Southern Caribbean azooxanthellate coral

communities off Colombia

Javier Reyes, Nadiezhda Santodomingo, Adriana Gracia, Giomar Borrero-Pérez,

Gabriel Navas, Luz Marina Mejía-Ladino, Adriana Bermúdez, Milena Benavides

Instituto de Investigaciones Marinas y Costeras, INVEMAR, Cerro Punta Betín,

AA 1016, Santa Marta, Colombia
([email protected])

Abstract. As a result of the explorations carried out by the Colombian Marine and
Coastal Research Institute (INVEMAR) between 1998-2002 along the Colombian
Caribbean continental shelf and upper slope, the occurrence of azooxanthellate
coral banks was suspected at three sites (from the northern to southern Colombian
Caribbean coast): off La Guajira Peninsula, at a water depth of 70 m; off Santa Marta,
at 200 m, and nearby the San Bernardo Archipelago, at 150 m). Each site exhibited
particular bottom features (relief and substrate), suggestive of reef structures. The
analysis of the fauna collected by bottom trawling at these sites showed that many
of the fishes, mollusks, echinoderms, crustaceans, antipatharians, soft corals and
bryozoans collected are characteristic dwellers of hard substrates or reef bottoms.
At the first site (Guajira) the hard coral Cladocora debilis, was the most abundant;
a total of 156 species of invertebrates and fishes were identified among the material
collected at this site. At the second site (Santa Marta), 13 scleractinian species were
collected, but Madracis myriaster, was the dominant species; another 102 species
of invertebrates and fishes were also found. At the third site (San Bernardo) 19
scleractinian species were found, M. myriaster being the dominant. A total of 135
species of invertebrates and fishes were collected at this site. It is presumed that
deep-sea coral banks have developed in these three settings, since many of the
collected species are known to be hard or reef bottom dwellers.

Keywords. Colombian Caribbean, biodiversity, deep-water communities, azoo-

xanthellate corals, mollusks, echinoderms, fishes, crustaceans

Introduction
The most extensive and unknown ecosystems on earth are on the sea floor
(Snelgrove et al. 1997). The majority of marine bottoms are sedimentary plains,
showing quite homogeneous features and similar oceanographic characteristics and
biota, which are considered cosmopolitan in distribution (Snelgrove et al. 1997).
However, within those extensive plains there are small areas like seamounts,

Freiwald A, Roberts JM (eds), 2005, Cold-water Corals and Ecosystems. Springer-Verlag

Berlin Heidelberg, pp 309-330
310 Reyes, Santodomingo, Gracia, Borrero-Pérez, Navas, Mejía-Ladino et al.

deep-sea trenches, reef-forming corals, submarine canyons, and cold seeps, whose
characteristics make possible a relative increase of the biodiversity from surrounding
soft bottom areas; these ecosystems are remarkable exceptions to the paradigm of
the deep sea as one of the most stable, and least productive biospheres on earth
(Mortensen et al. 1995; Koslow 1997; Koslow et al. 2001; Korn et al. 2003). They
exhibit unique characteristics and support a diverse community of organisms which
have been documented in limited biotic inventories and whose ecologic relationships
are so far understood (de Forges et al. 2000).
Deep-sea coral communities have been reported throughout the world oceans,
and although they seem to be constrained mainly to the North Atlantic and Pacific
temperate waters, there are also some other records in tropical waters off the African
coast, the Caribbean Antillean, and the Indopacific (Roberts and Hirschfield 2003).
Contrasting to shallow-water coral reefs, deep-sea corals do not depend on sunlight
to obtain energy, but feed on microscopic organisms from surrounding waters, and
their productivity may be related to their association with light hydrocarbon seeps
(de Forges et al. 2000). The discovery of these communities has been incidental,
and generally made by fishermen (Fosså et al. 2002). Nevertheless, during the last
decade the interest in the continental shelf communities has remarkably increased,
to the point, that there is current characterization and mapping information available
for the North Atlantic and the Pacific (Freiwald et al. 2002).
Although more than one hundred species can be associated with these unique reef
habitats, deep-sea coral communities have been classified by the main framework
species. Until now four deep-sea coral bank types have been described: Lophelia
pertusa banks (200-1000 m water depth), Oculina varicosa banks (45-150 m water
depth), Primnoa-Paragorgia forests (35-750 m water depth), and other living
habitats colonized not exclusively by corals, but also by sponges, sea anemones,
ascidians and bryozoans (Reed and Mikkelsen 1987; Freiwald et al. 2002; Krieger
and Wing 2002).
Similar to shallow-water coral reefs, whose high biodiversity is comparable
to that of the rain forest (Hubbell 1997), deep-sea corals have demonstrated that
coral framework can modify the seabed. Thus, these coral assemblages provide an
essential habitat for many species of invertebrates, including sea stars, nudibranchs,
octopuses, snails, crinoids, basket stars, sponges, and anemones, by providing
shelter, protection from currents and predators, breeding areas, spawning areas,
nurseries, food, and resting areas (Krieger 2001; Tunesi et al. 2001). Some of the
associated fauna are commercially and recreationally important such as the red
porgy, the greater amberjack, the rockfish, the Pacific Ocean perch, the flatfish,
the Atka mackerel, many species of snappers and groupers (Robins et al. 1986;
Reed and Mikkelsen 1987; Allen and Robertson 1994) and crustaceans like golden
king crab and shrimps. In addition, this biodiversity could be a potential source of
numerous marine natural products with medical, pharmaceutical and cosmetological
value (Bruckner 2002).
The Colombian continental shelf and break areas have been explored through few
 Southern Caribbean azooxanthellate coral communities off Colombia 311

