonation from the low-tide mark to a depth of 50 m Twenty-seven species of scleractinian corals and 39 species of gorgonians are foundhere and define a typical coral-reef community farther north than hasbeen acknowledged Gorgonian diversity is maximal at a depth of 15-20m while scleractinians are most diverse in shallower water Studies ofgorgonian biomass indicate a trend toward large numbers of small indi-viduals in low-diversity environments and a smaller number of largerindividuals in higher-diversity environments A mean density of 251coloniesjm2 gives these reefs the highest concentration of gorgoniansyet recorded in the Caribbean region

INTRODUCTION

A northward extension of the Florida Reef Tract was first proposedover 80 years ago when Shaler (1890) stated that corals were presentalong the Florida coast 40 miles north of Miami Vaughan (1910) dis-puted Shalers data and concluded that coral communities were absentnorth of the Florida Keys partly because of lower temperatures andpartly due to the southward transport of siliceous sand More recentlyseismic profiles have revealed a series of reeflike ridges or terraces parallelto the south Florida coastline (Duane amp Meisburger 1969) The exact

1Contribution No 1671 from the University of Miami Rosenstiel School of Marine and Atmo-spheric Sciencebull Present address Department of Biological Sciences Florida International University MiamiFlorida 33144

466 Bulletin of Marine Science

()

bull ~uPiter Inlet

bull PALM bull ~ BEACH COUNTY Boca RotoL bullbull

[23(3)

Dry [qrtugos

(~7 (U 0_1~~~~~I West I

820 81deg

I inch = 38miles

= reefs

I80deg

FIGURE 1 The area of investigation with localities mentioned in the text

nature of these structures is not known They may be coral reefs or theymay consist of a Recent coral veneer overlying Pleistocene beachrockFor the purposes of this paper the term reeflike ridges will be abbre-viated and the term reef along with patch reef reef tract etcwill be used where physiographically applicable without regard to thenature or depth of the antecedent platform This report intends to de-scribe the biological community presently living upon and presumablycontributing to these geological structures

1973] Goldberg Ecology of Coral-Octocoral Communities 467

ForeReefZone

FirstReef

PatchReef Sand Depression

Zone OuterReef

Platform

Outer ReefSlope

a 600 800 1200METERS

FIGURE2 General zonation and morphology of the reefs off the southeastFlorida coast

Three primary divisions of the Florida Reef Tract are recognized (Gins-burg 1956) (1) the back-reef zone which includes the relatively un-developed inshore and patch reefs (2) the outer-reef zone which includesthe well-developed reef platform and its slope (outer reef or seawardslope) into deeper water and (3) the fore-reef zone which includes thedeeper areas adjacent to the reef habitat proper Similar divisions can bedistinguished on the northern reef tract which extends from Fowey Rocksin the south to the Jupiter area of northern Palm Beach County Florida(Fig 1)

These reefs are divided into three terraces (Fig 2) The first is closestto shore and forms a fairly well-developed back reef some 100 metersoffshore in 4-5 meters of water in the Fort Lauderdale-Pompano Beacharea Off Boca Raton however the first reef is primarily littoral and iscomposed of ca1carenitic beachrock rather than coral limestone (Craiget al 1969)

A plain of gradually sloping sand lies between the first and second reefterraces The latter develops relief of 2-3 meters and is composed of abroad platform of gorgonians and flat coral colonies It lies in 7-8 metersdepth and is approximately 800 meters offshore in the Fort Lauderdale-Pompano Beach area A rocky ledge one or two meters high marks theseaward termination of this reef The second reef off Boca Raton liesin 9-10 meters depth and is composed mainly of a line of flat patch reefs

468 Bulletin of Marine Science [23(3)

of variable size In all patches observed this far north the substrate risesscarcely more than 03 to 06 meters above the sand flats In this paperthe term back reef will refer to both the first and second reef terraces

The second terrace is separated from the third by a sand depressionwhich slopes to a depth of 17-19 meters At this point it is abruptlyterminated in most areas by a sheer rocky ledge 3-4 meters high (Fig 2)The top of the ledge marks the crest of the third reef which generally liesin 16-18 meters of water The reef which begins here is biologically andgeologically the most well developed of all the terraces and extends to adepth of 30-40 meters The outer reef can be divided into two subareasThe first is termed the outer reef platform because of its low relief andgradual slope The platform extends to a depth of about 20 meters At thispoint the terrain becomes rugged forming knolls In many localities thesefuse into spurs some of which attain heights of over 8 meters Theseareas of relief are separated from one another by sand or rubble-filledgrooves which are oriented in an east-west direction This region slopesmore rapidly into deeper water and is termed the outer reef slope

The outer reef terminates in some areas by forming progressivelysmaller knolls or spurs while in other regions the framework has adefinite seaward limit In either case at a depth of 30-40 meters theterrain is composed of a flat plain of rubble This marks the fore-reefregion which extends to a depth of at least 50 meters

MATERIALS AND METHODS

The principal area of investigation was off Boca Raton Florida Thefirst reef here is close to shore and faunally impoverished Ecological in-vestigations were limited accordingly Two patch reefs were investigatedby collecting in a square~meter path along two transects which coveredmaximum length and width respectively The outer reef was studied byestablishing a linear series of successively deeper stations around whichcollections and square-meter-quadrat samples were taken These stationswere made at depths of 16 22 26 and 30 meters Collections were alsomade on the fore reef at 32 and 44 meters The number of samples wasdetermined by the number required to reveal no new species at a givenstation Ten per station was sufficient for this purpose Collected speci-mens were allowed to dry several weeks at room temperature and wereweighed to the nearest half gram Coral biomass was not measuredSpecies diversity was measured for both corals and gorgonians with theShannon-Weaver information function H = -~ Pi In Pi where Pi is theproportion of the individuals in the ith species (Pielou 1966) Theevenness components (1) which describe the ratio of observed to maxi-mum diversity were calculated as well (Pielou ibid)

