Northeast Gulf Science Volume 11 Article 1 Number 1 Number 1
7-1990 Observations on the Size, Predators and Tumor- Like Outgrowths of Gorgonian Octocoral Colonies in the Area of Santa Marta, Caribbean Coast of Colombia Leonor Botero Instituto de Investigaciones Marinas de Punta de Betin INVEMAR
DOI: 10.18785/negs.1101.01 Follow this and additional works at: https://aquila.usm.edu/goms
Recommended Citation Botero, L. 1990. Observations on the Size, Predators and Tumor-Like Outgrowths of Gorgonian Octocoral Colonies in the Area of Santa Marta, Caribbean Coast of Colombia. Northeast Gulf Science 11 (1). Retrieved from https://aquila.usm.edu/goms/vol11/iss1/1
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OBSERVATIONS ON THE SIZE, PREDATORS AND TUMOR-LIKE OUTGROWTHS OF GORGONIAN OCTOCORAL COLONIES IN THE AREA OF SANTA MARTA, CARIBBEAN COAST OF COLOMBIA
Leonor Botero
Institute de Investigaciones Marinas de Punta de Betin INVEMAR Apartado Aereo 1016 Santa Marta, Colombia South America
ABSTRACT: Gorgon ian communities of the Santa Marta area are dominated by species with large numbers of large colonies (height ;;.40 em) in contrast to that reported for other Caribbean sites where species with large numbers of small (<10 em in height) colonies are predominant. Cyphoma gibbosum and Hermodice caruncu/ata are the main predators of gorgonians in the area, but Cyphoma signatum and the fishes Alutera scripta, Acantho stracion quadricornis and Chaetodipterus faber were also observed feeding on gorgonians. Tumor-like outgrowths commonly appear in species of Pseudop/exaura and in Plexaura flexuosa and contain a chlorophyta Entocladia sp. Tubercle-like outgrowths in Plexaura homomalla, P. flexuosa, Pseudopterogorgia americana and Plexaurel/a grisea were found to contain a variety of invertebrates.
RESUMEN: Las communidades de gorgonaceos del area de Santa Marta estan dominadas por especies con gran numero de colonias grandes (altura ;;.40 em) en contraste a lo registrado en otros sitios del Caribe donde predominan especies con gran numero de colonias pequenas (altura ~10 em). Cyphoma gibbosum y Hermodice carunculata son los principales predadores de gorgonaceos en el area. Cyphoma signatum y los peces Alutera scripta, Acanthostracion quadricornis y Chaetodipterus faber tambien fueron vistos comiendo gorgonaceos. Formaciones parecidas a "tumores" aparecen comunmente en especies de Pseudoplexaura y en Plexaura flexuosa y contienen un alga clorofita, Entoc/adia sp. En Plex aura homomalla, P. flexuosa, Pseudopterogorgia americana y Plexaurella grisea, aparecen comunmente formaciones parecidas a "tuberculos" que contienen una variedad de invertebrados. Gorgonian octocorals of the area of and relatively low water transparency Santa Marta-Tayrona National Park are (compared to other Caribbean locations). abundant and diverse (Botero 1987a). The present paper adds to what is Dense gorgon ian growth most commonly known about octocoral communities of occurs (a) on hard, steep substrata adja the Santa Marta area by describing height cent to the coastline, (b) on shallow frequency distributions of species at two coral-octocoral patches in the middle of representative sites, where examples of inlets, (c) on and around large submerged the gorgonian communities of the area rocks whose tops emerge 2-3 m from the can be found. It also reports on the main water surface (piedras ahogadas) and (d) gorgonian predators of the area and on around islets. Botero (1987a,b) described tumor and tubercle-like outgrowths and quantified the zonation, density and appearing in certain species. diversity of gorgonian communities of the area. She concluded that the main MATERIALS AND METHODS environmental factors affecting their zonation and depth distribution are water The study area is located in the movement, general unavailability of hard Departamento del Magdalena, Caribbean consolidated substratum below 20-25 m Coast of Colombia (Fig. 1). Botero 1 Published by The Aquila Digital Community, 1990 1 Gulf of Mexico Science, Vol. 11 [1990], No. 1, Art. 1 2 Botero, L.