expeditions, the most relevant include the R/V Oregon in the 70´s made by BCF, R/
V Pillsbury in 1972 made by RSMAS and CIOH-INVEMAR-Smithsonian in 1995
on board at R/V ANCON, from which a systematic survey was done. Based on their
results the INVEMAR carried out the ʻMacrofaunaʼ cruises (1998-2002) on board at
R/V ANCON, whose primary aim was to fill the information gaps in the inventories
of the Colombian soft bottom fauna between 20 and 520 m depth. A total of 72400
specimens were collected during the ʻMacrofaunaʼ expeditions, of which 50 % of
the species were new records for Colombia (Saavedra-Díaz et al. 2000; Lattig and
Reyes 2001; Borrero-Pérez et al. 2002a, b; Cruz et al. 2002; Gonzalez et al. 2002;
Gracia et al. 2002; Roa-Varón et al. 2003; Borrero-Pérez and Benavides-Serrato
2004; Campos et al. 2004). In total 10 new species have been described (Lattig and
Cairns 2000; Lemaitre and Bermúdez 2000; Lemaitre et al. 2001; Saavedra-Díaz et
al. 2001; Ardila and Díaz 2002; Saavedra-Díaz et al. 2003).
The principal aim of the present study is to show and discuss the occurrence
of azooxanthellate coral banks in three localities of the southwestern Caribbean
sampled by the ʻMacrofaunaʼ cruises (INVEMAR 1998-2002). The results of
this study represent new evidence of coralline banks constructed by different
azooxanthellate corals species than those already reported (Freiwald et al. 2002).
The associated faunas are quite different in each one of the three coral settings,
represented primarily by anthozoans, crustaceans, mollusks, echinoderms and
fishes. In this way, these three unique azooxanthellate coral communities sustain
one of the highest species diversities found along the Colombian continental shelf.
Due to the fact that the sampling performed by the Macrofauna cruises focused on
macrobenthic fauna living on soft bottoms, the discovery of these three localities
rich in azooxanthellate corals and their associated fauna was an unexpected result.

Abbreviations
AC: Azooxanthellate corals
BCF: Bureau of Commercial Fisheries, U.S. Department of Interior
DSC: Deep-sea corals
INVEMAR: Instituto de investigaciones Marinas y costeras
GIS-SR Lab: Geographic information systems and remote sensors laboratory at
INVEMAR
MCZ: Museum of Comparative Zoology, Harvard University, Cambridge,
Massachusetts
MHNMC: Museo de Historia Natural Marina de Colombia
NMNH: National Museum of Natural History, Washington
SIBM: Colombian marine biodiversity database
RMNH: Nationaal Natuurhistorische Museum, Leiden, The Netherlands
RSMAS: Rosenstiel School of Marine and Atmospheric Science, Miami
Senckenberg: Senckenberg Museum Frankfurt, Germany
312 Reyes, Santodomingo, Gracia, Borrero-Pérez, Navas, Mejía-Ladino et al.

Materials and methods

The sampling was carried out on board the R/V ANCON in two phases:
Macrofauna I (October 1998 - April 1999) and Macrofauna II (March - April 2001).
In total 80 trawling stations were included between off La Guajira Peninsula (12º
34'N 71º 50' W) and off Gulf of Urabá (09º 02' N 76º 02' W) at five depth ranks (20,
70, 150, 300 and 500 m). A bottom area of about 25000 m2 was swept at each station
by using a semi-balloon trawl net (ca. 9 x 1 m opening) during 20 min, and at a
vessel speed of about 3 knots (5.5 km/h). Sea-bottom relief was differentiated using
a FURUNO FE 824 echo sounder (200 kHz). On board, the samples were washed
through a sieve, sorted and preserved in 70 % and 96 % ethanol (invertebrates)
and 10 % formalin (fishes). The locations of the three sites where a significant
abundance of azooxanthellate corals and their associated organisms were collected
are shown in Table 1 and Figure 1. The specimens were identified to species level by
consulting group specific monographs and through direct examination of museum
repositories from the NMNH, MCZ, RMNH, RSMAS, Senckenberg, and MHNMC
collections. Species relevant information was stored in the SIBM data base, hosted
at http://web.invemar.org.co/redcostera1/invemar/sib.jsp.