4691973] Goldberg Ecology of Coral-Octocoral Communities

RESULTS

I THE SURVEYS OF THE FIRST AND SECOND REEFS

The first reef in the study area is primarily beachrock and extends fromthe littoral zone to a depth of about 15 m below low tide This area iswell scoured by wave action and suspended sediments and is perenniallyturbid Most of the animals found just below low tide live in crevicesor hollows The crab Plagusia depressa is abundant as are sea urchinssuch as Diadema antillarum Echinometra lucunter Lytechinus variegatusand Eucidaris tribuloides In some areas the honeycomb worm Phrag-matopoma lapidosa forms broad flat colonies Large expanses of theboring sponge Ciona celata are also common Below this area broadplains of algae composed of species of Padina and Dictyota are found Theonly scleractinian here is the hardy Siderastrea radians Gorgonians arevery uncommon on the first reef Occasionally young colonies of Pseu-dopterogorgia americana P acerosa and Muricea muricata are foundbut they rarely reach mean size due to the pounding of winter stormsWhere the rock reef projects above low tide various intertidal formscongregate including large numbers of the false limpet Siphonaria pecti-nata and the chiton Acanthopleura granulata

The second reef terrace is about 570 meters directly offshore in 9 metersof water These flat patch reefs vary in size the largest being about 43 by32 meters The size of these reefs however may change considerablydue to shifting sands during storms (Goldberg unpublished)

A total of 15 species of coral was collected from the patch reefs (Table1) and the relative abundance of the species was noted Most of thesewere small and encrusting or lobate ranging in size from 3 to 15 cm indiameter Ocuina diffusa Solenastrea hyades Dichocoenia stokesii andMontastrea cavernosa were the most common scleractinians of this typeOn most of the back-reef patches a few large colonies of Diploria clivosaand Montastrea cavernosa were found D clivosa formed broad encrustingsheets which occasionally grew to over a meter in diameter while Mcavernosa formed low cone-shaped colonies the largest of which was 38cm in diameter However colonies of this size were uncommon

The most abundant animals were the gorgonians particularly those ofthe family Plexauridae (Table 2) Most of the plexaurids were of smallto medium size (20 cm or less) and were thus apparently either youngor stunted colonies A few specimens of Muricea and Plexaurella howeverreached heights of over 40 cm The family Gorgoniidae was the secondmost abundant family of gorgonians the most common member of whichwas Pseudopterogorgia americana The abundance was accentuated by thefact that P americana was larger on the average than any other gorgonianThe range in height was most commonly 35-40 em P acerosa a common


TABLE 1DISTRIBUTION AND ABUNDANCE OF THE REEF SCLERACTINIA

[23(3)

Location Abundance () t

Suborder AstrocoeniinaFamily Astrocoeniidae

Stephanocoenia michelinii M-E amp H PR 560ORP 125ORS 250

Family PocilloporidaeMadracis decactis (Lyman) ORP 050

ORS 250Family Acroporidae

Acropora cervicornis (Lamarck) ORP 050Suborder Fungiina

Family AgariciidaeAgaricia agaricites f purpurea (Linn) PR 100

ORP 560ORS 250

Agaricia lamarcki M-E amp H ORS 3000FR 1000

Family SiderastreidaeSiderastrea siderea (E amp S) PR 560

ORP 125ORS 466

Family PoritidaePorites astreoides Lesueur PR 560

ORP 560Suborder Faviina

Family FaviidaeSolenastrea hyades (Dana) PR 1500

ORP 125Diploria clivosa (E amp S) PR 560

ORP 2000Diploria labyrinthiformis (Linn) ORP 050Manicina areolata (Linn) PR 560

ORP 125Manicina mayori Wells ORP 050

ORS 466Cladocora arbuscula (Lesueur) PR 250Colpophyllia natans (MUlier) ORP 125Montastrea annularis (E amp S) PR 250

ORP 2000ORS 466FR 1400

1973] Goldberg Ecology of Coral-Octocorai Communities

TABLE 1 (Continued)

471


PR 1500ORP 2000ORS 3000FR 1400

PR 250ORP 125

PR 1500

PR 1500ORP 560ORS 466FR 200ORS 100PR 100ORP 560ORS 466FR 6000

PR 250ORP 560ORS 466ORP 125ORS 250ORS 100

ORP 125

Dichocoenia sellaris M-E amp HMeandrina meandries f meandrites (Linn)

Monasrea cavernosa (Linn)

Mussa angulosa (Pallas)

Meandrina meandrites f danae (Linn)

Family MussidaeMycetophyllia lamarckiana M-E amp H

Family RhizangiidaePhyllangia americana (M-E amp H)

Family OculinidaeOculina difJusa Lamarck

Family MeandrinidaeDichocoenia stokesi M-E amp H

Scolymia lacera (Pallas)

Suborder CaryophylliinaFamily Caryophylliidae

Eusmilia fastigiata (Pallas)

bull PR = Patch Reefs 9 meters deep Total species = 15ORP = Outer Reef Platform 16-20meters Total species = 20ORS = Outer Reef Slope 20-30 meters Total species = 14FR = Fore reef 30t meters Total species = 5

t Relative abundance lS calculated as the percentage contributed by each species to each reef zone

associate of P americana was not as common However an investigationof other patches only a few meters away revealed a great difference inthe population of Pseudopterogorgia On some of the smaller patcheshaving relief specimens of P acerosa one meter high or more occurredwhile specimens of P americana were relatively small and uncommonSmall colonies (IOta 11 em high) of the sea fan Gorgonia ventalinawere also fairly common but large specimens were rare

[23(3)472 Bulletin of Marine Science

II RESULTS OF THE OUTER REEF SURVEY

1 The Ledge Habitat- The vertical face of the ledge which began thethird reef was extensively bored particularly by organisms such as thedate mussel Lithophaga nigra and species of boring sponges in the genusCliona Encrusting organisms were also abundant Among those com-monly found were small encrusting colonies of the calcareous bryozoanTrematooecia turrita and encrustations of the foraminiferan Gypsina vesi-cularis The sponge Microciona juniperina was found commonly as an en-crusting form Most of the animals of the ledge were small andor cryptichowever two large organisms were common the comatulid crinoidNemaster rubiginosa and the anthothelid gorgonian 1ciligorgia schrammi

2 The Outer Reef Platform-The organisms characteristic of the plat-form include algae such as the red calcareous Galaxaura obtusata andthe calcareous green forms such as Halimeda opuntia H discoidea andUdotea flabellum Dictyota bartayresii however is the most commonalga The barrel sponge Xestospongia muta is very common on the reefplatform and upper-reef slope (Fig 5) This sponge can be as large as oneby two meters and serves as a home for the usual variety of spongeinquilines among which specimens of the ophiuroid genus Ophiactis arethe most prominent In some areas the zoanthid Palythoa sp formsbroad encrusting sheets particularly on the small sloping mounds ofrocky substrate