TAYRONA NATIONAL.. PARI<
+N I ,;: ~ANT~ ''I .-:~ :·:·,, ....;·.: 0 I Z IKM ·<':J:';•
Figure 1. Study area showing the location of Stations 1 and 2. (1987a,b) gives a detailed description of mainly hard metamorphic rock covered the prevailing environmental conditions by scleractinian and gorgonian corals. affecting the area. Fieldwork was carried The most abundant scleractinian species out during 1984. are Montastrea annularis, M. cavernasa, Station 1. Located on the western Dip/aria sp. and Siderastrea sp. Below side of Cinto Bay, it is exposed to strong 14-15 m depth the substratum changes trade winds blowing from the northeast to coarse sand and pebbles with some and to strong wave agitation. Its depth algal growth, and gorgonian and scler is 9-12 m. The bottom substratum is a actinian corals disappear. Gorgon ian col hard metamorphic rock platform covered onies occupy a depth range of 0.5-14 m. with small sheet (encrusting) and mound At each station, a 30 x 15m plot was like scleractinians intermingled with cal demarcated with nylon rope and sub careous sand patches and sheet and divided into 5 x 5 m quadrats. All colonies mound-like sponges. The most abundant in each quadrat were identified to species scleractinians are Mantastrea cavernosa, and their height was measured with a Dip/aria sp., and in a smaller proportion metered stick. Ca/paphyllia natans. Gorgonian colonies For coverage determinations a point are very abundant and tall, and the com -line transect method was used (Loya, munity as a whole has a characteristic 1978). Since gorgonians have a tree-like pattern given mainly by the shape, height pattern of growth, both substratum and and predominance of the gorgon ian Plex spatial (canopy) coverage determinations aurella grisea. are important. A 50 m polypropylene line Station 2. Located on the eastern marked at 0.5 m intervals was laid 1.5 m side of Chengue Bay, it is protected from above the bottom and counts were made strong wave agitation caused by the north of the number of points intercepting gor east tradewinds. Bottom slope is approx gon ian branches and stolons. Intercep imately 20-25° and the substratum is tion of branches was used for spatial https://aquila.usm.edu/goms/vol11/iss1/1 2 DOI: 10.18785/negs.1101.01 Botero: Observations on the Size, Predators and Tumor-Like Outgrowths of Octocorals in Colombia 3
(canopy) coverage determinations while and branching pattern, On the other stolon interception was used for sub hand, these may vary with age of the col stratum coverage determinations. ony, from species to species and even Counts of snails (Cyphoma gib from habitat to habitat within the same bosum) on colonies of the different species. At Station 1 coverage of the species were made once within the substratum by gorgonians was 35% and 30 x 15m submarine plot. Each colony was dominated by Plexaurella grisea, within the plot was examined. Eunicea tusca and Eunicea succinea Qualitative observations were made (Table 1), the rest of t~e substratum of tumor and tubercle-like outgrowths being covered by scleractinian corals, and of predation on gorgonians by other sponges and sand. At station 2, where a organisms. depth gradient is noticeable, highest substratum coverage by gorgonians RESULTS occurred at 8 m depth, was equal to 28% and was dominated by Gorgonia ven Coverage talina (Table 2). Canopy coverage was in Canopy coverage is defined here as all cases much higher than substratum the area of substratum shaded by, or coverage and this was expected since lying beneath the gorgonian "foliage". gorgonians occupy much more space Substratum coverage is defined as the (which is reflected on larger shaded area encrusted or directly covered by the areas on the substratum) in the water colony's basal stolon or by branches in column than on the substratum (Tables direct contact with the substratum as 1 and 2). Sites with high frequencies of can happen with very short colonies. large-sized species such as Pseudoplex Percentage coverage of gorgonian aura flagel/osa, Pseudoplexaura porosa, species is not necessarily directly Pseudoplexaura wagenaari, Plexaurella related to their frequency of occurrence grisea, Gorgonia ventalina and Plexaura or to their density. It is a measurement flexuosa have higher canopy coverage that depends mostly on colony size, size than sites where small species such as of basal attachment or stolon, shape, Eunicea succinea, Muricea muricata, TABLE 1. Substratum and canopy coverage of the different species of octocorals found at Station 1. Data indicate number of points in the line transect intercepted by octocorals. Transect was layed at 10m depth.
