(2013) - Santodomingo - Corals colombia.pdf

BIODIVERSITY OF CORAL REEFS
Diversity and distribution of azooxanthellate corals
in the Colombian Caribbean
Nadiezhda Santodomingo & Javier Reyes & Paola Flórez &
Isabel Cristina Chacón-Gómez & Leen P. van Ofwegen &
Bert W. Hoeksema
Received: 29 May 2012 /Revised: 31 August 2012 /Accepted: 23 September 2012 /Published online: 23 October 2012
# The Author(s) 2012. This article is published with open access at Springerlink.com
Abstract During the last decade, knowledge of azooxanthel-
late corals in the Colombian Caribbean has increased through
exploration campaigns by the Marine and Coastal Research
Institute (INVEMAR). The distribution of 142 species of
corals, including hard corals (Scleractinia 64 species), black
corals (Antipatharia 18 species), and soft corals (Octocorallia
60 species) is assessed. Statistical analyses were performed to
examine the coral species distribution through a geographic
gradient (210 stations in 8 sectors) and a bathymetric range
(10–520 m depth). Four principal patterns were observed: (1)
northeastern distribution (46 species), (2) southwestern distri-
bution (11 species), (3) association with azooxanthellate coral
bioherms (37 species), and (4) widespread (44 species). In
addition, 4 species were only found around the San Andres
Archipelago (insular pattern). Two main oceanographic fac-
tors were identified to play a role in the northeast versus
southwest coral fauna separation, La Guajira upwelling sys-
tem and the Magdalena River influx. These patterns appear to
be depth-related, since the separation between northeast and
southwest was mainly shown by the shallow-water coral
fauna, whereas most of the deep-water corals (>200 m depth)
were widely distributed along the Colombian Caribbean
coastline. These data were also analyzed from a conservation
perspective in order to propose new strategies for the protec-
tion of the Colombian Caribbean coral fauna.
Keywords Azooxanthellate corals . Colombian Caribbean .
Diversity . Scleractinia . Octocorallia . Antipatharia
Introduction
In response to the worldwide decline in marine diversity
(Gaston 2000; Pandolfi et al. 2005), a paradigm shift has
occurred towards better and more precautionary conservation
and management of marine resources (Ludwig et al. 1993;
Dayton 1998). These conservation strategies are designed
through ecological models (Gering et al. 2003) based on
which species occur within a particular area and on what
factors control their distribution (Friedlander et al. 2003).
The accomplishment of this task receives special attention in
Colombia, which is also recognized for its megadiverse ter-
restrial biota (Mittermeier et al. 1997). Owing to Colombia’s
position between the Atlantic and the Pacific, each with dis-
tinct geological, oceanographic and climatic features, it is one
of the countries with the highest marine biological diversity in
South America (Díaz and Acero 2003; Miloslavich et al.
2010). Although marine biodiversity research of Colombia
has a short history, considerable knowledge has been gathered
in the last 10 years, particularly with regard to species inven-
tories and ecosystem characterizations (e.g., Campos et al.
2004; Reyes et al. 2005, 2010; Benavides et al. 2011).
During the last decades, the Colombian continental shelf
has been explored through a few research expeditions. The
Electronic supplementary material The online version of this article
(doi:10.1007/s12526-012-0131-6) contains supplementary material,
which is available to authorized users.
N. Santodomingo
Department of Earth Sciences, Natural History Museum,
Cromwell Road,
SW7 5BD London, UK
e-mail: nsantodomingo@gmail.com
J. Reyes :P. Flórez :I. C. Chacón-Gómez
Museo de Historia Natural Marina de Colombia,
Marine and Coastal Research Institute (INVEMAR),
Cerro Punta Betin,
Santa Marta, Colombia
N. Santodomingo :L. P. van Ofwegen :B. W. Hoeksema (*)
Department of Marine Zoology, Naturalis Biodiversity Center,
P.O. Box 9517, 2300 Leiden, The Netherlands
e-mail: bert.hoeksema@naturalis.nl
Mar Biodiv (2013) 43:7–22
DOI 10.1007/s12526-012-0131-6

most important ones are those made by the R/V “Oregon” in
the 1960s, the R/V “Pillsbury” in 1972, and the B/I “Ancón” in
1995. Consequently, INVEMAR carried out six “Macrofauna”
cruises (1998–2002) on board of the B/I “Ancón” to fill the
information gap concerning the Colombian soft bottom fauna
between 10 and 520 m depth (Saavedra-Díaz et al. 2000; Lattig
and Reyes 2001; Borrero-Pérez et al. 2002a, b; Cruz et al. 2002;
Gracia et al. 2002, 2004; Reyes and Santodomingo 2002;
Roa-Varón et al. 2003; Borrero-Pérez and Benavides-Serrato
2004; Campos et al. 2004; Flórez-Romero et al. 2007).
The discovery of azooxanthellate coral communities
(Santodomingo et al. 2007) and methane seep ecosystems
(Gracia et al. 2012) were among the most interesting results
of these expeditions.
Earlier research on scleractinian azooxanthellate corals
inhabiting Colombian shallow waters was carried out by von
Prahl and Erhardt (1985, 1989), Werding and Sánchez (1989),
Sánchez (1995), and Díaz et al. (2000), and regarding deep
waters by Cairns (1979, 2000). The most representative stud-
ies on the taxonomy and ecology of Colombian octocorals
were carried out by Botero (1987), Sánchez (1994, 1998,
1999, 2001), and Sánchez et al. (1997, 1998). Ecological
and taxonomic research on antipatharians were performed
by Sánchez (1999) and Opresko and Sánchez (1997, 2005).
As a result of the recent marine macrofauna expeditions,
knowledge about the Colombian coral fauna showed an im-
portant growth with three new species for science (Lattig and
Cairns 2000; Reyes et al. 2009) and new records for the region
(Reyes 2000; Lattig and Reyes 2001; Reyes et al. 2005;
Chacón-Gómez et al. 2008, 2010, 2012).
Most results of these expeditions have only been pre-
sented in technical reports with no attempt to understand
species distribution patterns. Therefore, the main goal of the
present study is to analyze and compare the spatial distribu-
tions of Colombian Caribbean azooxanthellate anthozoans
(orders Scleractinia, Alcyonacea, Pennatulacea, and
Antipatharia) along a geographical gradient (210 stations
in eight sectors) and a bathymetric range (10–520 m).
Materials and methods
Specimens were collected during the INVEMAR expedi-
tions Macrofauna I (1999), Macrofauna II (2000), Uraba
(2003), Corpoguajira (2005), and Marcoral (2005) on board
of B/I “Ancón” using an epibenthic trawl net (9×1 m open-
ing; 3 knots by 10 min), a Van Veen grab (60 l, 0.03 m2
),
and a heavy chained rocky dredge (1×0.4 m opening; 1.5
knots by 5 min). Specimens collected during previous bio-
diversity projects and material donated to INVEMAR by the
Smithonian Institution’s National Museum of Natural
History at Washington, DC (NMNH-SI) were also included
in the analysis (specimens from San Andres and three
stations at depths beyond 520 m). In this way, the study
comprised 210 sampling localities primarily covering a
depth range from 10 to 520 m depth in the Colombian
Caribbean (Table 1; Fig. 1a; ESM 2 Table S1).