Coral species reach their maximum number at the reef crest (Table 1)Some such as Acropora cervicornis are found only at this location Ingeneral however the species of coral found are otherwise not appreciablydifferent from those found on the back reefs Notable exceptions to thisinclude the greater abundance of Meandrina meandrites forma meandritesand Agaricia agaricites on the outer reef Oculina difJusa so common onthe sandy back reefs is conspicuously absent from the outer reef Themajor difference between corals found on the platform and the coralsfound on the back reefs is size Larger colonies of the same species arefound in deeper water than in shallow but colonies reaching a width ofover 30 em are not common Largest species (greater than 30 em indiameter) are as on the patch reefs Diploria clivosa Montastrea cavernosaand M annularis

The most conspicuous organisms on the platform are the gorgoniansIn contrast to the coral populations there is a distinct difference betweengorgonians occurring on the back reef and those occurring on the outerreef Not only are some of the species different (Table 2) but individualsizes and population densities are distinctly dissimilar The specimens onthe platform were half the number but twice the mass per square meter ofreef when compared to those on the patch reefs This is primarily due to


TABLE 2DISTRIBUTIONANDAn UNDANCEOF THE REEF GORGONACEA

Location Abundance ()i

Suborder ScleraxoniaFamily Briareidae

Briarellin asbestinum (Pallas) PR 166ORP 333SS 560DS 125

Family AnthothelidaeErythropodium caribaeorum (D amp M) PR 040lciligorgia schrammi Duchassaing ORP 560

SS 1500DS 2000FR 500

Diodogorgia noduliera (Hargitt) FR 500

Suborder HolaxoniaFamily Plexauridae

Plexallra flexuosa Lamouroux PR 1500ORP 750SS 560DS 125

Plexallrella dichotoma (Esper) PR 166ORP 560SS 200

Plexaurella grisea Kunze PR 166ORP 333

Plexaurella fusifera Kunze ORP 750SS 1500

Plexaurella pumila Verrill DS 067Muricea muricata (Pallas) PR 1500

ORP 750Muricea pendula Verrill FR 033

Muricea laxa Verrill FR 033

Muriceopsis petila Bayer DS 2000Eunicea calyculata (E amp S) PR 1500

ORP 560SS 1500FR 033

Eunicea laciniata (D amp M) PR 166S8 200

Eunicea laxispica (Lamarck) PR 040

474 Bulletin at Marine Science

TABLE 2 (Continued)

[23(3)

Location~ Abundance ()i

Eunicea palmeri Bayer

Eunicea fusca (D amp M)

Eunicea succinea (Pallas)Eunicea tourneforti (M-E amp H)

Eunicea clavigera Bayer

Eunicea pinta Bayer amp Deichmann

Pseudoplexaura spPseudoplexaura crucis Bayer

Family GorgoniidaePseudopterogorgia rigida (Bielschowsky)

Pseudopterogorgia acerosa (Pallas)

Pseudopterogorgia americana (Gmelin)

Pseudopterogorgia elisabethae Bayer

Pseudopterogorgia navia BayerPseudopterogorgia spl

Pseudopterogorgia sp2

Pterogorgia citrina (Esper)

Pterogorgia guadalupensis (D amp M)

Gorgonia ventalina Linn

Lophogorgia cardinalis Bayer

PRORPSSDSPRORPDSPRORPSSDSORPSSSSDSDSFR

PRORPDSPRORP

PRORPSSSSFRDS

PRS5ORP

PR

PRSSPRORPS5FR

700560560

2000166560125700750

1500125

750560200

1400125033

700750125700333

1500750200040033067

040040

067040

040040

166750040033


TABLE 2 (Continued)

475

Location Abundance ()t

Family EllisellidaeEllisella barbadensis (D amp M)

Ellisella sp

Nicella schmitti Bayer

Family ParamuriceidaeSwijtia exserta (E amp S)

ORP 06755 560DS 1400FR 140055 040D5 067FR 1400ORP 0675S 200D5 125FR 3000

DS 125FR 3000

bull PR = Patch Reefs 9 meters deep Total pecies = 19ORP = Outer Reef Platform 16-20 meters Total species = 19SS = Shallow slope 20-25 meters Total species = 19DS = Deep slope 25-30 meters Total species 16FR Fore reef 30+ meters Total species = 12

t Relative abundance is calculated as the percentage contributed by each species to each reef zone

a few abundant large species Plexaurella fusifera and Eunicea taumefortifor example were sometimes 80 em or more in height and such coloniescould weigh as much as 300 grams (dry weight) A comparison of popu-lations between the platform and the patch reefs (Table 2) shows thatalthough the number of species is similar the species composition is notlciligorgia sehrammi Plexaurella fusifera Eunieea toumeforti and Eunieeac1avigera were reasonably common on the platform but were absent fromany of the back reefs

3 The Outer Reef Slape-The outer reef slope comprises the bulk ofthe third reef terrace This subdivision extends from a depth of about 20meters to the end of the reef at 30 meters The transition between outerreef platform and outer reef slope occurs gradually within a depth ofabout 4 meters and a horizontal distance of approximately 50 metersThe low sandy slopes of the platform gradually give way to the rockyknolls which characterize this region These structures support a profusefaunal assemblage quite different in composition from that of the plat-form (Fig 5) Many of the algae which are prominent in the previouszone are not as common on the slope although masses of Dietyota bartay-resii and Galaxaura obtusata are still present Populations of the tlagelli-form antipatharian Cirripathes luetkeni become quite abundant below 22 m

47630

Bulletin of Marine Science

951

[23(3)

~o

10

middot5

---- --- -134 --_

egt---ltgt GORGONIANS----- CORALS

-bull

115

PATCH REEF OUTER SHALLOW DEEP FORE-REEFAREA REEF SLOPE SLOPE

PLATFORM

FIGURE 3 Species diversity (R) vs vertical zonation on the reef tractNumbers associated with graph points indicate the evenness (J)

Coral diversity decreases on the outer reef slope (Fig 3 Table 1) andthe population becomes largely dominated by Montastrea cavernosa (FigS) In many areas one of the leaf corals Agaricia lamarcki codomi-nates and is always found as an encrusting form morphologically similarto A agaricites forma purpurea (Fig 6) The outer reef slope is notreadily divisible into subzones on the basis of its coral fauna