SPECIES CANOPY COVERAGE SUBSTRATUM COVERAGE Gorgonia ventalina 2 0 Pseudopterogorgia acerosa 2 0 Pseudopterogorgia americana 8 1 Plexaura f!exuosa 4 2 Plexaurella grisea 21 8 Plexaurella dichotoma 6 3 Pseudoplexaura wagenaari 1 0 Eunicea fusca 7 7 Eunicea succinea 7 6 Eunicea tourneforti 1 1 Eunicea asperu/a 6 3 Eunicea knighti 3 3 Eunicea ca/yculata 2 1 Subtotal 70 35 Corals, sponges, sand 30 65 Total 100 100
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TABLE 2. Substratum and canopy coverage of the different species of octocorals found at Station 2. Data indicate number of points in the line transect intercepted by octocorals. C = canopy; S = sub stratum. Transects were layed at 4, 6, 8 and 12 m depth.
DEPTH 4m 6m am 12m SPECIES c s c s c s c s Gorgonia ventalina 14 9 10 4 13 9 3 1 Pseudopterogorgia americana 1 0 0 0 4 1 13 4 Pseudopterogorgia acerosa 0 0 0 0 1 2 0 0 Plexaura f/exuosa 13 5 3 2 7 4 0 2 Plexaura homomal/a f. hom. 2 0 0 0 0 0 0 0 Plexaura homomal/a f. kuk. 4 0 1 1 5 2 2 1 Pseudoplexaura wagenaari 3 2 3 1 5 2 3 2 Pseudoplexaura flagel/osa 3 2 0 0 3 1 0 0 Pseudoplexaura porosa 0 0 0 0 9 4 0 0 Plexaurel/a dichotoma 1 0 1 0 0 0 2 0 Eunicea fusca 0 0 0 0 0 0 1 3 Eunicea succinea 0 0 0 0 0 0 1 1 Eunicea asperula 0 0 2 1 4 1 1 0 Eunicea ca/yculata 0 0 1 1 3 1 1 0 Muriceopsis f/avida 2 1 2 0 0 0 3 0 Muricea muricata 0 0 1 1 1 1 1 1 Muricea pinnata 0 0 0 0 0 0 1 0 Subtotal 43 19 24 11 55 28 32 15 Corals, sponges, sand 57 81 76 89 45 72 68 85 Total 100 100 100 100 100 100 100 100
Muriceopsis flavida and Pterogorgia Observations on Cyphoma and other citrina are most abundant. At Station '1, predators canopy coverage was 70% and was Figure 4 presents information on the dominated by Plexaurella grisea. At frequency of occurrence of C. gibbosum Station 2, canopy coverage varied on different gorgonian species and on between 30-55% being highest at 8 m the relative number of gorgonian colonies depth and was comprised mostly of a inhabited by C. gibbosum. G. ventalina combination of several species (P. f/ex and P. acerosa were the only species uosa, P. homomal/a, G. ventalina, P. having more than '10 snails on the same wagenaari, M. flavida). colony at any one time. Cyphoma sig Height natum was found only on colonies of Height frequency distributions tor Plexaurella dichotoma or Plexaurella some of the most common species in nutans. Stations '1 and 2 are shown in Figures 2 Egg capsules of C. gibbosum were and 3. P/exaurella grisea and observed on colonies of G. ventalina, P. Pseudoplexaura porosa exhibited the flexuosa, E. fusca, P. grisea, and P. largest heights (1.75-3.00 m); small dichotoma in September and October colonies of these species were rare. when water temperature was highest. Gorgonia ventalina, Plexaura flexuosa, Predation of gorgonians by other Plexaura homomalla, Pseudoplexaura species was also observed. The poly wagenaari, P/exaurella dichotoma, and chaete Hermodice carunculata was Pseudopterogorgia americana had abundant in the study area and was mostly intermediate heights (0.50-'1.25 m) often seen eating the tips of gorgonian while Muricea muricata, Eunicea fusca, branches by introducing as much as Muriceopsis f/avida, and Eunicea suc 2 em of a branch into its buccal cavity. cinea were small species (0.20-0.25 m). When the worm had finished eating, the https://aquila.usm.edu/goms/vol11/iss1/1 4 DOI: 10.18785/negs.1101.01 Botero: Observations on the Size, Predators and Tumor-Like Outgrowths of Octocorals in Colombia 5