Corals with polyps (in 70 or 96 % ethanol) and dry coral
skeletons were kept in the collection of the Museo de Historia
Natural Marina de Colombia (MHNMC). Specimens were
identified to species or to genus level in the case of some
octocorals and antipatharians. Some identifications were con-
firmed by comparison with type material deposited in the
NMNH-SI. Identifications were based on: Scleractinia (Cairns
1979, 2000; Zibrowius 1980); Octocorallia (Deichman 1936;
Bayer 1961; Bayer et al. 1983; Verseveldt and Bayer 1988;
Williams 1995, 1999; Sánchez and Wirshing 2005); and
Antipatharia (Brook 1889; Opresko 2001, 2002, 2003, 2004,
2006; Opresko and Sánchez 2005). The database comprised a
total of 1,226 records (online at http://siam.invemar.org.co/
siam/sibm/index.htm, accessed 30 October 2011).
Systematic surveys carried out by INVEMAR since 1999
along the Colombian Caribbean comprised eight sectors or
seascapes (INVEMAR 2000; Díaz and Acero 2003), which
are based on the topography and width of the continental
shelf, the geomorphology of the coastal zone, upwelling,
terrestrial run-off through major river outlets, and the distri-
bution of the main marine ecosystems. These eight sectors
are: La Guajira (GUA), Palomino (PAL), Tayrona (TAY),
Magdalena (MAG), Coralline Archipelagos (ARCO), Gulf
of Morrosquillo (MOR), Southern Colombian Caribbean
(DAR), and San Andres and Old Providence islands (SAN).
Bathymetric ranges include samples taken at 20, 70, 150, 200,
300, and 500 m depth. Therefore, the data was summarized in
a matrix of presence/absence records of species in each sector
of the Colombian Caribbean with their respective bathymetric
ranges (Appendix 1). Sectors and depth ranges (>2 species)
were grouped using group-average clustering based on Bray-
Curtis similarities calculated on presence/absence of coral
Table 1 Number of sampling stations of various projects carried out in
the Colombian Caribbean
Project Localities Methods
Macrofauna I (1) 61 T
Macrofauna II (1) 74 T
Uraba 2003 (1) 9 T
Corpoguajira (1) 31 T
Marcoral (1) 9 T,G,D
Biodiversity 15 S
NMNH-SI (2,3) 11 G,D
Total 210
Research vessels: 1 B/I “Ancón”, 2 R/V “Pillsbury”, 3 R/V “Oregon”
Methods: T trawling net, G van Veen grab, D rock dredge, S scuba
diving
8 Mar Biodiv (2013) 43:7–22

Fig. 1 a Distribution of sampling stations in the Colombian Caribbean in
the sectors (red bars) La Guajira (GUA), Palomino (PAL), Tayrona (TAY),
Magdalena (MAG), Coralline Archipelagos (ARCO), Gulf of Morros-
quillo (MOR), and Southern Colombian Caribbean (DAR). b–f Example
maps of distribution patterns for azooxanthellate corals: b northeastern
distribution of the octocoral Bellonella rubistella (family Alcyoniidae); c
southwestern distributions of the octocoral Nicella guadalupensis (family
Ellisellidae) and the free-living scleractinian Deltocyathus cf. italicus
(family Caryophylliidae); d associated to coral bioherms: distribution of
the octocoral Acanthogorgia aspera (family Acanthogorgiidae); e wide-
spread: distribution of the hard coral Deltocyathus calcar (family Caryo-
phylliidae); f insular pattern: distribution of the solitary scleractinian
Polymyces wellsi (family Flabellidae), from the additional material col-
lected in the sector San Andres and Providencia Archipelago (SAN)
shown in red box; white line indicates the Colombian Economic Exclu-
sive Zone (dotted line, borders in dispute)
Mar Biodiv (2013) 43:7–22 9

species data using PRIMER 6 software (Clarke and Gorley
2006). This information was sorted using non-metric multidi-
mensional scaling (MDS). The MDS analysis was performed
100 times. Distribution maps of selected species were
obtained after the incorporation of the dataset in a georefer-
enced matrix with ArcGIS™ v.9.3 software.
Results
A total of 142 species was listed (Appendix 1), consisting of
64 Scleractinia (hard corals), 55 Alcyonacea (soft corals and
gorgonians), 5 Pennatulacea (sea pens), and 18 Antipatharia
(black corals). Over 50 % of the scleractinians belonged to the
Caryophylliidae (n034; ESM 1 Figs. S1–S3), which were
followed by the Dendrophylliidae (n011; Fig. S4), the
Flabellidae (n04), and the Pocilloporidae (n04; Fig. S5), the
Oculinidae (n03; Fig. S5), the Fungiacyathidae (n02), and
the Turbinoliidae (n02), and the Gardineriidae, Guyniidae,
Rhizangiidae, and Schizocyathidae, each with only 1 species
(Fig. S5). Except for Cladocora arbuscula, all observed scler-
actinians were non-symbiotic (azooxanthellate) or facultative
symbiotic (apozooxanthellate, see “*” in Appendix 1) (Cairns
et al. 1999; Cairns and Kitahara 2012). The antipatharians
were represented by 18 species belonging to four families
(ESM 1 Fig. S6).
Octocorals (soft corals, gorgonians and sea pens) were
represented by 60 species (ESM 1 Figs. S7–S12). The order
Alcyonacea was represented by 55 species, belonging to 12
families: Plexauridae (n019; Fig. S12), Gorgoniidae (n08;
Fig. S10), Ellisellidae (n07; Fig. S9), Nidaliidae (n06;
Fig. S11), Chrysogorgiidae (n05; Fig. S8), Acanthogorgiidae
(n04; Fig. S7), and the Alcyoniidae, Anthothellidae,
Clavularidae, Nephtheidae, Keroeididae and Primnoidae, each
with only 1 species. The order Pennatulacea was represented by
5 species, which belong to three families (Fig. S13). Among the
observed alcyonaceans, Muricea elongata, Pterogorgia sp. and
Eunicea sp. are recognized as zooxanthellate (“+” in
Appendix 1), 9 other species are apozooxanthellate (“*” in
Appendix 1), while the remaining 43 species were azooxan-
thellate (Sánchez and Wirshing 2005).
Bathymetric distribution
The species richness varied along the depth range, reaching a
peak of 86 species at the 100–200 m depth range (Fig. 2). The
diversity distribution over depth intervals is symmetrical, exhib-
iting a lower number of species at the shallowest areas (<50 m)
and deepest bottoms (>200 m). However, it is important to
mention that our knowledge on azooxanthellate corals of the
Colombian Caribbean can only be considered as comprehensive
down to the 500 m depth, because only a few localities have
been sampled below this isobath (Fig. 2, ESM 2 Table S1).