Results
During the expeditions, three localities with a significant abundance of AC and
associated organisms were discovered unexpectedly; however, AC species in other
localities have recorded in small amounts. The AC stations were located along the
Colombian continental shelf (Fig. 1), off La Guajira Peninsula, off Santa Marta, and
off San Bernardo Islands (Table 1). Unlike other sampled bottoms of continental
shelves and upper slope, the species richness found for these three AC stations (338
species) was relatively high, with a value representing around 40 % of the species
collected throughout the whole study. Species list is shown in Appendix 1. Although
the most abundant AC and dwellers found in the three settings belong to different
species (in terms of taxonomic identification), the number of species observed in
each station was similar, the species common to all sectors remains low (11 species).
The most diverse and abundant group sampled corresponds to mollusks (Fig. 2).
Some of the echinoderms collected from these three coralline formations constitute

Table 1 Station list, samples collected in each of the coralline communities. Locality
(INVEMAR Macrofauna project station name), depth range, and initial (superior coordinates)
and final (inferior coordinates) trawl position are shown
Station Depth (m) Lat (N) Long (W)
11º 23.53' 73º27.78'
La Guajira (INV 048) 70-71
11º24.40' 73º27.62'
11º23.25' 74º12.46'
Santa Marta (INV 019) 200–220
11º23.61' 74º12.37'
San Bernardo 9º47.12' 76º13.45'
155-160
(INV 073) 9º46.61' 76º13.72'
 Southern Caribbean azooxanthellate coral communities off Colombia 313

Fig. 1 Map of the Colombian Caribbean showing the three localities where the coral
communities were discovered (stars), and where Lophelia pertusa was recorded (triangles)

the first records for the Colombian Caribbean. Many fish species associated with
coral reef and rocky bottoms were also collected in the areas mentioned.

Common species
The stony coral Madracis myriaster (Fig. 3a), common in Santa Marta and San
Bernardo, but rare in La Guajira (the shallowest site), appears to be the main coralline
matrix builder in the two first areas. Other species common to all localities were
the stony coral Anomocora fecunda (Fig. 3c), the black coral Antipathes lenta, the
alcyonarian Stereonephthya portorricensis (Fig. 3k) and the brittle-star Ophiothrix
suensonii, both of them coralline ground dwellers. Paracyathus pulchellus
(Scleractinia) was commonly found attached to other stony corals skeletons; crabs
belonging to the genera Mithrax and Stenorhyncus, and the fishes Synagrops bella
and Antigonia capros were also recorded.

Associated biota, general characteristics

La Guajira. Water depth 70 m. Cladocora debilis (thin tube coral; Fig. 3b)
was the main coral matrix builder, its sympodial budding and recumbent shapes
provide more surfaces on which sponges, bryozoans, octocorals and tunicates
can settle. Other scleractinian species such as Madracis myriaster (Fig. 3a),
314 Reyes, Santodomingo, Gracia, Borrero-Pérez, Navas, Mejía-Ladino et al.

Fig. 2 Species number by major taxa (cnidarians, mollusks, crustaceans, echinoderms and
fishes) in each of the coral communities: La Guajira, Santa Marta, and San Bernardo

Anomocora fecunda (Fig. 3c), Anomocora prolifera, and Paracyathus pulchellus

also occasionally occur.
A total of 156 species were collected in this locality, where mollusks were the
most diverse taxon (59 species). Bivalves like Chlamys munda and Arca zebra,
and gastropods like Petaloconchus erectus, and Vermicularia spirata were the most
abundant mollusk species. V. spirata was often found attached to sponges and to C.
debilis in small clusters with bryozoans. V. spirata is a facultative coral associate,
using it as refuge or preying on it (Reed and Mikkelsen 1987). Echinoderms were
the most abundant taxon, with 1367 specimens (Fig. 2). The crinoid Analcidometra
armata was the dominant echinoderm, it has been reported as a common species
attached to the gorgonians in the Caribbean Sea (Meyer 1973; Hendler et al. 1995).
Ophiothrix angulata (Fig. 3u) was found in several stations along the sampling area,

Fig. 3 Invertebrates and fishes collected in the Southern Caribbean deep-sea coral banks
off Colombia. The measures given in parenthesis following the species names correspond
to the specimen approximate height (invertebrates) and length (fishes). The species shown
in this figure were selected to be hard bottom dwellers or associated to their surrounding
areas. Cnidarians: a Madracis myriaster [10 cm]; b Cladocora debilis [3 cm]; c Anomocora
fecunda [3 cm]; d Thalamophyllia riisei [5 cm]; e Caryophyllia berteriana [5 cm]; f Nicella
cf. guadalupensis [4 cm]; g Chrysogorgia desbonni [8 cm]; h Callogorgia sp. [11 cm];
i Nidalia rubripunctata [7 cm]; j Antipathes n. sp. [4 cm]; k Stereonephthya portorricensis
[11 cm]. Mollusks: l Coralliophila caribaea [2 cm]; m Coralliophila squamosa [2 cm];
 Southern Caribbean azooxanthellate coral communities off Colombia 315

n Pseudosimnia vanhyningi [1.3 cm]; o Babelomurex dalli [2.4 cm]; p Sthenorytis

pernobilis [1.6 cm]. Crustaceans: q Uroptychus uncifer; r Anomalothir frontalis; s Latreillia
elegans. Echinoderms: t Asteroschema cf. oligactes [3 cm]; u Ophiothrix angulata [4 cm];
v Ophioderma rubicundum [6 cm]; w Astrocnida cf. isidis [10 cm]; x Pentacrinus sp. [14
cm]. Fishes: y Apogon pseudomaculatus [3 cm]; z Fistularia petimba [15 cm]; aa Bollmannia
sp. [2 cm]
316 Reyes, Santodomingo, Gracia, Borrero-Pérez, Navas, Mejía-Ladino et al.