00 M. murleoto 50 G. venfolino
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~ ·- ._,----riJ 0 0 50 l''l-1-1 P. dleholoma 00 M. murlcoto ,. 0 ,. 0 u u z II! 00 E. eolyeulalo liJ 00 1\1. flovldo :>"' :> 0 0 liJ 0: 0 0 II."' IL IZ II. E. fueeo ~ 00 50 P. waoonoorl
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0 ~ 0 n n 0 n 0 0 n 0 0 0 0 .. 0 0 0 0 0 N n 0 0 ...... " .. :: .. "N 0 "N .. .." 0 ~ N N ~ " .. .." .. " " N N N N .. I I 0 n n' ,; n' b n 0 ~ 0 0' ' 0 ' 0 0' ' ' :;: N ' ' .. .. 0 ...... 0 .. .." "N ~ " 0 "N " 0 N" ~ .." N N N ...... " N .. N N HEIGHT (em) HEIGHT (em) Figure 2. Height frequency distributions of some of Figure 3. Height frequency distributions of some of the most abundant gorgon ian species at Station 1. the most abundant gorgonian species at Station 2. 100
:I! w 90 X f- 80 z 0 TO ~ :::> CJ) 0 eo CD CD (!) 00 u 40 :X: !:: n:all 3::: 30 "w n x3!1 CJ) II !:!:! II n::: 1!14. 20 II n=6 z n n" 14 g n"IOO n = 24 n '"~II N 10 " 8 II
cfl 0 P. flu P. ho10 P.wag P. grla !. oucc e:. tuoc M. plnn P. a ear
Published by The Aquila Digital Community, 1990 5 Gulf of Mexico Science, Vol. 11 [1990], No. 1, Art. 1 6 Botero, L. branch tip would be completely denuded of its coenenchyme and polyps, and only the bare horny axis remained. The fishes
Alutera scripta, Acanthostracion quadri- aiQOO cornis and Chaetodipterus faber were seen feeding on gorgonians, and stomach contents of C. faber revealed a substan- tial amount of G. ventalina (A.Acero comm. pers.). Tumors and tubercles Colonies of Pseudoplexaura porosa, P. flagel/osa, P. wagenaari and Plexaura gorgonin chamber flexuosa often exhibited tumor-like out
growths very similar to the "algal tumors polyp or nodules" described by Goldberg & Figure 6. Schematic representation, in cross section Makemson (1981). Tumors were flabby of the inside part of tumor-like outgrowths appearing (soft) and varied in size between 3-8 em in Pseudoplexaura porosa. in longest dimension and 2-4 cin in thick ness (Fig. 5). The external tissue or coen coenenchyme with spicules was found enchyme may or may not have polyps; (Fig. 6). In P. flexuosa, infesting algae when present on the surface, polyps were surrounded by coenenchyme, but were always retracted even if those on no chamber-like spaces surrounded by the rest of the colony were extruded. The gorgonin were observed. infesting alga was identified as a Chloro Colonies of P. homomalla, P. f/ex phyte, Entoc/adia sp. It was contained uosa, P. americana and P. grisea com within "chamber-like" spaces that were monly exhibited hard, tubercle-like out surrounded by a layer of gorgonin. In growths (Fig. 7). After dissection in the between these chambers, normal-looking laboratory, they were found to contain a variety of organisms some of which were alive and some dead. Dead barnacles and empty molluscan shells were com mon, but small crabs, shrimps, bivalves (Lithophaga sp.), sipunculids and sponges were usually found alive. They were all enclosed by a thick horny layer of gorgonin that was itself covered by coenenchyme and polyps. Communica tion with the external water column was only through very small openings. I spec ulate that these organisms had probably settled on exposed gorgonian axes and were subsequently overgrown by the octocoral's axial and cortical tissues.