Geographic distribution
A maximum number of 76 species per sector was observed in
Tayrona (Fig. 3), 17 of which were only found in this area
(Appendix 1). Coralline archipelagos (n072) were ranked
second in species richness. La Guajira (n062) and Palomino
(n061) showed almost similarly high species numbers,
whereas Magdalena (n021) and Darien (n019) scored much
lower. The lowest diversity was observed in the Gulf of
Morrosquillo sector with only 4 species of hard corals, and a
total absence of octocorals and black corals (Fig. 3). Although
no systematic surveys have been performed by INVEMAR in
the sector of San Andres and Old Providence islands (n08),
data retrieved from previous expeditions illustrate the lack of
knowledge of this area and the need of filling in this gap.
Distribution patterns
Our statistical approach clustered the coral fauna into two
main groups: (1) coral communities from deep bottoms
(>200 m) on the continental slope and (2) coral communities
on the continental shelf ≤200 m (Fig. 4a). Within the shallow
fauna group, three main clusters were observed: (2a) south-
western, conformed by stations<200 m from sectors MAG,
ARCO, and DAR; (2b) northeastern, all the stations <150 m
from sectors GUA, PAL and TAY; and (2c) coral fauna asso-
ciated to the bioherms represented by stations ARCO_150 and
TAY_200. Exceptionally, the third azooxanthellate coral bio-
herm located in the Palomino sector (PAL_70) was more
similar to the surrounding shallow-water fauna at the north-
eastern bioherm than to the other two. These four major
groups were also observed in the MDS analysis (Fig. 4b).
Based on these clusters, four principal distributional patterns
of coral species in the Colombian Caribbean are proposed:
1. Northeast: 46 coral species (32 %) with occurrences at
sectors GUA/PAL/TAY, mainly in shallow waters (32
species). Carijoa riisei (Fig. S9b), Renilla muelleri
(Fig. S14a–g), and Stichopathes sp., which exhibited a
GUA/PAL/TAY/MAG distribution, were also included
in this category since their records were restricted to a
single station at the north of the Magdalena River
mouth. The distribution of the soft coral Bellonella
rubistella is shown as an example (Figs. 1b, 5a).
2. Southwest: 11 species (8 %) distributed over sectors
MAG/ARCO/DAR. Six species inhabit shallow waters
(e.g., Nidalia occidentalis, Fig. S11f), while the other five
occurred mainly in deep waters (e.g., Stephanocyathus
diadema Fig. S3a, b). The distributions of Deltocyathus
cf. italicus and Nicella guadalupensis are shown as exam-
ples (Figs. 1c, 5b).
3. Azooxanthellate coral bioherms: 37 coral species (26 %)
exclusively living in azooxanthellate coral bioherms
10 Mar Biodiv (2013) 43:7–22

(Reyes et al. 2005; Santodomingo et al. 2007). These
bioherms are mainly constructed by Madracis asperula
(Fig. S5t), M. myriaster (Fig. S5u), Anomocora fecunda
(Fig. S1a, b), Cladocora debilis (Fig. S1n), and/or
Eguchipsammia cornucopia (Fig. S4o), and were found
in sectors Tayrona (150 m), Coralline Archipelagos
(200 m), and Palomino (70 m). Examples of this distri-
bution are the octocorals Acanthogorgia aspera (Figs. 1d,
5c) and Stereonephthya portoricensis (Fig. S11b, c), the
solitary scleractinian Coenocyathus parvulus (Fig. S1o,
p), and the antipatharian Tanacetipathes barbadensis.
4. Widespread: 44 species (31 %), 10 of which are mainly
from deep water (e.g., Stephanocyathus paliferus,
Schizocyathus fissilis), 24 mainly from shallow-water
(e.g., Diodogorgia nodulifera, Fig. S9a, and Trichogorgia
lyra, Fig. S10e), while the other 10 species occur in both
depth ranges (e.g., Caryophyllia berteriana, Fig. S1k, l,
and Oxysmilia rotundifolia, Fig. S2j, k). Deltocyathus
calcar was the most widely distributed species (Figs. 1e,
Fig. 3 Number of azooxanthellate species of Scleractinia, Alcyonacea,
Pennatulacea, and Antipatharia in eight sectors of the Colombian
Caribbean: La Guajira (GUA), Palomino (PAL), Tayrona (TAY), Coral-
line Archipelagos (ARCO), Gulf of Morrosquillo (MOR), Darien
(DAR), and San Andres and Old Providence islands (SAN)
TAY_500
GUA_500
DAR_500
PAL_500
TAY_300
DAR_300
MAG_300
ARCO_300
ARCO_500
MAG_500
DAR_150
GUA_300
MAG_20
MAG_70
DAR_20
MOR_20
ARCO_70
GUA_150
ARCO_150
TAY_200
PAL_300
DAR_70
PAL_150
TAY_150
GUA_70
PAL_70
TAY_20
TAY_70
GUA_20
PAL_20
Deep
waters
100
80
60
40
20
0
Similarity
Shallow
waters
Southwest
Northeast
Bioherms
ARCO_70
ARCO_150
ARCO_300
ARCO_500
DAR_20
DAR_70
DAR_150
DAR_300
DAR_500
GUA_20
GUA_70
GUA_150
GUA_300
GUA_500
MAG_20
MAG_70
MAG_300
MAG_500
MOR_20
PAL_20
PAL_70
PAL_150
PAL_300 PAL_500
TAY_20
TAY_70
TAY_150
TAY_300
TAY_500
TAY_200
2D Stress: 0.2
Deep
waters
Deep
waters
Shallow
waters
SW
NE
Bioherms
a
b
Fig. 4 a Bray-Curtis similarity analysis for the Colombian Caribbean
sectors based on presence/absence of azooxanthellate corals; b MDS
analysis based on presence/absence of azooxanthellate corals in the
Colombian Caribbean. Codes for sectors were simplified to La Guajira
(GUA), Palomino (PAL), Tayrona (TAY), Coralline Archipelagos
(ARCO), and Darien (DAR). Depth is indicated besides the sector
code, e.g. Palomino at 500 m 0 PAL_500
0 20 40 60 80 100
> 500 m
> 300 to 500 m
> 200 to 300 m
> 100 to 200 m
> 50 to 100 m
> 20 to 50 m
20
Depth
interval
Number of species
Number of stations
Fig. 2 Species richness distribution of azooxanthellate corals in the
Colombian Caribbean over depth intervals. The number of stations
sampled (diamonds) per depth interval is also indicated
Mar Biodiv (2013) 43:7–22 11

5f-g), followed by Caryophyllia ambrosia caribbeana
(Figs. S1g, h), and Deltocyathus eccentricus (Fig. S2c, d).
Despite a lack of systematic sampling around San
Andres and Old Providence Archipelago, four of the eight
species from here represent a distribution restricted to
these islands: i.e. Stephanocyathus (Odontocyathus) coro-
natus (Fig. S3g, h), Balanophyllia hadros, Polymyces
wellsi (Figs. 1f, S5g, h), and Fungiacyathus symmetricus
(Fig. S5k). These islands consist of oceanic atolls that are
located near Jamaica.