The members of the order Gorgonacea were most conspicuous membersof the slope assemblage lciligorgia schrammi was encountered occasionallyon the platform where it reached moderate lengths but on the slope it wasoften 1-15 meters high and was one of the most common and mostprominent gorgonians For this reason the outer reef slope is also referredto as the lciligorgia zone The abundance and strikingly uniform orienta-tion (normal to current) characteristic of this species is shown in Figure7 (d Wainwright amp Dillon 1969)

Although many large forms were found much of the gorgon ian popu-lation was composed of the smaller species This was particularly true onthe deepest parts of the slope where many of the large species of Eunicea


40 bull bull bull Deep slopebull 0 o Potch reefsbull bull bullbull 0N 0 bullOuter reef platform~ bull 0 0

- bull bull Upper part of outeren 0 00 reef slope-I bullltl 0J 30 0agtazu0

Ciz 20

10 400 500 600100 200 300

BIOMASS (g)

FIGURE 4 Gorgonian biomass vs vertical zonation Each point representsa square meter sample Average number of coloniesjm2 is 337 for the deepslope 343 for the patch reefs 152 for the outer reef platform and 174 forthe outer reef slope The areas of higher diversity tend to have greater biomassdistributed among fewer individuals

and Plexaurella were absent At a depth of 22-25 m some species ofgorgonians previously common were absent or becoming increasingly rareThe total number of species however does not change due to gradualreplacement by deep-reef species The loss of shallow-water taxa is mostreadily demonstrated on the outer reef slope below 25 m Here 50per cent of the species are confined to depths below 20 m InterestinglyEunicea palmeri the most abundant gorgonian at 30 m is abundant in1- to 2-m depths in the Florida Keys (Bayer 1961 131) No investiga-tions have been made of deep-water gorgonians in the Keys however

At the reefs edge most of the gorgonians present with the exceptionof Iciligorgia schrammi and some Ellisella barbadensis are small This isthe reason for the reduced biomass here (Fig 4) A corresponding dropin number of species is seen upon comparison of the shallow and deepends of the slope (Table 2) Although 16 species were found at 30meters only five could be considered common


FIGURE 5 View of a knoll on the outer reef slope (depth 24 m) Thebarrellike sponges (left foreground and top center) are Xestospongia muta(Schmidt) The flagelliform gorgonian growing from left sponge is Eliselabarbadensis (Duch amp Mich) The large gorgonian in the center is lciligorgiaschrammi (Duch) All of the coral colonies are Montastrea cavernosa (Linn)


FIGURE 6 The encrusting form of Agaricia lamarcki Milne-Edwards ampHaime typical of the deep-reef habitat in Florida (depth 28 m)

4 The Fore-Reef Zone-The fore-reef zone usually begins abruptly at adepth of 30 meters where the outer-reef knolls or grooves do not persistThe terrain is similar to the rubble of the grooves which run between thespurs of the outer reef slope The rubble appears to be talus but isactually a series of nodular encrustations The nucleus can be anythingfrom bryozoan to coral This is subsequently overgrown by serpulidscoralline algae sponges ascidians and other organisms Usually only theoutermost layer is composed of living material The inner remains arepermeated by loosely cemented to uncemented sand grains The forereef is virtually without relief and descends to the deepest area investigated(50 m)

The more conspicuous benthic invertebrates of this area are found inscattered groupings composed of various sponges didemnid ascidiansgorgonians and small corals The one coral that appears to be character-istic of the fore reef is Meandrina meandrites forma danae

Despite the reduction in numbers of species the Gorgonacea are still themost conspicuous invertebrates of the fore reef Groups of five or sixgorgonians define the islandlike aggregations which stand out prominentlyagainst the barren rubble As is shown in Table 2 12 species are presenton the fore reef but of these six are rare two are occasional and only fourare characteristic of the area Three of the four characteristic species


FIGURE 7 View of the outer reef slope (lciligorgia zone depth 28-30 m)showing orientation of I schrammi Current flow is normally from left toright

are of the family Ellisellidae and most of these were found only occasionallyon the reef proper

DISCUSSION

1 The Gorgonacea- The highest gorgon ian diversity was found on theouter reef platform Species here are fairly evenly distributed with eightabundant five common three uncommon and three rare taxa resultingin the high evenness (J) components shown in Figure 3 The correlationof this area with highest diversity is not unexpected The depth range(16-20 m) is not great enough to eliminate shallow-water species but isgreat enough to thin their ranks so that the population is not dominatedby one or two species as in shallow water It is also significant that deeper-water species are found here which adds to the heterogeneity of the plat-forms gorgonian fauna

1973J Goldberg Ecology of Coral-Octocoral Communities 481

The decline of diversity in deeper water is a function of not only thefewer species present but the increase in the proportion of rare to commonspecies The sorting mechanism is not known at present but it is probablethat the low diversity and extreme patchiness of the fore reef is attributableto the lack of firm substrate in that zone rather than depth per se

Table 3 shows the relative dominance of all reef gorgonians The firsteight species account for over 50 per cent of all the individuals sampledIt is interesting to note that four of these eight dominants are restrictedentirely to below 16 m Their prominence is due to greater abundance in alow-diversity situation The other four are vertically cosmopolitan rcefspecies whose abundance is contributed to by both outer-reef and patch-reef environments

The quadrat samples provide another aspect of vertical distribution inthe Gorgonacea Figure 4 shows the number of individualsjm2 plottedagainst biomass From this it is apparent that the margins of the reefhabitat (ie the deep slope and the second reef) contain a large number ofsmall individuals while the outer reef platform and upper slope contain arelatively small number of very large individuals per m2bull Perhaps thisdistribution is indicative of the unstable conditions on the patch reef onone hand and the increasing diminution of light or lack of suitable substratewith depth on the other The noteworthy feature is the correlation of lowerdiversity with conditions promoting high density and low biomass Theapplicability of this observation to other reef situations however is notknown at present

With rare exceptions symbiotic gorgonians (species with zooxanthellae)are the only forms found above 16 m In fact many species such asPseudopterogorgia acerosa and Gorgonia ventalina appear to be restrictedto this zone Below 20 m various asymbiotic species such as Iciligorgiaschrammi and some of the eIliseIlids become fairly numerous Symbioticforms however are still most abundant although they are generally notconspecific with those encountered in shallower water Species such asEunicea tourneforti E pinta Pseudopterogorgia navia and P Elisabethaeall have zooxanthellae but are found only on the outer reef particularlyon the slope below 20 m (Table 2) Only six species are found both at9 and 30 m These are Briareum asbestinum Plexaura flexuosa Euniceapalmeri E fusca E calyculata and Pseudopterogorgia rigida all of whichare symbiotic