but it was especially abundant in the La Guajira community. O. angulata has also
been reported on Oculina varicosa banks off Florida (Reed and Mikkelsen 1987).
In the La Guajira setting, coral reef dweller fishes such as the family Apogonidae
(Apogon affinis, A. quadrisquamatus and A. pseudomaculatus) were common
(Palacio 1974; Uyeno et al. 1983; Böhlke and Chaplin 1993; Cervigón 1993; Allen
and Robertson 1994). Seagrass-inhabitant species like Paradiplogrammus bairdi,
were also found there.
Santa Marta. Water depth 200 m. 13 scleractinian species were collected.
Madracis myriaster (Fig. 3a), Coenosmilia arbuscula, Anomocora fecunda and
the solitary Polymices fragilis, were the most abundant species. Here, the stony
corals are characterized by bushy shaped colonies (M. myriaster, A. fecunda and
C. arbuscula) or individual polyps (Polymices fragilis and Javania cailleti), both
forms have strong bases attached directly to the rock.
Another 102 species of fishes, echinoderms, mollusks, crustaceans, and cnidarians
were also found. However, in contrast to La Guajira, other organisms did not
significantly encrust the coral skeletons. The species of black corals and octocorals
were also numerous, especially Antipathes columnaris, Aphanipates abietina; and
species of the genus Stichopathes, and the octocorals Chrysogorgia desbonni (Fig. 3g),
Trychogorgia lyra, Nidalia sp. (Fig. 3i) and Nicella cf. guadalupensis (Fig. 3f). The
abundance of mollusks was low; the gastropods found were anthozoan´s dwellers,
which usually prey on it; Sthenorytis pernobilis (Epitonidae) (Fig. 3p) on sea
anemones; Pseudosimnia vanhyningi (Ovulidae) (Fig. 3n) on octocorals;
and Babelomurex dalli (Fig. 3o), and Coralliophila squamosa (Fig. 3m)
(Coralliophilidae) on stony corals (Keen 1971; Cate 1972; Reed and Mikkelsen
1987), all of these constitute first records for the Colombian Caribbean (Gracia et
al. 2004).
In Santa Marta, Ophiothrix suensonii, commonly found on gorgonians, recorded
by Hendler et al. (1995) from shallow to deep reef zones and the sea cucumber
Holothuria lentiginosa enodis previously recorded off Florida from Oculina varicosa
banks (Pawson et al. 1982), were found (Borrero-Pérez et al. 2003). Another 14
species were found exclusively in this area; three of them, Stephanasterias cf.
albula, Trigonocidaris albida, and Palaeobrissus hilgardi were the first records for
the southern Caribbean.
The crustacean fauna found in the three coralline formations presents no
major exclusive coral associations, however, in Santa Marta species of the family
Chyrostilidae like Uroptychus sp., were recorded as hosted in octocorals of the
Chysogorgia genus (Pequegnat and Pequegnat 1970). Some crab species collected,
like those of the genera Anomalothir and Latreillia, are known to be common
inhabitants of rubble shells and sandy bottoms (Williams 1984). Palicus affinis,
P. alternatus, P. gracilipes and P. sicus, exhibit no particular habitat preferences
(Rathbun 1918), but were found associated with coral grounds only in the Santa
Marta and San Bernardo areas.
Rocky-bottom fishes like Gymnothorax poligonius, Chlorophthalmus
agassizi, Bellator egretta, Antigonia capros, Antigonia combatia, and Pontinus
nematophthalmus were the first records for the Colombian Caribbean.
 Southern Caribbean azooxanthellate coral communities off Colombia 317

San Bernardo. Water depth: 155-160 m. 19 scleractinian species were found,

among which, Anomocora fecunda, Coenosmilia arbuscula, Thalamophyllia riisei
(Fig. 3d) and Madracis myriaster were dominant. Eguchipsammia cornucopia,
mostly devoid of living tissue, was also common. Other less abundant scleractinians
were also found attached to the four species mentioned above. Three different
growth forms of stony corals were present: recumbent (T. riisei and E. cornucopia);
bushy (M. myriaster and A. fecunda); and solitary (Caryophyllia berteriana, Fig.
3e, and Coenocyathus parvulus). The San Bernardo site corresponds to an extensive
carbonate deposit formed during the Pleistocene epoch over older strata deformed
by tectonism and diapiric intrusion (Vernette 1989a).
Beside corals, a total of 115 species of invertebrates and fishes were collected
at this site. San Bernardo was the most diverse sector in echinoderms with 38
species (Fig. 2); brittle stars such as Ophioderma appressum, Astrocnida cf. isidis
(Fig. 3w), Asteroporpa cf. annulata, Asteroschema cf. laeve and Asteroschema cf.
oligactes (Fig. 3t) were found attached to stony corals and gorgonians. Also, rare
species, like Ophiosyzygus disacanthus, previously recorded as dwelling in hard
bottoms (Turner and Heyman 1995; Borrero-Pérez and Benavides-Serrato in press),
the pedunculate crinoids Democrinus conifer (Macurda and Meyer 1976), and the
coral reef crinoid Nemaster rubiginosus (Hendler et al. 1995) were also present in
San Bernardo. The mollusk fauna here has no restricted coral dwellers, except the
gastropod Coralliophila caribaea, 10 species of bivalves were found.
In San Bernardo, five fish species associated with coral reefs (Allen and
Robertson 1994; Cervigón 1994) were caught. Fistularia petimba, Bollmannia sp.,
Serranus atrobranchus, Decodon puellaris and Pristigenys alta, although they were
not the most abundant species collected (Fig. 3). While Citharichthys cornutus,
Lipogramma evides and Neomerinthe beanorum were the most frequent species
caught.