Endemism and similarities with the Pacific fauna
Heterocyathus antoniae (Fig. 5d, e), the only Colombian
Caribbean scleractinian endemic, has a northeastern distri-
bution pattern. This species is remarkable since congeneric
extant species were only known from the Pacific, mainly on
sandy bottoms adjacent to coral reefs in the eastern Pacific
and Indian oceans, from Mozambique to Japan (Hoeksema
and Best 1991; Cairns and Zibrowius 1997; Zibrowius
1998; Cairns 1999) and the Gulf of California (Durham
and Barnard 1952). The last occurrence of Heterocyathus
Fig. 5 Diversity of azooxanthellate corals of the Colombian Caribbe-
an. Octocorals: a Bellonella rubistella, b Nicella guadalupensis, and c
Acanthogorgia aspera. Scleractinians: d, e the endemic species
Heterocyathus antoniae, and f, g morphological variability of Delto-
cyathus calcar. Antipatharian: h Aphanipathes salix. Scales (a–c)
1 cm, (d–g) 5 mm, (h) 2 mm. More illustrations in ESM 1
12 Mar Biodiv (2013) 43:7–22

in the Tethys Ocean is known from the Miocene (20–15
MYA, Burdigalian) based on fossils recovered in southwest-
ern France (Stolarski et al. 2001).
Tethocyathus prahli represents an interesting case of
shared biota with the Pacific. This scleractinian species is
an exclusive trans-isthmic species that does not exhibit a
cosmopolitan distribution. It has been found off the
Magdalena River delta (Colombia), Cocos Island (Costa
Rica), and the northern coast of the Colombian Pacific
(Lattig and Cairns 2000; Reyes et al. 2009). Concerning
octocorals, it is noteworthy that Muricella (ESM 1 Fig. S7c,
d) and Astrogorgia (ESM 1 Fig. S12a, b) were hitherto only
known from the Indo-Pacific. In addition, specimens of
Verrucella and Ctenocella, known from Pacific shallow
waters (Hoeksema and van Ofwegen 2008) and western
Atlantic deep waters (Bayer and Grasshoff 1995), are
reported here for the first time in southern Caribbean shal-
low waters. Detailed taxonomic revisions of these four
genera and the subsequent description of potentially new
Colombian species are still needed to address further dis-
cussion on the similarities and divergences between the
Tropical Eastern Pacific and Caribbean fauna.
Discussion
Corals thrive in shallow waters, mainly on coral reefs, which
harbor the highest diversity of species known in the marine
realm (Veron 2000; Fabricius and Aldersdale 2001), espe-
cially in the center of maximum marine species richness, the
so-called Coral Triangle (Hoeksema 2007). The construc-
tion of reef structures is assisted by the symbiotic relation-
ship with zooxanthellae. However, almost 66 % of the
scleractinian coral species lack this symbiosis and are found
deeper than 50 m (Cairns et al. 1999; Cairns 2007), some of
which form reefs on cold and deep-sea bottoms (Roberts et
al. 2006). Only a few zooxanthellate coral species can be
found deeper than 50 m in tropical seas, on so-called mes-
ophotic reefs (Bongaerts et al. 2010).
During the present study, a high diversity of exclusively
azooxanthellate coral species (138 spp.) was found exploit-
ing a wider range of marine habitats such as soft-bottoms,
hard-grounds, and deep-sea coral bioherms, far beyond the
boundaries of shallow coral reefs. Thus, a higher species
richness of azooxanthellate scleractinian corals (n063) in
contrast to their zooxanthellate reef-dwelling counterparts
(n054) has been observed in the Colombian Caribbean
(Díaz et al. 2000; Reyes 2000; Reyes et al. 2010). The
difference in species richness is even more remarkable for
the antipatharians, with almost twice (n018) the number for
mesophotic and aphotic deep waters (≤520 m) in compari-
son with the 10 species previously known in shallow-water
Colombian reefs (Sánchez 1999; Opresko and Sánchez
1997, 2005). Regarding octocorals, around 70 species were
known from studies on reef communities, with about half of
the species belonging to only four zooxanthellate genera
Eunicea (n012), Pseudopterogorgia (n010), Plexaura (n0
5), and Pseudoplexaura (n05) (Botero 1987; Sánchez 1994,
1998, 1999, 2001; Sánchez et al. 1997, 1998); although the
number of azooxanthellate octocoral species is lower (n0
57), it is remarkable that most of the 33 genera were repre-
sented by one or two species, and only a few genera by more
than four (e.g. Chrysogorgia, Nidalia, and Leptogorgia).
Diversity with depth
Depth has been recognized as one of the major parameters
controlling coral species distribution (Houston 1985;
Adjeroud 1997; Dawson 2002). In this study, the coral fauna
varies in the bathymetric gradient, showing the lowest di-
versities in the shallowest and deepest bottoms, and a peak
of high diversity at 100 to 200 m depth (Fig. 2). This pattern
resembles those observed in other biogeographic provinces,
where high species richness occurs on the border of conti-
nental margins between 100 and 300 m depth, as for exam-
ple in azooxanthellate scleractinians of the Indo-Pacific
(Cairns and Zibrowius 1997) and Brazil (Kitahara 2007),
and octocorals of Japan (Matsumoto et al. 2007). The bathy-
metric pattern of Colombian azooxanthellate corals is also
similar to that exhibited by azooxanthellate scleractinians at
a global scale, which shows a high species richness at 50–
200 m depth (Cairns 2007). Despite the global diversity
peak for this group at 200–1,000 m depth (Cairns 2007),
more surveys would be required to attempt direct compar-
isons of our data with those ranges, as faunas >500 m are
still poorly sampled (Fig. 2).
Light is not a determinant factor in the distribution of
azooxanthellate corals, whereas changes in salinity, temper-
ature, nutrients, and sediments across the depth gradient can
play a significant role in the presence and richness of species
(Reyes-Bonilla and Cruz-Piñón 2000; Kitahara 2007).
Oceanographic data reported for the Colombian Caribbean
indicate that at 100–200 m depth, where the highest coral
diversity is observed (61 %), the salinity is around 36.5 psu
and temperatures oscillate between 18 and 21 °C (Andrade
and Barton 2000, 2005; Andrade et al. 2003).
Some studies have suggested that seawater calcium car-
bonate could play a major role in the biogeography of corals,
as scleractinians deposit aragonite to build their skeletons and
octocorals use calcite to form their sclerites (Buddemeier and
Fautin 1996; Buddemeier and Smith 1999). Although this
hypothesis is supported by a major development of species-
rich coral reefs in saturated waters above the aragonite satu-
ration horizon (Guinotte et al. 2006; Cairns 2007), some corals
with unique microstructural adaptations can live below car-
bonate saturation levels, such as micrabaciids, up to 5,000 m
Mar Biodiv (2013) 43:7–22 13

depth (Janiszewska et al. 2011) and Fungiacyathus, up to
6,000 m (Cairns 2007). So far, all Colombian corals have
been surveyed above the aragonite saturation horizon, includ-
ing records for Fungiacyathus crispus (up to 318 m) and F.
symmetricus (up to 842 m). There are no records for micraba-
ciid corals, and Stephanocyathus diadema, the deepest-living
coral (up to 1,257 m) in this area.