As one might expect the number of asymbiotic individuals and speciesincreases directly with depth Nicella schmitti and Swiftia exserta becomemost common below 30 m and with Ellisella spp dominate the gorgonianpopulation Kinzie (1970) noted a similar phenomenon in Jamaica butsymbiotic domination persisted to a depth of 55 m in that locality


TABLE 3SUMMARYOF THE REEF GORGONACEAANDRANK OF DOMINANCE

Depth range Rank Species (m) Dominance

1 lciligorgia schrammi Duchassaing 16-32 9122 Eunicea calyculata (Ellis amp Solander) 9-32 7183 Ellisella barbadensis (Duch amp Mich) 16-32 7054 Nicella schmitti Bayer 16-32 6655 Eunicea palmeri Bayer 9-30 6426 Swiftia exserta (Ellis amp Solander) 30- 6257 Plexaura fiexuosa Lamouroux 9-30 5878 Pseudopterogorgia americana (Gmelin) 05-22 4909 Eunicea tourneforti Milne-Edwards amp Haime 16-30 475

10 Plexaurella fusifera Kunze 16-22 45011 Muricea muricata (Pallas) 05-16 45012 Muriceopsis petila Bayer 30- 40013 Eunicea pinta Bayer amp Deichmann 22-30 32014 Pseudopterogorgia rigida (Bielschowsky) 9-30 31515 Ellisella sp 22-32 30116 Eunicea clavigera Bayer 16-22 26217 Briareum asbestinum (Pallas) 9-30 23718 Pseudopterogorgia acerosa (Pallas) 05-16 23719 Gorgonia ventalina Linn 9-22 19120 Eunicea fusca Duch amp Mich 9-30 17021 Plexaurella dichotoma (Esper) 9-22 16522 Eunicea succinea (Pallas) 9 14023 Plexaurella grisea Kunze 9-16 10024 Diodogorgia noduliera (Hargitt) 30- 10025 Erythropodium caribaeorum (Duch amp Mich) 9 08026 Eunicea laxispica (Lamarck) 9 08027 Eunicea laciniata (Duch amp Mich) 9-22 07328 Pseudoplexaura sp 30 02529 Pseudopterogorgia spl 9-22 01630 Pterogorgia guadelupensis Duch amp Mich 9-22 01631 Plexaurella pumila Verrill 30 01332 Pseudopterogorgia navia Bayer 30- 01333 Pseudopterogorgia sp2 16 01334 Pseudopterogorgia elisabethae Bayer 22-32 00835 Pterogorgia citrina (Esper) 9 00836 Muricea pendula Verrill 30- 00737 Muricea laxa Verrill 30- 00738 Pseudoplexaura crucis Bayer 30- 00739 Lophogorgia cardinalis Bayer 30 007


TABLE 4SUMMARYOF THE REEF SCLERACTINIAAND RANK OF DOMINANCE


1 Montastrea cavernosa (Linn) 9-32 19752 Meandrina meandrites f danae (Linn) 30- 15003 Montastrea annularis (Ellis amp Solander) 9-32 10294 Agaricia lamarcki Milne-Edwards amp Haime 22-32 10005 Dichocoenia stokesii Milne-Edwards amp Haime 9-32 6816 Diploria clivosa (Ellis amp Solander) 9-16 6407 Soenastrea hyades (Dana) 9-16 4068 Oculina difJusa Lamarck 9 3879 Mycetophyllia lamarckiana

Milne-Edwards amp Haime 9-32 31910 Meandrina meandrites f meandrites (Linn) 9-32 28111 Porites astreoides Lesueur 9-16 28012 Siderastrea siderea (Ellis amp Solander) 05-32 28013 Stephanocoenia michelinii

Milne-Edwards amp Haime 9-32 23314 Agaricia agaricites f purpurea (Linn) 9-30 22715 Manicina areolata (Linn) 9-32 17116 Manicina mayori Wells 16-30 12817 Phyllangia americana Milne-Edwards amp Haime 9-16 09418 Mussa angulosa (Pallas) 16-30 09419 Madracis decactis (Lyman) 16-30 07520 Cladocora arbuscula (Lesueur) 9 06221 Eusmilia fastigiata (Pallas) 16 03122 Colpophyllia natans (MUlier) 16 03023 Dichocoenia stellaris Milne-Edwards amp Haime 24 02524 Scolymia lacera (Pallas) 18 02525 Diploria labyrinthiformis (Linn) 16 01226 Acropora cervicornis (Lamarck) 16 01227 Siderastrea radians (Pallas) 05-2 003

At least 39 species of gorgonians are found on the reefs studied hereBayer (1961) has listed the number of gorgonian species known fromother shallow-water Caribbean areas For comparison some of his figuresare as follows east coast of Florida to Dry Tortugas 47 species BahamaIslands 32 species Cuba 25 species Jamaica 22 species Bermuda 18species and Puerto Rico 16 species

The low figures cited for these areas are probably due to scattered col-lections rather than faunal impoverishment A recent list by Gonzales-Brito(1970) for example has brought the number of shallow-water PuertoRican gorgonians to 35 48 per cent of which are also found in this study


Similarly Kinzie (1970) has shown after intensive collecting that atleast 43 species of gorgonians are present on reefs off northern JamaicaSixty-two per cent of these Jamaican species are also found off southernPalm Beach County The total cumulative-area diversities for gorgoniansin these two areas are similar mathematically but a comparison of habitatsis not quite as simple First the reefs differ topographically In Jamaicathe reefs begin just offshore grade through a number of subzones and dropinto deep water (Gareau 1959 1967) Many of these subzones such ashigh-energy shallow-water reefs shallow Acropora rubble areas andprecipitous slopes effectively lower gorgonian diversity (Kinzie 1970)Such environments are absent from the reefs being considered here SecondKinzie noted that gorgonian diversity is inversely related to the areacovered by living coral (A similar situation between Indo-Pacific coralsand a1cyonaceans was alluded to by Wells [1957]) In an area with ex-tensive coral development such as Jamaica it is not surprising that thedensity component of gorgon ian diversity is somewhat decreased ThusKinzie found an average of 19 individualsjm2 while Cary (1918) found87 individualsjyd2 off the Dry Tortugas Florida where there is con-siderably less coral cover Preliminary data indicate that the reefs off thesoutheast Florida coast have a coral cover that amounts to only a fifth ofthe available substrate Therefore the observed average density of 251individualsm2 may not be too high for this area even in consideration ofthe patchiness mentioned above