Discussion
The Colombian seabed still remains poorly known. The efforts addressed to the
study the shallow-water coral reefs and seagrasses during the last three decades have
been increased, and data about characterization and mapping of them are already
available (Díaz et al. 2000, 2003), but shallow-coral reefs and seagrasses, represent
only less than 1 % of the Colombian marine bottoms (SIG-SR Lab). Recently,
other complex ecosystems occurring on the continental shelf and slope around the
world are being discovered and studied mainly in North Atlantic and Pacific oceans.
DSC banks belong to those recently described ecosystems (Mortensen et al. 1995;
Freiwald et al. 2002; Korn et al. 2003). According to the results of this study, three
new records of azooxanthellate coral communities occurring on the Colombian
continental shelf are documented. The records registered off La Guajira, Santa
Marta and San Bernardo were fortuitous, because the Macrofauna expeditions were
focused to explore the soft bottoms of the continental shelf and slope between 20
and 500 m. In 77 of the 80 stations sampled no major amounts of corals on hard
318 Reyes, Santodomingo, Gracia, Borrero-Pérez, Navas, Mejía-Ladino et al.

substrates were found. Occasionally, in some sampling areas, where rough relief
was observed, trawling was avoided to prevent net damage; for that reason, more
areas where deep-sea coral communities occur would be expected. Further research
aiming to map and study their biodiversity is necessary to assess the value of these
deep-sea coral ecosystems and to propose conservation measures.
The results suggest that the deep-sea coral communities observed on the
Colombian continental shelf have a similar ecological role (in terms of dwelling
and food supply) to those reported in temperate regions. The fauna composition of
the stony coral framework here however is different. Lophelia spp., Oculina spp.
and Madrepora spp. were uncommon species; Madracis myriaster and Cladocora
debilis seem to be the main frame building species found. M. myriaster belongs to
the apozooxanthellate coral group defined by Best (2001), which developed adaptive
strategies to survive in colder waters; this adaptive response could derive from the
facultative presence of zooxanthellae, followed by the migration to deep waters.
This kind of adaptation explains the extensive bathymetric range of M. myriaster
namely 20-700 m in the West Atlantic (Best 2001).
On the other hand, the Lophelia pertusa records in the Oceana Report (Roberts
and Hirschfield 2003) give some evidence of the possible occurrence of Lophelia
banks off Venezuela. In addition, some other non-living Lophelia records from the
R/V Pillsbury (Cairns 1979) and INVEMAR expeditions in Colombian waters at
500 m depth, suggest that Lophelia could have occurred there in the recent past, or
could be alive there now and forming coral banks in the surrounding waters.
The differences in stony coral species composition found among the surveyed
localities indicate some differences in environmental conditions. Cladocora debilis
is the most important coral established in La Guajira formation. This suggests that
the growth of recumbent species is favored of a homogeneous non-consolidated
bottom in this locality. In fact, sedimentary data sets indicate that deposited material
on the coast off La Guajira comes from upper Tertiary and Quaternary alluviums
(Thomas and MacDonald 1969). Metamorphic rocks from the Sierra Nevada de
Santa Marta mountain system (Gonzáles and Rodríguez 2003) form a favourable
substrate for the corals settled at Santa Marta site. There, the coralline fauna was
mainly represented by species with corallums showing an extended base attached
to the rock (Polymices fragilis and M. myriaster). In San Bernardo both corallum
shapes were common, suggesting a heterogeneous substrate that probably promoted
the variability in the corallum forms settled there. Apparently, the attached species
grow on limestone substrate, which is composed mainly of a fossil reef terrace built
during the last marine regression about 20 ky ago when the sea level was 120 m
below the current one (Vernette 1989b). This fossil reef terrace has developed on
diapiric domes or mud vulcanoes which can be related to the active plate tectonics
off the Colombian Caribbean continental margin (Vernette 1989c).
Although the hard corals that build framework belonged to different species,
they exhibit a similar shape characterized by recumbent growth with secondary
corallites located both sides of the principal axis (see Figs. 3a-e). The branching
pattern displayed by Anomocora fecunda (Fig. 3c) and Coenosmilia arbuscula,
 Southern Caribbean azooxanthellate coral communities off Colombia 319