The distinction between shallow and deep waters is arbi-
trary and its extent depends on the scope of each study.
According to Cairns (2007), deep-water species are those
occurring >50 m depth based on the argument that few
zooxanthellate coral species are present below this depth
(but see Kahng and Maragos 2006; Bongaerts et al. 2010;
Kahng et al. 2010). This limit was supported by Lindner et al.
(2008) arguing that a 50-m wave disturbance boundary is used
to determine the division between ‘onshore’ and ‘offshore’
environments in the fossil record (Bottjer and Jablonski 1988),
and this corresponds approximately with the maximum depth
at which regular storm-generated waves may cause sediment
resuspension and disturbance to the benthos. In this study, a
200-m boundary was adopted to distinguish shallow- from
deep-water, based on scleractinian studies by Cairns (1979,
2000) and because it corresponds with the continental shelf
edge (Lakewood 1999; Stewart 2008). In this sense, the
presence of an ecotone area with fauna elements of the conti-
nental shelf and the upper shelf slope may explain the maxi-
mum species diversity at 100–200 m depth.
Biogeographic patterns
The coral faunas of the northeastern (GUA, PAL, and TAY)
and the southwestern (MAG, MOR, and DAR) sectors differ
in terms of species richness and composition due to the
synergy of two ecological factors. First, the cold-water
upwelling off La Guajira increases the concentration of
nutrients (Andrade et al. 2003; Andrade and Barton 2005)
and therefore creates suitable conditions to the occurrence of
suspension feeders such as corals. Second, the presence of
the Magdalena River Delta and its freshwater and sediment
discharge (Restrepo et al. 2006) could act as a barrier for the
larval dispersion of coral species.
Coral fauna in shallow waters (<200 m) seem to follow this
distribution pattern (northeastern vs. southeastern), while most
of the deep-water species are widely distributed in the
Colombian Caribbean. The relative constancy of deep-sea
environmental conditions might explain the more homoge-
neous distribution of species in deep bottoms (Grassle 1991).
The diversity of scleractinian corals, solitary ones in
particular, together with octocorals and antipatharians, was
higher in areas where azooxanthellate coral bioherms occur
(Roberts et al. 2006, 2009). This relationship is partly
explained by the presence of hard-bottom substrates where
solitary coral species and many other sessile suspension
feeders live. It is evident that branching azooxanthellate corals
(e.g., Madracis spp. Cladocora debilis, Thalamophyllia riisei,
among others) are the principal components of such hard
substrates as either living colonies or accumulated debris
(Santodomingo et al. 2007).
It is remarkable that the Morrosquillo sector has only four
species, including two common dwellers of artificial sub-
strates, Phyllangia americana americana and Astrangia sol-
itaria. This low diversity could be due to natural factors such
as high sedimentation rates on shallow (<40 m) sea floors
inside the Gulf of Morrosquillo, or to destructive fisheries,
with the latter being the most plausible. The Gulf of
Morrosquillo has been swept by shrimp trawling nets over
more than 30 years. The primary gears have been trawling
nets, both demersal and pelagic, that on average operate for
about 9 h/day (Herazo et al. 2006). Octocorals are included in
their bycatch (García et al. 2008). Consequently, it is possible
that coral diversity and abundances were higher in the past.
Unfortunately, there are no historical coral collections to sup-
port this hypothesis (see Hoeksema et al. 2011).
Relationship with the Pacific fauna
The closure of the Central American Isthmus (12–2.8 MYA)
led to a great schism in the marine realm resulting in the
extinction of some species and the development of two sepa-
rate and distinctive marine faunas: the Tropical Eastern
Atlantic from the Caribbean (Collins et al. 1996; Knowlton
and Weigt 1998; Lessios 2008). Therefore, the occurrence of
Tethocyathus prahli at both the Atlantic and Pacific Colombian
coastal areas is remarkable (Lattig and Cairns 2000; Reyes et
al. 2009) as a relict of the trans-isthmian fauna in deep waters
and could be used in further phylogeographic studies.
The presence of a center of endemism in the Colombian
Caribbean is suggested for the area surrounding La Aguja
Canyon, located between sectors PAL and TAY. This hy-
pothesis is based on the occurrence of the free-living coral
Heterocyathus antoniae (Reyes et al. 2009), belonging to a
genus previously only known from the Indo-Pacific
(Hoeksema and Best 1991), as well as the fish Quadratus
ancon (Mok et al. 2001), belonging to a West Pacific genus,
and the ophiuroid Ophiosizygus disacanthus (Borrero-Pérez
and Benavides-Serrato 2004), described from Japan. In ad-
dition, the particular distribution of these taxa suggests that
they probably belong to an Atlantic relict fauna in this
specific area of the Colombian Caribbean.
Colombian corals in the regional context
The geographic distribution patterns of azooxanthellate
scleractinian corals described by Cairns (1979, 2000) based
on geopolitical regions were successfully tested by using
statistical analyses (see fig. 1 in Dawson 2002). However,
14 Mar Biodiv (2013) 43:7–22

those analyses were carried out on only the 42 Colombian
azooxanthellate coral species known at that time. The recent
addition of 17 new species records and 3 new species (Reyes
2000; Lattig and Reyes 2001; Reyes et al. 2005; Santodomingo
et al. 2007) significatively increased the knowledge on azoox-
anthellate scleractinians in the Colombian Caribbean, which
appears to have one of the most diverse coral faunas in the
region with representatives of almost 50 % of the described
species in the western Atlantic region. Thus, although the
patterns proposed by Cairns (1979, 2000) and Dawson (2002)
explain general trends of scleractinian distribution in the
Caribbean region, they do not resemble the patterns observed
in the present study. Furthermore, distributions of octocorals
and antipatharians are also included in the present analysis as an
effort to cover the main groups of azooxanthellate anthozoans.
The current geographic distribution patterns revealed by
Colombian azooxanthellate corals seem to correspond better
with the delimitation of Marine Ecoregions of the World
(MEOW) proposed by Spalding et al. (2007). In their proposal,
the southern Caribbean MEOW (0northeastern Colombia) only
includes La Guajira sector, while the remaining sectors,
Palomino, Tayrona, Magdalena, Morrosquillo, Coralline
Archipelagos and Darien, belong to the southwestern
Caribbean MEOW (0southwestern Colombia). Based on the
criteria used to define MEOWs as “areas of relatively homoge-
neous species composition, clearly distinct from adjacent sys-
tems” (Spalding et al. 2007), our results could contribute to a
more precise delineation of MEOWs at the Colombian
Caribbean. Consequently, a new proposal for the delimitation
of MEOWs would include sectors under the influence of La
Guajira upwelling system (La Guajira, Palomino and Tayrona
sectors) within the southern Caribbean MEOW (0northeastern
Colombia), and would include the remaining sectors
(Magdalena, Morrosquillo, Coralline Archipelagos and
Darien) within the southwestern Caribbean MEOW
(0southwestern Colombia), also establishing the Magdalena
River delta as a barrier.