Thus the data presently available indicate that Florida has one of themost diverse gorgonian faunas in the Caribbean by measures of eitherdensity number of species or H

2 The Scleractinia-Twenty-seven species of coral have been found onthese reefs (Table 4) A list of species numbers from various other tropicalAtlantic localities is given below for comparison

LocationJamaicaFloridaBahamasCubaCura~aoPuerto RicoBarbadosBermudaBrazilMexico

No of species62423737363533201918

ReferenceGareau amp Wells (1967)Smith (1948)Smith (1948)Duarte-Bello (1961)Roos (1971)Almy amp Carrion-Torres (1963)Lewis (1960)Laborel (1966)Laborel (1970)Kornicker et al (1959)

Many of the 42 species reported from Florida are not found north ofthe Miami area These include such important hermatypic species as


Porites porites Acropora palmata and Diploria strigosa Others such asD labyrinthiformes Colpophyllia natans and Acropora cervicornis arenot common and do not form large colonies when present Nonethelesslarger hermatypes such as Diploria clivosa A garicia lamarcki andMontastrea spp are present upon the slopes of the outer reef in quantityThe prominence of M meandrites f danae is due entirely to its abundanceon the fore-reef rubble a substrate unsuitable for colonization by othercorals A similar situation has also been noted for this species in Jamaicaby Gareau (1960)

As shown in Table 4 Montastrea cavernosa accounts for nearly 20 percent of all coral colonies Its relative abundance increases with depthand is maximal below 20 m This species also appears to be a commoncomponent of other Caribbean reefs although it is normally subordinateto M annularis However Roos (1971) has noted that the situation isreversed on the deep reefs of Cura~ao Similarly Lewis (1960) noted apredominance of M cavernosa on the turbid windward reefs of Barbadosas did Laborel (1966) on the reefs of southern Bermuda The dominanceof M cavernosa in nonoptimal outer-reef situations is perhaps no betterillustrated than on the reefs of Brazil where it completely replaces Mannularis as the dominant hermatype (Laborel 1969) Laborel citedthe relatively high degree of turbidity and concomitant low levels ofillumination for this reversal The Brazilian situation seems applicable toFloridas reefs The ridges off the southeastern Florida coast are a mileor less from the shore with its associated runoff and canal effluentsFurthermore several coastal communities operate sewage outfall pipes thatterminate directly on the outer reef These factors account for the apparentincrease in turbidity over these reefs and may be the agents responsiblefor the elimination or reduction of photophilic reef corals noted in thisstudy The reduced temperatures and southward flow of siliceous sandscited by Smith (1948) inhibit coral growth farther north but appear to beinsignificant in the area studied (Goldberg unpublished)

Preliminary observations in the Florida Keys indicate that the usualdominance of M annularis resumes This may be a function of theincreased distance from the Florida mainland to the outer reefs (roughly10 miles) as well as the presence of intervening islands Experimentalevidence for this hypothesis however remains for future work

ACKNOWLEDGMENTS

I thank Drs R M Adams W R Courtenay Jr S Dobkin and R BGrimm of Florida Atlantic University and Drs L G Abele F M BayerL P Thomas and G L Voss of the University of Miami for helpfulcriticism of the manuscript Mr John Larson of Florida Atlantic Uni-versity supplied the photographs and was a considerable help in the field


SUMARIO

LA ECOLOGfA DE COMUNIDADES DE CORALES-OCTOCORALES

FRENTE A LA COSTA SUDESTE DE LA FLORIDA

GEOMORFOLOGfA COMPOSICI6N DE

ESPECIES y ZONACI6N

Se describen tres terrazas submarinas paralelas localizadas a 10 largode la costa sudeste de la Florida extendiendose desde Miami hasta elCondado de Palm Beach La parte central de esta area cerca del sur delCondado de Palm Beach fue analizada con respecto a su geomorfologfacomposici6n de la comunidad y zonaci6n de la marca de la marea bajahasta una profundidad de 50 m Aquf se encuentran 27 especies de coralesescleractinios y 39 especies de gorgonias que definen una comunidad dearrecife coralino tfpico mas al norte de 10 que hasta ahora se habfaconocido La diversidad de gorgonias es maxima a una profundidad de15-20 m mientras las escleractinias son mas diversas en aguas mas bajasLos estudios de biomasas de gorgonias indican una inclinaci6n hacianumeros mayores de individuos pequeuos en ambientes de baja diversidady un numero menor de individuos mayores en ambientes de alta diversidadUna densidad media de 251 coloniasjm2 da a estos arrecifes la mayorconcentraci6n de gorgonias hasta ahora registrada en la regi6n del Caribe

LITERATURE CITEDALMY C AND C CARRION-ToRRES

1963 Shallow water stony corals of Puerto Rico Caribb J Sci 3(2-3)133-162

BAYER F M1956 Octocorallia Pp 166-231 in Moore R C (Ed) Treatise on in-

vertebrate paleontology Part F Coelenterata Geo Soc AmNew York

1961 The shallow-water Octocorallia of the West Indian Region MartinusNijhoff The Hague 373 pp

CARY L R1918 The Gorgonacea as a factor in the formation of coral reefs Pubis

Carnegie Instn No 213 341-362CHAMBERLAIN C

1966 Some Octocorallia of Isla de Lobos Veracruz Mexico BrighamYoung Univ Stud (Geo) 13 47-54

CRAIG A K S DOBKIN R B GRIMM AND J B DAVIDSON1969 The gastropod Siphonaria pectinata A factor in the destruction of

beachrock Am Zool 9(3) 895-901DUANE D B AND E P MEISBURGER

1969 Morphology and sediments of the nearshore Continental ShelfMiami to Palm Beach Florida Tech Memo US Army Corps ofEngineers Coastal Engineering Research Center No 29

1973] Goldberg Ecology of Coral-Octocoral Communities 487DUARTE-BELLO P P

1961 Corales de los arrecifes cubanos Ser Educac Acuario NacionalNo2 85 pp

GINSBURG R N1956 Environmental relationships of grain size and constituent particles in

some south Florida carbonate sediments Bull Am Ass PetrolGeol 40(10) 2384-2427

GONZALEZ-BRITO P1970 Una lista de los Octocorales de Puerto Rico Caribb J Sci 10(1-2)