was quite similar, and sometimes establishing differences between them was very
difficult. Some specimens exhibit a transitional form, which indicates a possible
hybridization process, analogous to that described for shallow coral reefs (Vollmer
and Palumbi 2002). Other evidence suggests that corallum shape convergence is a
common phenomenon; species previously described by Cairns (2000) as reptoid
colonies (Thalamophyllia riisei; Fig. 3d) or as individual polyps (Caryophyllia
berteriana; Fig. 3e), showed a branching pattern similar to that of Cladocora debilis
(Fig. 3b). Such convergence suggests an environmental factor response, analogous
to those corallum-shape changes showed by some species in shallow coral reefs,
responding to hydrodynamic factors.
The differences among the faunas associated with the three DSC communities
give some hints about the environmental conditions of the surrounding waters. At
La Guajira site, sponges and ascidians (suspension feeders) were abundant; this
fact indicates an environment enriched with suspension material, perhaps produced
by the upwelling conditions typical of the area. Epizoic fauna was common in La
Guajira coral community; sponges, ascidians, bryozoans, antipatharians, octocorals,
and scleractinians were settled on Cladocora debilis (Fig. 3b) skeletons. Sponges
also provide substrates to other fauna; the scleractinian Paracyathus pulchellus
which was often observed settled on them. In San Bernardo some epizoites were
noticed whereas in Santa Marta the epizoites were rare.
Tropical coral reefs present a fragmented distribution due to their specific
ecological requirements to develop complex structures (Birkeland 1997). Some
typical shallow reef species were found associated with these deep-sea coral
communities. The brittle star Ophioderma spp., the gobiid fish Bollmannia spp. and
Apogon spp. increased their bathymetric range to deeper waters, suggesting that there
may be an ecological relationship (i.e. breeding areas) between shallow-water reef
habitats and their deep-water continental shelf counterparts. This example suggests
that distributional patterns of some coral reef species may be characterized by a
continuum rather than by isolated populations, using these deep-water environments
as stepping stone habitats between shallow reefs in order to disperse.
Different facultative or obligate epizoic interactions within these communities
were observed, including stony corals, gorgonians and sponges as substrates for
bryozoans, mollusks, echinoderms, other stony corals and sponges. Some studies
on invertebrate fauna associated with corals have demonstrated that while corals
play an important role in framework building, crinoids, sea anemones, bryozoans,
sponges, and other scleractinians, participate in the accumulation of carbonate
sediments and the accretion of coralline matrix (Sulak and Ross 2003).
Further investigation will be focused on studying the biological and ecological
relationships in these settings. It is necessary to carry out research on habitat
characterization and mapping, as well as on the assessment of impacts caused by
trawl fishing in order to support arguments for the sustainable use and conservation
of these unique biological communities.
320 Reyes, Santodomingo, Gracia, Borrero-Pérez, Navas, Mejía-Ladino et al.

Acknowledgements
Instituto Colombiano para el Desarrollo de la Ciencia y la Tecnología “Francisco
José de Caldas” (COLCIENCIAS), INVEMAR through the projects Macrofauna
(Grants No. 210509-10401 and 210509-11248). Stephen Cairns, David Pawson,
Cynthia Ahearn, Rafael Lemaitre, Thomas Munroe and Jerry Harasewich of the
National Museum of Natural History, Smithsonian Institution. Patricia Lattig, Juan
Manuel Díaz, Adela Roa, Néstor Ardila, Lina Saavedra, Luz Stella Mejía, Andrea
Polanco, Arturo Acero, Néstor Campos and the work team of the Museo de Historia
Natural Marina de Colombia. José E. Polo, Jóse González, Jorge Viaña, Franklin
and Gregory, crew members of the R/V Ancón. Special thanks to Georges Vernette
and Blanca Posada (Geosciences team), Daniel Rozo (GIS-RS lab), and Alexandra
Hiller (Justus Liebig Universität). To Jaime Garzón, Sven Zea, Don Wilson and
Kathryn Scanlon for their valuable comments which improved this paper.

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Appendix 1

Species list of invertebrates and fishes collected in the coralline settings off Colombia. La
Guajira (GUA), Santa Marta (SMA) and San Bernardo (SBE). The new records (NR) for
Colombian Caribbean are shown
Species Coral Setting
Group NR
GUA SMA SBE
Cnidarians Acanthogorgia schrammi × × +
Acryptolaria conferta × +
Amphiantus caribaea × +
Anomocora fecunda × × ×
Anomocora prolifera × ×
Antipathes atlantica ×
Antipathes cf. salix × +
Antipathes columnaris × × +
Antipathes furcata ×
Antipathes gracilis × ×
Antipathes lenta × × ×
Antipathes n. sp. ×
Aphanipathes abietina × +
Balanophyllia bayeri × +
Balanophyllia cyathoides × × +
Balanophyllia palifera × × +
Balanophyllia pittieri ×
Balanophyllia wellsi × +
Bellonella rubistella × +
Callogorgia sp. × +
Caryophyllia ambrossia ×
Caryophyllia barbadensis × +
Caryophyllia berteriana × ×
Caryophyllia sp. × ×
Chrysogorgia desbonni × +
Chrysogorgia sp. ×
Cladocora debilis ×
Coenocyathus parvulus × +
Coenosmilia arbuscula × ×
Deltocyathus calcar ×
Diodogorgia sp. ×
Eguchipsammia cornucopia × +
Javania cailleti × ×
Leipsiceras pollens ×
Madracis asperula ×
Madracis myriaster × × ×
Madracis pharensis ×
Madrepora carolina × +
Nicella guadalupensis × +
Nicella sp.1 ×
Nicella sp.2 ×
Nidalia cf. occidentalis × +
Nidalia dissidens × × +
Oxysmilia rotundifolia ×
Paracyathus pulchellus × ×
Placogorgia tenuis × +
Polycyathus mayae × +
Polymyces fragilis × +
Renilla muelleri ×
Riisea sp. ×
Stereonephthya portorricensis × × × +
 Southern Caribbean azooxanthellate coral communities off Colombia 325