Our comprehensive analysis highlights not only the im-
portance of rigorous taxonomic studies but also indicates the
need for more faunistic sampling in poorly studied areas in
order to overcome Linnean and Wallacean shortfalls (Brown
and Lomolino 1998; Lomolino 2004) and to better under-
stand distribution patterns in the marine realm.
Conservation topics
Concerning the shallow-water coral reefs of Colombia, most
reef areas have been included within Marine Protected Areas
MPAs (Díaz et al. 2000), and the conservation status of nine
coral species from Colombia was emphasized in the IUCN Red
List of endangered species of Colombia (Ardila et al. 2002). The
recent discovery of three azooxanthellate coral communities of
the Colombian Caribbean and their high associated diversity of
molluscs, crustaceans, echinoderms, bryozoans, and fishes
(Reyes et al. 2005; Santodomingo et al. 2007) has promoted
the inclusion of this ecosystem in the conservation priorities for
the design and management of some MPAs (Alonso et al.
2008a, b). For instance, the establishment of the 200-m isobath
border for the Rosario and San Bernardo Coralline
Archipelagos MPA was supported by the presence of azooxan-
thellate Madracis coral bioherms between 150 and 160 m depth
within its boundaries (MAVDT 2005). Although the second
azooxanthellate coral bioherm is located in the area adjacent to
the Tayrona National Park, the management plans of this MPA
do not so far include this ecosystem. Moreover, the high coral
diversity found in La Guajira and Palomino sectors could be
affected by the current increment of trawling fishing registered
in these sectors (Viaña et al. 2002). Some strategies have been
designed for the future establishment of an MPA network in this
area (Alonso et al. 2008b), including the Palomino sector, which
contains the third azooxanthellate coral community.
The need for conservation of many reef coral species has
been established because they are relatively well known and
relatively few species are considered data-deficient (Carpenter
et al. 2008). The present study indicates that information about
the deep-water coral fauna is scarce. Future campaigns in areas
with special biodiversity values, such as La Aguja Canyon in
the Tayrona sector (down to 3,000 m depth) or around the
Lophelia pertusa records of La Guajira (500–1,000 m), would
not only provide important information to the knowledge of the
Colombian coral fauna and its deep-water coral communities
but also increase our knowledge on the global deep-water coral
fauna, which may be important for its protection.
Acknowledgements Special thanks to S. Cairns for his advice, support,
and allowing the use of the National Museum of Natural History coral
collection. Thanks to D. Opresko (Oak Ridge National Laboratory) and G.
Williams (California Academy of Science) for giving advice and literature
for the identification of antipatharians and pennatulaceans, respectively.
Thanks to J. Sánchez (Universidad de Los Andes) for fruitful discussions.
Thanks to P. Lattig for her help on identifications. G. Navas, A. Gracia, L.
S. Mejia, N. Ardila, G. Borrero, N. Cruz, M. Benavides, A. Polanco, A.
Bermudez, A. Roa, M. Díaz, and E. Montoya participated in the expedi-
tions (INVEMAR). T. López and B. Rodriguez provided some pictures of
corals. S. Braden (Smithsonian Institution) helped during SEM sessions of
Guyniidae and Pennatulacea specimens, and A. Freiwald facilitated the
use of the SEM laboratory at the Institute of Palaeontology (Erlangen-
Nuremberg University). J. Bohórquez, L. Arias and C. Garcia (LAB-SI,
INVEMAR) facilitated the access to the marine biodiversity database. The
study of deep-sea corals of the Colombian Caribbean was possible thanks
to the financial and logistic support of INVEMAR and the Instituto
Colombiano para el Desarrollo de la Ciencia y la Tecnología “Francisco
José de Caldas (COLCIENCIAS) given to the projects Macrofauna I,
Macrofauna II (2105-13-07997) and MARCORAL (2115-09-16649).
The MSc study of NS was funded by the Alβan Programme (code
E07M402757CO). The valuable comments of S.E.T. van der Meij and
three anonymous reviewers helped to improve this manuscript.
Open Access This article is distributed under the terms of the
Creative Commons Attribution Noncommercial License which permits
any noncommercial use, distribution, and reproduction in any medium,
provided the original author(s) and source are credited.
Mar Biodiv (2013) 43:7–22 15

Appendix
Table 2 Distribution of 142 species of azooxanthellate corals: 64
Scleractinia, 18 Antipatharia, 55 Alcyonacea, and 5 Pennatulacea
collected during INVEMAR expeditions in the Colombian Caribbean.
Sectors: La Guajira (GUA), Palomino (PAL), Tayrona (TAY), Coralline
Archipelagos (ARCO), Darien (DAR) and San Andres and Old Provi-
dence islands (SAN); exclusive species to one of the sectors (Ex.0•).
Geographical distribution pattern (GeoP): Widespread (Wide),
Northeast mainly in sectors GUA/PAL/TAY (NE), Southwest mainly
in sectors MAG/ARCO/DAR (SW), associated to azooxanthellate coral
bioherms present in the sectors PAL, TAY and ARCO (Bioherm),
Insular (Is) and Endemic to the Colombian Caribbean (Endemic).