DISCUSSION
Figure 5. Photograph of a tu~or-like outgrowth on Lasker & Coffroth (1983) noted that a branch of Pseudoplexaura porosa (fixed specimen). among octocorals at Carrie Bow Cay https://aquila.usm.edu/goms/vol11/iss1/1 6 DOI: 10.18785/negs.1101.01 Botero: Observations on the Size, Predators and Tumor-Like Outgrowths of Octocorals In Colombia 7
(Belize) species with low recruitment fusca, Eunicea succinea, Muricea muri rates (as evidenced by the absence of cata and Muriceopsis flavida are small small colonies) were never the most species, which are reproductively mature abundant members of the community. In at maximum heights of 30-40 em. Larger contrast, species with large numbers of colonies of these species were never small (<10 em) colonies were always the found in the area, and colonies smaller predominant species. The authors pro than 10 em were low in numbers. Colonies pose that this is due to the unstable sub of large-sized species (Bayer, 1961) such stratum of their study area which will no as Pseudoplexaura porosa, Pseudoplex longer hold colonies after they reach a aura flagellosa, Plexaurella grisea, Plex certain size. In my study area, height fre aurella nutans, Plexaurella dichotoma, quency distributions are very different Plexaura flexuosa and Plexaura from those found at Carrie Bow Cay; this homomalla were most commonly found probably reflects differences in environ in my study area with heights ~50 em. mental conditions between the two areas, Colonies of Plexaurella grisea and P. particularly substratum. Species with dichotoma smaller than 40 em in height large numbers of large colonies (>30 em) were never found, even though they were were always predominant and, in general, the dominant species at Station 1. Here, communities were dominated by large most colonies were ~1 m. Pseudoplex colonies (~40 em). The maximum height aura showed a low proportion of small of adult colonies of the different species colonies at the different stations. I sug must be taken into account. Eunicea gest that recruitment to octocoral com munities of Tayrona National Park does not occur as often as in Carrie Bow Cay or other locations where calcareous sub stratum is unstable and easily bioerod able. At Tayrona National Park, hard metamorphic rock substratum is very stable and catastrophic storms (hurri canes) have not occurred, thus, mortality of high colonies because of toppling is low, even at exposed wave surge sites. In general, I found that in agitated places in Tayrona National Park, gorgonians remain firmly attached to the substratum. Overall, colonies are longer-lived and recruitment is not common or constant. Bak & Engel (1979) described three life history patterns in scleractinians: 1) abundant juvenile recruitment; 2) abun dant vegetative reproduction and 3) low rates of recruitment and high survival. If applied to gorgonians of Tayrona National Park, I hypothesize that low rates of recruitment and high survival of colonies is the dominant pattern. Figure 7. Photograph of a tubercle-like outgrowth on Opresko (1973) reported maximum a branch of Plexaurella grisea (dried specimen). heights for species in the area of Miami.