Appendix 1 continued
Species Coral Setting
Group NR
GUA SMA SBE
...Cnidarians Stichopathes cf. lutkeni × +
Stichopathes cf. occidentalis ×
Stichopathes pourtalessi × +
Stichopathes n. sp. ×
Swiftia sp. ×
Thalamophyllia riisei × +
Thelogorgia vossi ×
Thesea sp.1 ×
Trichogorgia lyra ×
Villogorgia nigrensis × +
Viminella sp. × +
Crustaceans Aepinus septemspinosus × +
Agaricochirus alexandri ×
Anasimus latus ×
Anomalothir frontalis × × +
Anomalothir furcillatus × +
Anomalothir sp. ×
Arachnopsis sp. ×
Batrachonotus fragosus ×
Calappa sulcata × ×
Callidactylus asper ×
Chasmocarcinus cylindricus × ×
Chyrostilidae sp.2 ×
Chyrostilidae sp.3 ×
Clythrocerus moreirai × +
Collodes trispinosus × +
Cyclodorippe antenaria × +
Ethusa mascarone ×
Euchirograpsus americanus × ×
Eucratopsis crassimanus × +
Euphrognata rastellifera ×
Homola barbata × +
Iliacantha subglobosa ×
Latreilliia elegans × +
Macrocoeloma eutheca × +
Macrocoeloma septemspinosus × +
Mesorhoea sexspinosa ×
Mithrax cornutus × ×
Munida benedicti × +
Munida evermani × +
Munida flinti × +
Munida sp.3 ×
Munida sp.4 ×
Munidopsis platirostris × +
Munidopsis sp.1 ×
Osachila antillensis ×
Paguristes sp.1 ×
Palicus affinis × +
Palicus alternatus ×
Palicus gracilipes × +
Palicus sicus × × +
Panoplax depressa ×
Pantomus sp. × ×
Parapontophylus gracilis × +
Parthenope agona ×
Parthenope fraterculus ×
Parthenope pourtalessi × +
326 Reyes, Santodomingo, Gracia, Borrero-Pérez, Navas, Mejía-Ladino et al.

Appendix 1 continued
Species Coral Setting
Group NR
GUA SMA SBE
...Crustaceans Persephona crinita × +
Petrochirus diogenes ×
Plesionika acanthonotus × +
Plesionika longicauda × +
Plesionika longipes × +
Plesionika miles × +
Plesionika tenuipes × +
Podochela gracilipes ×
Pylopaguropsis atlantica × +
Pylopagurus discoidalis × × +
Pyromaia acanthina ×
Pyromaia propinqua × +
Rhodochirus rosaceus ×
Stenocionops furcata ×
Stenorhynchus seticornis × ×
Stenorhynchus yangii × × × +
Uroptychus sp. ×
Xylopagus tayrona ×
Echinoderms Agassizia excentrica × ×
Amphioplus sp.1 ×
Amphioplus sp.2 ×
Analcidometra armata ×
Asteroporpa cf. annulata ×
Asteroschema cf. laeve ×
Asteroschema cf. oligactes × ×
Astrocnida cf. isidis ×
Astropecten alligator ×
Bathyplotes natans × +
Brissopsis elongata ×
Centrostephanus sp. ×
Clypeaster euclastus ×
Clypeaster lamprus ×
Coccometra sp. ×
Coelopleurus floridanus × ×
Comactinia meridionalis × ×
Coronaster sp. ×
Crinometra brevipinna ×
Democrinus conifer ×
Dipsacaster sp. ×
Pentacrinus sp. × ×
Eucidaris tribuloides ×
Genocidaris maculata ×
Histampica cf. duplicata × ×
Holothuria lentiginosa enodis × +
Holothuria occidentalis ×
Hypalometra defecta ×
Leptonemaster venustus × ×
Marginaster cf. pectinatus ×
Mediaster sp. ×
Nemaster discoidea ×
Nemaster rubiginosus ×
Neocomatella cf. pulchella ×
Ophiacantha sp.1 ×
Ophiacantha sp.2 ×
Ophiacantha sp.3 ×
Ophiactis algicola ×
Ophiactis savignyi ×
 Southern Caribbean azooxanthellate coral communities off Colombia 327