Bathymetric range (BatR): (1) shallow waters, (2) shallow and deep
waters, (3) deep waters. Most are azooxanthellate species (unmarked),
except for (+) zooxanthellate and (*) apozooxanthellate
Species Code GUA PAL TAY MAG ARCO MOR DAR SAN Ex. GeoP BatR Depth
Order Scleractinia
Family Caryophylliidae
Anomocora fecunda (Pourtalès, 1871) Afec GUA PAL TAY – ARCO – DAR – Wide 1 10–200 m
Anomocora marchadi (Chevalier, 1966) Amar GUA PAL – – – – – – NE 1 50 m
Anomocora prolifera (Pourtalès, 1871) Apro GUA PAL TAY – ARCO – – – Wide 1 50–200 m
Caryophyllia ambrossia caribbeana Cairns, 1979 Caca – PAL TAY MAG ARCO – DAR – Wide 3 200–510 m
Caryophyllia barbadensis Cairns, 1979 Cbar – – – – ARCO – – – • Bioherm 1 160 m
Caryophyllia berteriana Duchassaing, 1850 Cber GUA – TAY – ARCO – – SAN Wide 2 22–293 m
Caryophyllia crypta Cairns, 2000 Ccry – – TAY – – – – – • NE 1 17 m
Cladocora arbuscula (Lesueur, 1821) +
Carb GUA PAL TAY – – – – – NE 1 10–60 m
Cladocora debilis Milne Edwards and Haime, 1849 Cdeb GUA PAL TAY MAG ARCO – – – Wide 1 10–153 m
Coenocyathus parvulus (Cairns, 1979) Cpar – PAL – – ARCO – – – Bioherm 1 21–160 m
Coenosmilia arbuscula Pourtalès, 1874 Coar – – TAY – ARCO – – – Bioherm 1 72–218 m
Colangia immersa Pourtalès, 1871 Cimm GUA – – – – – – – • NE 1 73 m
Deltocyathus calcar Pourtalès, 1874 Dcal GUA PAL TAY MAG ARCO – DAR – Wide 2(3) 107–520 m
Deltocyathus eccentricus Cairns, 1979 Decc GUA – TAY MAG ARCO – DAR – Wide 3 270–507 m
Deltocyathus italicus (Michelotti, 1838) Dita – – – MAG ARCO – – – SW 2(3) 70.9–500 m
Heterocyathus antoniae Reyes, Santodomingo and Cairns, 2009 Hant – PAL TAY – – – – – Endemic 1 21–76 m
Lophelia pertusa (Linnaeus, 1758) Lper GUA – – – – – – – • NE 3 305–314 m
Oxysmilia rotundifolia (Milne Edwards and Haime, 1848) Orot – – TAY – ARCO – – SAN Wide 2 107–238 m
Paracyathus pulchellus (Philippi, 1842) Ppul GUA PAL – – ARCO – – – Wide 2 10–269 m
Phacelocyathus flos (Pourtalès, 1878) Pflo – – – – ARCO – – – SW 1 150–180 m
Phyllangia americana (Milne Edwards and Haime, 1848) Pame GUA PAL TAY – – MOR DAR – Wide 1 10–73 m
Polycyathus mayae Cairns, 2000 Pmay – – – – ARCO – – – • Bioherm 1 113–160 m
Polycyathus senegalensis Chevalier, 1966 Psen GUA – – – – – – – • NE 1 73–152 m
Rhizosmilia maculata (Pourtalès, 1874) Rmac – – TAY – – – – – • NE 1 15–42 m
Stephanocyathus (S.) diadema (Moseley, 1876) Sdia – – – MAG ARCO – – – SW 3 502–1,257 m
Stephanocyathus (S.) isabellae Reyes, Santodomingo and Cairns
2009
Sisa GUA PAL – – ARCO – – – Wide 3 493–504 m
Stephanocyathus (S.) paliferus Cairns, 1977 Spal GUA – – MAG ARCO – DAR – Wide 3 269–510 m
Stephanocyathus (S.) laevifundus Cairns, 1977 Slae – – – – – – DAR – • SW 3 1,158–
1,225 m
Stephanocyathus (O.) coronatus Cairns, 1977 Scor – – – – – – – SAN • Is 3 750–768 m
Tethocyathus prahli Lattig and Cairns, 2000 Tprah GUA – – MAG – – – – Wide 3 152–310 m
Tethocyathus variabilis Cairns, 1979 Tvar – – – – ARCO – – – • Bioherm 1 113–160 m
Thalamophyllia riisei (Duchassaing and Michelotti, 1864) Trii – – – – ARCO MOR – SAN Wide 1 22–160 m
Trochocyathus cf. fasciatus Cairns, 1979 Tfas – – TAY – – – – – • NE 2 218 m
Trochocyathus rawsonii Pourtalès, 1874 Traw GUA PAL – – ARCO – – – Wide 2 70–308 m
Family Dendrophylliidae
Balanophyllia bayeri Cairns, 1979 Bbay – – TAY – – – – – • Bioherm 1 200 m
Balanophyllia caribbeana Cairns, 1977 Bcar GUA – TAY – ARCO – – – Wide 1 20.4–107 m
Balanophyllia cyathoides (Pourtalès, 1871) Bcya GUA – TAY – ARCO – – – Wide 1 70.5–200 m
Balanophyllia dineta Cairns, 1977 Bdin GUA – – – – – – – • NE 1 151 m
Balanophyllia hadros Cairns, 1979 Bhad – – – – – – – SAN • Is 3 238–247 m
Balanophyllia palifera Pourtalès, 1878 Bpal GUA – TAY – ARCO – – – Wide 1 22–218 m
Balanophyllia pittieri (Cairns, 1977) Bpit GUA – TAY – – – – – NE 1 70.1–200 m
Balanophyllia wellsi Cairns, 1977 Bwel GUA – – – ARCO – – – Wide 1 73–160 m
16 Mar Biodiv (2013) 43:7–22

Table 2 (continued)
Eguchipsammia cornucopia Pourtalès, 1871 Ecor – – – – ARCO – – – • Bioherm 1 117–160 m
Rhizopsammia goesi (Lindstroem, 1877) Rgoe GUA – – – – – – – • NE 1 73–152 m
Tubastraea coccinea Lesson, 1829 Tcoc GUA PAL TAY – ARCO – – – Wide 1 10 m
Family Flabellidae
Flabellum moseleyi Pourtalès, 1880 Fmos – PAL TAY MAG ARCO – DAR – Wide 3 304–520 m
Javania cailleti (Duchassaing and Michelotti, 1864) Jcai GUA – TAY – ARCO – – SAN Wide 2 113–238 m
Polymyces fragilis Pourtalès, 1868 Pfrag – – TAY – – – – – • Bioherm 2 200–218 m
Polymyces wellsi Cairns, 1991 Pwel – – – – – – – SAN • Is 3 548 m
Family Fungiacyathidae
Fungiacyathus crispus (Pourtalès, 1871) Fcri – PAL TAY MAG – – DAR – Wide 3 276–318 m
Fungiacyathus symmetricus (Pourtalès, 1871) Fsym – – – – – – – SAN • Is 3 576–842 m
Family Gardineriidae
Gardineria minor Wells, 1973 Gmin – – TAY – ARCO – – – Bioherm 1 17–107 m
Family Guyniidae
Guynia annulata Duncan, 1872 Gann – PAL TAY – – – – – NE 1 150–153 m
Family Oculinidae
Madrepora carolina (Pourtalès, 1871) Mcar – – – – ARCO – – – • Bioherm 1 108–155 m
Madrepora oculata Linnaeus, 1758 Mocu – – – – ARCO – – – • SW 3 924–950 m
Oculina tenella Pourtalès, 1871 Oten GUA PAL TAY – – – – – NE 1 21.4–150 m
Family Pocilloporidae