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His values are lower, especially for the family Gorgoniidae, especially Gorgonia tall species, than those measured in spp. to be preferred by C. gibbosum over Tayrona National Park. While Plexaurella plexaurid (family Plexauridae) gorgon grisea reached 1.5-1.7 m in my study ians, and preference was not clearly, area, maximum heights in Miami were influenced by previous meals. Behety 25-46 em. Pseudoplexaura porosa attained (1980) reported that C. gibbosum pre heights of 3 m in Tayrona National Park ferred plexaurid over gorgoniid species while in the Miami area it reached a maxi in agitated places because of their mum height of 1.16 m and its average higher rigidity. In contrast, in deeper and size was 36.5 em. calmer places, C. gibbosum selected Dominance of large colonies in Tay species in relation to their texture, i.e. rona National Park can account for the softest species were preferred regard general low densities if compared to less of their rigidity. At my study site, 2 other studied Caribbean sites (Belize, species of Gorgoniidae (Pseudoptero Jamaica, Miami) where densities are gorgia acerosa and Gorgonia ventalina) higher but colonies are smaller. Wahle were the only ones found to have more (1985) and Lasker & Coffroth (1983) than 10 snails per colony at any one time reported densities of 16-30 colonies/m 2 in and they also produced the largest per Jamaica and Belize, respectively. Opresko centage of colonies (45% and 33%, (1973) and Goldberg (1973) measured respectively) with snails on them. Quali densities of gorgonian colonies as high tative observations in several sites of the as 25 colonies/m 2 in the Miami area. In study area showed more snails on G. Tayrona National Park maximum recorded ventalina than on any other species, 2 densities were 6 colonies/m • I hypothe regardless of agitation conditions and of size that the large sizes of gorgonians in the presence of plexaurid species. Where Tayrona National Park and their conse G. ventalina was low in numbers (as in quent low density are due to their higher Station 1), C. gibbosum was most com survival (longer life span), which in turn mon on the most abundant species (P/ex is a result of the stable substratum and aurella grisea at Station 1). lack of hurricanes or catastrophic storms Theodor (1964), Morse eta/. (1977, (which reportedly account for high mor 1981), Goldberg & Makemson (1981) and talities at other Caribbean sites, Cary, Goldberg et a/. (1984) reported and 1914; Woodley eta/., 1981). Cary (1914) described algal tumors similar to those concluded that protection from storms is reported here, appearing in axes of one of the major factors determining the certain gorgonians, mainly Gorgonia size of gorgonian individuals in a popu ventalina and Pseudop/exaura spp. At lation. my study area, tumors were never found Since the number of colonies of on G. ventalina. In Pseudoplexaura spp. each gorgonian species observed dif the infecting alga was identified as a fered, it is difficult to conclude whether Chlorophyte, Entocladia endozoica by the snails preferred one or several Goldberg eta/. (1984). In Tayrona National species of gorgonians over others. Fre Park the alga was identified as Ento quency of occurrence of snails on the dif c/adia sp, and possibly it could be the ferent gorgonian species could have same species. The tumors consisted of depended on the availability of species masses of this filamentous green alga rather than on preferences. Birkeland & that separated normally tightly apposed Gregory (1975) found gorgonians of the laminae of the gorgon ian horny skeleton. https://aquila.usm.edu/goms/vol11/iss1/1 8 DOI: 10.18785/negs.1101.01 Botero: Observations on the Size, Predators and Tumor-Like Outgrowths of Octocorals in Colombia 9