Appendix 1 continued
Species Coral Setting
Group NR
GUA SMA SBE
…Echinoderms Ophiochondrus cf. convolutus ×
Ophioderma appressum ×
Ophioderma rubicundum ×
Ophiomitra cf. valida ×
Ophiomitrella cf. laevipellis ×
Ophiomyxa cf. tumida ×
Ophiomusium acuferum × × ×
Ophiomusium eburneum ×
Ophiomusium cf. testudo × ×
Ophiomusium validum ×
Ophionereis dolabriformis ×
Ophiopaepale cf. goesiana ×
Ophiophragmus riisei ×
Ophioplax cf. ljungmani × ×
Ophiopristis cf. hirsuta × ×
Ophiopsila hartmeyeri ×
Ophiostigma isocanthum ×
Ophiosyzygus disacanthus × +
Ophiothrix angulata ×
Ophiothrix orstedii ×
Ophiothyreus cf. goesi ×
Ophiothrix suensonii × × ×
Ophiura cf. acervata × × ×
Palaeobrissus hilgardi × +
Paleopneustes cristatus ×
Paleopneustes tholoformis ×
Persephonaster echinulatus ×
Plutonaster agassizi agassizi × ×
Pteraster sp. ×
Rosaster cf. alexandri ×
Stephanasterias cf. albula ×
Stylocidaris affinis × ×
Stylocydaris lineata × × +
Stylometra cf. spinifera ×
Tamaria cf. halperni ×
Trigonocidaris albida × +
Fishes Ancylopsetta cycloidea ×
Antigonia capros × × × +
Antigonia combatia × × +
Apogon affinis ×
Apogon pseudomaculatus ×
Apogon quadrisquamatus ×
Bathyanthias mexicana × +
Bellator brachychir ×
Bellator egretta × +
Bembrops anatirostris × ×
Bollmannia sp. ×
Bregmaceros atlanticus × ×
Chlorophthalmus agassizi × +
Citharichthys cornutus ×
Citharichthys gymnorhinus ×
Decodon puellaris ×
Engyophrys sentus ×
Fistularia petimba ×
Gymnothorax polygonius × +
Halieutichthys aculeatus ×
Holanthias martinicensis × ×
328 Reyes, Santodomingo, Gracia, Borrero-Pérez, Navas, Mejía-Ladino et al.

Appendix 1 continued
Species Coral Setting
Group NR
GUA SMA SBE
…Fishes Kathetostoma cubana ×
Lipogramma evides × +
Lophiodes reticulatus ×
Monacanthus ciliatus ×
Pontinus nematophthalmus × × +
Priacanthus arenatus ×
Prionotus beani ×
Prionotus stearnsi ×
Pristigenys alta ×
Saurida brasiliensis ×
Saurida caribbaea × +
Scorpaena agassizii ×
Scorpaena calcarata ×
Serranus atrobranchus × ×
Serranus chionaraia ×
Serranus sp.1 ×
Serranus sp.2 ×
Serranus tortugarum ×
Steindachneria argentea ×
Syacium micrurum ×
Syacium gunteri ×
Symphurus piger × × ×
Synagrops bella × × × +
Synagrops spinosus ×
Synodus poeyi ×
Thalassophryne maculosa ×
Mollusks Abra sp. ×
Abra longicallis ×
Acesta colombiana ×
Acteon sp. ×
Anadara sp. ×
Arca zebra ×
Arcopsis adamsi ×
Arene sp. ×
Babelomurex dalli × +
Barbatia candida × ×
Bellaspira pentagonalis ×
Calliostoma fernandezi ×
Calliostoma sp. ×
Capulus sp.1 ×
Capulus sp.2 ×
Cardiomya sp. ×
Chama congregata ×
Chama macerophylla ×
Cheilea equestris ×
Chlamys munda ×
Circomphalus strigillinus ×
Cocculina sp. ×
Cochlespira radiata × +
Conus cf. daucus ×
Conus cf. granarius ×
Coralliophila caribaea ×
Coralliophila squamosa × +
Cosmioconcha nitens ×
Crucibulum cf. marense ×
Crucibulum sp. ×
Cuspidaria sp. ×
 Southern Caribbean azooxanthellate coral communities off Colombia 329

Appendix 1 continued
Species Coral Setting
Group NR
GUA SMA SBE
…Mollusks Cypraea cassis ×
Cypraea cinerea ×
Dentalium sp. ×
Dentimargo sp. ×
Diodora cayennensis × ×
Diodora sp.1 ×
Diodora sp.2 ×
Discotectonica discus × +
Distorsio cf. mcgintyi ×
Douglassia sp. ×
Entodesma beana ×
Eratoidea hematita ×
Eudolium crosseanum ×
Eudollium thompsoni ×
Fusinus lightbourni ×
Glyphostoma gabbi ×
Hypselodoris sp. ×
Laevicardium sybariticum ×
Laevidentalium callipeplum × +
Limaria sp. × ×
Limopsis antillensis ×
Loligo sp. ×
Macoma tenta ×
Mitrolumna biplicata ×
Myrtea prystiphora ×
Nassarius scissuratus ×
Nassarius hotessieri ×
Nemocardium sp. ×
Nemocardium tinctum ×
Notocrater sp. ×
Nuculana cestrota ×
Octopus sp. ×
Olivella cf. acteocina ×
Parvamussium pourtalesianum ×
Pecten chazaliei ×
Persicula pulcherrima ×
Petaloconchus erectus ×
Pitar albida ×
Pitar arestus ×
Pitar cf. albida ×
Pleurobranchus sp. ×
Plicatula cf. gibbosa ×
Polystira tellea × × +
Poromya rostrata × ×
Prunum marginatum ×
Pseudosimnia vanhyningi × +
Pyrgospira sp. ×
Semirossia tenera × +
Sthenorytis pernobilis × +
Tellina persica ×
Trachipollia didyma ×
Tucetona pectinata ×
Ventricolaria listeroides ×
Vermicularia spirata ×
Volvarina cf. avena ×
Volvarina sp.1 ×
Volvarina sp.2 ×
330 Reyes, Santodomingo, Gracia, Borrero-Pérez, Navas, Mejía-Ladino et al.

Appendix 1 continued
Species Coral Setting
Group NR
GUA SMA SBE
…Mollusks Xenophora caribaea ×
Xenophora conchyliophora ×
TOTAL 338 species 156 115 134 84