63-69GaREAu T F

1959 The ecology of Jamaican coral reefs I Species composition andzonation Ecology 40( 1) 67-90

]960 On the physiological ecology of the coral M eandrina braziliensis(Milne Edwards amp Haime) in Jamaica (Abstract) Trans 3rdConf Ass Island Marine Labs

GaREAu T F AND J W WELLS1967 The shallow-water Scleractinia of Jamaica Revised list of specIes

and their vertical distribution range Bull Mar Sci 17(2) 442-453

KINZIE R A1970 The ecology of the gorgonians (Cnidaria Octocorallia) of Discovery

Bay Jamaica Unpublished Dissertation Yale University 107 ppKORNICKER L S F BONET R CANN AND C M HOSKIN

1959 Alacran Reef Campeche Bank Mexico Pubis lnst mar SciUniv Tex 6 1-22

LABOREL J1966 Contribution a lt~tude des madreporaires des Bermudes (systematique

et repartition) Bull Mus natn Hist nat Paris Ser 2 38 281-300]969 Les peuplements de madreporaires des cotes tropicales du Bresil

Ann Univ Abidjan Ser E Ecologie 2(3) 1-260] 970 Madreporaires et hydrocoralliaires recifaux des cotes bresiliennes

systematique ecologie repartition vertic ale et geographique AnnisInst oceanogr Paris 47(1) 171-229

LEWIS J B1960 The coral reefs and coral communities of Barbados WI Can 1

Zoo 38 1133-1145PIE LOU E C

1966 The measurement of diversity in different types of biological collec-tions J theor BioI 13 131-144

Roos P J1971 The shallow-water stony corals of the Netherlands Antilles Stud

Fauna Curacao 37(130) 1-108SHALER N S

] 890 The topography of Florida Bull Mus compo ZooI Harv 16 139-158

SMITH F G W1948 Atlantic reef corals University of Miami Press Miami Florida

112 ppVAUGHAN T W

1910 A contribution to the geologic history of the Floridian PlateauPap Tortugas Lab 4(33) 99-185


WAINWRIGHT S A AND J R DILLON]969 On the orientation of sea fans (genus Gorgonia) BioI Bull mar

bioI Lab Woods Hole 136 130-139WELLS 1 W

1957 Coral Reefs In Treatise on marine ecology and paleoecology Vol1 Ecology Mem geol Soc Am 67 609-632

NOTE ADDED IN GALLEYS

While this manuscript was in press the following pertinent paper was pub-lishedKINZIE R A III

1973 Coral Reef Project-Papers in Memory of Dr Thomas F Goreau 5The zonation of West Indian gorgonians Bull Mar Sci 23(1)93-155


()

bull ~uPiter Inlet

bull PALM bull ~ BEACH COUNTY Boca RotoL bullbull

[23(3)

Dry [qrtugos

(~7 (U 0_1~~~~~I West I

820 81deg

I inch = 38miles

= reefs

I80deg

FIGURE 1 The area of investigation with localities mentioned in the text

nature of these structures is not known They may be coral reefs or theymay consist of a Recent coral veneer overlying Pleistocene beachrockFor the purposes of this paper the term reeflike ridges will be abbre-viated and the term reef along with patch reef reef tract etcwill be used where physiographically applicable without regard to thenature or depth of the antecedent platform This report intends to de-scribe the biological community presently living upon and presumablycontributing to these geological structures


ForeReefZone

FirstReef


Zone OuterReef

Platform

Outer ReefSlope

a 600 800 1200METERS












RESULTS








[23(3)








ORP 560ORS 250



ORP 125ORS 466








ORP 2000ORS 466FR 1400


TABLE 1 (Continued)

471


PR 1500ORP 2000ORS 3000FR 1400

PR 250ORP 125

PR 1500



ORP 125



























SS 1500DS 2000FR 500











ORP 560SS 1500FR 033




TABLE 2 (Continued)

[23(3)



















PRORPDSPRORP

PRORPSSSSFRDS

PRS5ORP

PR

PRSSPRORPS5FR

700560560

2000166560125700750

1500125

750560200

1400125033

700750125700333

1500750200040033067

040040

067040

040040

166750040033


TABLE 2 (Continued)

475



Ellisella sp




DS 125FR 3000





47630


951

[23(3)

~o

10

middot5

---- --- -134 --_


-bull

115


PLATFORM








Ciz 20

10 400 500 600100 200 300

BIOMASS (g)














DISCUSSION


























No of species62423737363533201918







ACKNOWLEDGMENTS



SUMARIO




ESPECIES y ZONACI6N


















63-69GaREAu T F



















112 ppVAUGHAN T W










ForeReefZone

FirstReef


Zone OuterReef

Platform

Outer ReefSlope

a 600 800 1200METERS












RESULTS








[23(3)








ORP 560ORS 250



ORP 125ORS 466








ORP 2000ORS 466FR 1400


TABLE 1 (Continued)

471


PR 1500ORP 2000ORS 3000FR 1400

PR 250ORP 125

PR 1500



ORP 125



























SS 1500DS 2000FR 500











ORP 560SS 1500FR 033




TABLE 2 (Continued)

[23(3)



















PRORPDSPRORP

PRORPSSSSFRDS

PRS5ORP

PR

PRSSPRORPS5FR

700560560

2000166560125700750

1500125

750560200

1400125033

700750125700333

1500750200040033067

040040

067040

040040

166750040033


TABLE 2 (Continued)

475



Ellisella sp




DS 125FR 3000





47630


951

[23(3)

~o

10

middot5

---- --- -134 --_


-bull

115


PLATFORM








Ciz 20

10 400 500 600100 200 300

BIOMASS (g)














DISCUSSION


























No of species62423737363533201918







ACKNOWLEDGMENTS



SUMARIO




ESPECIES y ZONACI6N


















63-69GaREAu T F



















112 ppVAUGHAN T W
















RESULTS








[23(3)








ORP 560ORS 250



ORP 125ORS 466








ORP 2000ORS 466FR 1400


TABLE 1 (Continued)

471


PR 1500ORP 2000ORS 3000FR 1400

PR 250ORP 125

PR 1500



ORP 125



























SS 1500DS 2000FR 500











ORP 560SS 1500FR 033




TABLE 2 (Continued)

[23(3)



