LITERATURE CITED Goldberg, W.M., J.C. Makemson & S.B. Colley. 1984. Entoc/adia endozoica Bak, R.P.M. & M.S. Engel. 1979. Distribu sp. nov., a pathogenic chlorophyte: tion, abundance and survival of structure, life history, physiology and juvenile hermatypic corals (Sclerac effect on its coral host. Bioi. Bull., tinia) and the importance of life 166:368-383. history strategies in the parent coral Lasker, H.R. & M.A. Coffroth. 1983. Oc community. Mar. Bioi., 54:341-352. tocoral distributions at Carrie Bow Bayer, F.M. 1961. The shallow water Oc Cay, Belize. Mar. Ecol. Progr. Ser., tocorallia of the West Indian region. 13:21-28. Martinus Nijhoff, The Hague, 373 pp. Loya, Y. 1978. Plotless and transect Behety, P. 1980. Variaciones en Ia dieta methods, In: Coral reefs: Research de Cyphoma gibbosum Linnaeus en methods. D.R. Stoddart & R.E. Johan funcion de Ia profundidad. Cienc. nes (eds.). Unesco, Paris, p. 197-217. Bioi., 5:75-84. Morse, D.A., N.C. Morse & H. Duncan. Birkeland, C. & B. Gregory. 1975. Forag 1977. Algal "tumors" in the Caribbean ing behavior and rates of feeding of sea fan Gorgonia ventalina. Proc. 3rd the gastropod Cyphoma gibbosum Int. Coral Reef Symp.: 623-629. (Linnaeus). Bull. Nat. Hist. Mus. Los Morse, D.E., A. Morse, H. Duncan & R.K. Angeles County, 20:57-67. Trench. 1981. Algal tumor in the Carib Botero, L. 1987a. Gorgonian octocoral bean octocorallian Gorgonia ven communities of the Santa Marta area, talina. II. Biochemical characteriza Caribbean Coast of Colombia: tion of the algae and first Species composition, patterns of epidemiological observations. Bull. zonation and quantitative structure. Mar. Sci., 31:399-409. Ph.D. dissertation, University of Opresko, D.M. 1973. Abundance and Delaware, 122 p. distribution of shallow-water gorgo Botero, L. 1987b. Zonaci6n de octo nians in the area of Miami, Florida. corales gorgonaceos en el area de Bull. Mar. Sci., 23:535-558. Santa Marta y Parque Nacional Theodor, J. 1964. Contribution a l'etude Tayrona, Costa Caribe Colombiana. des gorgones. II: Ecologie: La faune An. lnst. lnv. Mar. Punta Betfn, et Ia flare contenues dans des ex 17:61-80. croissances de l'axe d' Eunicel/a stric Cary, L.R. 1914. Observations upon the ta. Vie et Milieu Supp., 17:157-163. growth rate and oecology of gorgo Wahle, C.M. 1985. Habitat related pat nians. Pubis. Carnegie lnst. Wash., terns of injury and mortality among 182:79-90. jamaican gorgonians. Bull. Mar. Sci., Goldberg, W.M. 1973. The ecology of the 37(3):905-927. coral-octocoral communities off the Woodley, J.D., E.A. Chornesky, P.A. Clif Southeast Florida coast: Geomor ford, J.B.C. Jackson, L.S. Kaufman, N. phology, species composition and Knowlton, J.C. Lang, M.P. Pearson, zonation. Bull. Mar. Sci., 23:465-488. J.W. Porter, M.C. Rooney, K.W. Goldberg, W.M. & J.C. Makemson. 1981. Rylaarsdam, V.J. Tunnicliffe, C.M. Description of a tumorous condition Wahle, J.L. Wulff, A.S.G. Curtis, M.D. in a gorgonian coral associated with Dallmeyer, B.P. Jupp, M.A.R. Koehl, J. a filamentous green alga. Proc. 4th Neigel and E.M. Sides. 1981. Hur Int. Coral Reef Symp., 2:685-697. ricane Allen's impact on Jamaican
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Coral Reefs. Science, 214:749-755.
https://aquila.usm.edu/goms/vol11/iss1/1 10 DOI: 10.18785/negs.1101.01