PRORPDSPRORP

PRORPSSSSFRDS

PRS5ORP

PR

PRSSPRORPS5FR

700560560

2000166560125700750

1500125

750560200

1400125033

700750125700333

1500750200040033067

040040

067040

040040

166750040033


TABLE 2 (Continued)

475



Ellisella sp




DS 125FR 3000





47630


951

[23(3)

~o

10

middot5

---- --- -134 --_


-bull

115


PLATFORM








Ciz 20

10 400 500 600100 200 300

BIOMASS (g)














DISCUSSION


























No of species62423737363533201918







ACKNOWLEDGMENTS



SUMARIO




ESPECIES y ZONACI6N


















63-69GaREAu T F



















112 ppVAUGHAN T W











[23(3)








ORP 560ORS 250



ORP 125ORS 466








ORP 2000ORS 466FR 1400


TABLE 1 (Continued)

471


PR 1500ORP 2000ORS 3000FR 1400

PR 250ORP 125

PR 1500



ORP 125



























SS 1500DS 2000FR 500











ORP 560SS 1500FR 033




TABLE 2 (Continued)

[23(3)



















PRORPDSPRORP

PRORPSSSSFRDS

PRS5ORP

PR

PRSSPRORPS5FR

700560560

2000166560125700750

1500125

750560200

1400125033

700750125700333

1500750200040033067

040040

067040

040040

166750040033


TABLE 2 (Continued)

475



Ellisella sp




DS 125FR 3000





47630


951

[23(3)

~o

10

middot5

---- --- -134 --_


-bull

115


PLATFORM








Ciz 20

10 400 500 600100 200 300

BIOMASS (g)














DISCUSSION


























No of species62423737363533201918







ACKNOWLEDGMENTS



SUMARIO




ESPECIES y ZONACI6N


















63-69GaREAu T F



















112 ppVAUGHAN T W










TABLE 1 (Continued)

471


PR 1500ORP 2000ORS 3000FR 1400

PR 250ORP 125

PR 1500



ORP 125



























SS 1500DS 2000FR 500











ORP 560SS 1500FR 033




TABLE 2 (Continued)

[23(3)



















PRORPDSPRORP

PRORPSSSSFRDS

PRS5ORP

PR

PRSSPRORPS5FR

700560560

2000166560125700750

1500125

750560200

1400125033

700750125700333

1500750200040033067

040040

067040

040040

166750040033


TABLE 2 (Continued)

475



Ellisella sp




DS 125FR 3000





47630


951

[23(3)

~o

10

middot5

---- --- -134 --_


-bull

115


PLATFORM








Ciz 20

10 400 500 600100 200 300

BIOMASS (g)














DISCUSSION


























No of species62423737363533201918







ACKNOWLEDGMENTS



SUMARIO




ESPECIES y ZONACI6N


















63-69GaREAu T F



















112 ppVAUGHAN T W





















SS 1500DS 2000FR 500











ORP 560SS 1500FR 033




TABLE 2 (Continued)

[23(3)



















PRORPDSPRORP

PRORPSSSSFRDS

PRS5ORP

PR

PRSSPRORPS5FR

700560560

2000166560125700750

1500125

750560200

1400125033

700750125700333

1500750200040033067

040040

067040

040040

166750040033


TABLE 2 (Continued)

475



Ellisella sp




DS 125FR 3000





47630


951

[23(3)

~o

10

middot5

---- --- -134 --_


-bull

115


PLATFORM








Ciz 20

10 400 500 600100 200 300

BIOMASS (g)














DISCUSSION


























No of species62423737363533201918







ACKNOWLEDGMENTS



SUMARIO




ESPECIES y ZONACI6N


















63-69GaREAu T F



















112 ppVAUGHAN T W










TABLE 2 (Continued)

[23(3)



















PRORPDSPRORP

PRORPSSSSFRDS

PRS5ORP

PR

PRSSPRORPS5FR

700560560

2000166560125700750

1500125

750560200

1400125033

700750125700333

1500750200040033067

040040

067040

040040

166750040033


TABLE 2 (Continued)

475



Ellisella sp




DS 125FR 3000





47630


951

[23(3)

~o

10

middot5

---- --- -134 --_


-bull

115


PLATFORM








Ciz 20

10 400 500 600100 200 300

BIOMASS (g)














DISCUSSION


























No of species62423737363533201918







ACKNOWLEDGMENTS



SUMARIO




ESPECIES y ZONACI6N


















63-69GaREAu T F



















112 ppVAUGHAN T W










TABLE 2 (Continued)

475



Ellisella sp




DS 125FR 3000





47630


951

[23(3)

~o

10

middot5

---- --- -134 --_


-bull

115


PLATFORM








Ciz 20

10 400 500 600100 200 300

BIOMASS (g)














DISCUSSION


























No of species62423737363533201918







ACKNOWLEDGMENTS



SUMARIO




ESPECIES y ZONACI6N


















63-69GaREAu T F



















112 ppVAUGHAN T W









47630


951

[23(3)

~o

10

middot5

---- --- -134 --_


-bull

115


PLATFORM








Ciz 20

10 400 500 600100 200 300

BIOMASS (g)














DISCUSSION


























No of species62423737363533201918







ACKNOWLEDGMENTS



SUMARIO




ESPECIES y ZONACI6N


















63-69GaREAu T F



















112 ppVAUGHAN T W












Ciz 20

10 400 500 600100 200 300

BIOMASS (g)














DISCUSSION


























No of species62423737363533201918







ACKNOWLEDGMENTS



SUMARIO




ESPECIES y ZONACI6N


















63-69GaREAu T F



















112 ppVAUGHAN T W



















DISCUSSION


























No of species62423737363533201918







ACKNOWLEDGMENTS



SUMARIO




ESPECIES y ZONACI6N


















63-69GaREAu T F



















112 ppVAUGHAN T W

































No of species62423737363533201918







ACKNOWLEDGMENTS



SUMARIO




ESPECIES y ZONACI6N


















63-69GaREAu T F



















112 ppVAUGHAN T W













ACKNOWLEDGMENTS



SUMARIO




ESPECIES y ZONACI6N


















63-69GaREAu T F



















112 ppVAUGHAN T W










SUMARIO




ESPECIES y ZONACI6N


















63-69GaREAu T F



















112 ppVAUGHAN T W














63-69GaREAu T F



















112 ppVAUGHAN T W









articulos octocorales white papers cientificos investigaciones fauna arrecifal

Documents