Madracis asperula Milne Edwards and Haime, 1849 Masp GUA PAL – – ARCO – – – Wide 1 20–153 m
Madracis brueggemanni (Ridley, 1881) Mbru – – – – ARCO – – – • Bioherm 1 107–160 m
Madracis myriaster (Milne Edwards and Haime, 1849) Mmyr GUA – TAY – ARCO MOR DAR – Wide 2 21.4–300 m
Madracis pharensis (Heller, 1868)*
Mpha – – TAY – ARCO – – – Bioherm 1 20–107 m
Family Rhizangiidae
Astrangia solitaria (Lesueur, 1817) Asol GUA PAL TAY MAG – MOR DAR – Wide 1 10–154 m
Family Schizocyathidae
Schizocyathus fissilis Pourtalès, 1874 Sfis – – TAY MAG ARCO – DAR – Wide 2(3) 158–507 m
Family Turbinoliidae
Sphenotrochus auritus Pourtalès, 1874 Saur – – TAY – – – – – • NE 1 10 m
Sphenotrochus lindstroemi Cairns, 2000 Slind – – TAY – – – – – • NE 1 10 m
Order Antipatharia
Family Antipathidae
Antipathes atlantica Gray, 1857 Atla – PAL – – – – – – • Bioherm 1 71.6 m
Antipathes furcata Gray, 1857 Afur – PAL – – – – – – • NE 1 50–70 m
Antipathes gracilis Gray, 1860 Agra – PAL TAY – ARCO – – – Bioherm 1 21–160 m
Antipathes lenta Pourtalès, 1871 Alen GUA PAL TAY – – – – – NE 2 20–300 m
Antipathes sp. Atsp – – TAY – – – – – • NE 2 200–494 m
Cirripathes paucispina (Brook, 1889) Cpau – – TAY – – – – – • NE 1 150 m
Stichopathes filiformis (Gray, 1860) Sfil – PAL – – – – – – • NE 1 150 m
Stichopathes luetkeni (Brook, 1889) Slue – PAL – – ARCO – – – Bioherm 2 160–300 m
Stichopathes occidentalis (Gray, 1860) Socc – PAL TAY – ARCO – – – Bioherm 1 70–160 m
Stichopathes pourtalesi Brook, 1889 Spou GUA PAL TAY – – – – – NE 2 70–300 m
Stichopathes sp. Stich GUA PAL TAY MAG – – – – NE 2 50–502 m
Family Aphanipathidae
Aphanipathes salix (Pourtalès, 1880) Asal – – TAY – – – DAR – Wide 1 160–200 m
Elatopathes abietina (Pourtalès, 1874) Aabi – – – – ARCO – – – • Bioherm 1 155–160 m
Rhipidipathes colombiana (Opresko and Sánchez, 1997) Aclm – – TAY – – – – – • NE 3 296 m
Family Myriopathidae
Tanacetipathes barbadensis (Brook, 1889) Tbar – PAL TAY – ARCO – – – Bioherm 1 50–160 m
Tanacetipathes spinescens (Gray, 1860) Aspi – – – – ARCO – – – • Bioherm 1 160 m
Tanacetipathes tanacetum (Pourtalès, 1880) Ttan GUA – – – – – – – • NE 3 305 m
Family Stylopathidae
Stylopathes columnaris (Duchassaing, 1870) Acol – PAL TAY – ARCO – – – Bioherm 2 150–520 m
Order Alcyonacea
Family Acanthogorgiidae
Mar Biodiv (2013) 43:7–22 17

Table 2 (continued)
Acanthogorgia aspera Pourtalès, 1867 Aasp – PAL TAY – ARCO – – – Bioherm 1 70–160 m
Acanthogorgia schrammi Duchassaing and Michelotti, 1864 Asch – PAL TAY – – – – – NE 2 70–304 m
Acanthogorgia sp. Acsp GUA PAL – – – – – – NE 1 50–50 m
Muricella sp. Muri – PAL TAY – ARCO – – – Bioherm 2 20–504 m
Family Alcyoniidae
Bellonella rubistella (Deichmann, 1936) Brub GUA PAL TAY – – – – – NE 1 70–152 m
Family Anthothelidae
Diodogorgia nodulifera (Hargitt and Rogers, 1901) Dnod GUA PAL TAY – – – DAR – Wide 2 10–300 m
Family Chrysogorgiidae
Chrysogorgia desbonni Duchassaing and Michelotti, 1864 Cdes – – – – ARCO – – – • SW 3 296–296 m
Chrysogorgia elegans Verrill, 1883 Cele GUA PAL TAY – ARCO – – – Wide 3 484–510 m
Chrysogorgia sp. Chry – – – MAG ARCO – – – Wide 2 70.5–475 m
Chrysogorgia thyrsiformis Deichmann, 1936 Cthy – – – – ARCO – – – • Bioherm 1 160–160 m
Trichogorgia lyra Bayer and Muzik, 1976 Tlyr GUA PAL TAY MAG ARCO – DAR – Wide 1 20–150 m
Family Clavulariidae
Carijoa riisei Duchassaing and Michelotti, 1860*
Crii GUA PAL TAY MAG – – – – NE 2 21.4–500 m
Family Ellisellidae
Ctenocella sp.1 *a
Cteno GUA PAL – – – – – – NE 1 20–152 m
Ellisella barbadensis (Duchassaing and Michelotti, 1864)*
Ebar GUA – – – – – – – • NE 1 50 m
Nicella guadalupensis (Duchassaing and Michelotti, 1860) Ngua – – – – ARCO – – – • SW 1 107–160 m
Nicella sp. Nicel – – TAY – ARCO – – – Bioherm 1 107–200 m
Riisea paniculata Duchassaing and Michelotti, 1860 Rpan – – TAY – ARCO – – – Bioherm 1 160–200 m
Verrucella sp.a
Verru GUA PAL TAY – ARCO – – – Wide 1 50–200 m
Viminella sp. Vimin GUA PAL TAY – ARCO – DAR – Wide 2 20–300 m
Family Gorgoniidae
Leptogorgia cardinalis (Bayer, 1961)*
Lcar GUA PAL – – – – – – NE 2 50–498 m
Leptogorgia medusa Bayer, 1952*
Lmed – PAL – – – – – – • NE 1 70 m
Leptogorgia punicea (Milne Edwards and Haime, 1857)*
Lpun GUA – – MAG – – – – Wide 1 10–20.9 m
Leptogorgia setacea Pallas, 1766*
Lset GUA PAL TAY – – – DAR – Wide 1 10–152 m
Leptogorgia sp. 1*
Lepto GUA – – – – – – – • NE 1 50 m
Leptogorgia sp. 2*
Lopho GUA PAL – MAG – – – – Wide 2 10–475 m
Pterogorgia sp.+
Ptero GUA – – – – – – – • NE 1 10 m
Tobagogorgia hardyi Sánchez and Acosta de Sánchez, 2004 Thar GUA PAL TAY – – – – – NE 1 26.6–76 m
Family Keroeididae
Thelogorgia vossi Bayer, 1991 Tvos – – TAY – – – – – • Bioherm 1 200 m
Family Nephtheidae
Stereonephthya portoricensis (Hargitt, 1901) Spor – PAL TAY – ARCO – – – Bioherm 1 20–200 m
Family Nidaliidae
Nidalia deichmannae Utinomi, 1954 Ndei – – – – ARCO – – – • SW 1 107 m
Nidalia dissidens Verseveldt and Bayer, 1988 Ndis – – TAY – – – – – • Bioherm 1 200 m
Nidalia occidentalis Gray, 1835 Nocc – – – – ARCO – – – • SW 1 107–160 m
Nidalia rubripunctata Verseveldt and Bayer, 1988 Nrub GUA PAL TAY – ARCO – – – Wide 1 20–155 m
Nidalia sp. Nidal GUA PAL – – – – – – NE 1 50–70 m
Siphonogorgia agassizii (Deichmann, 1936) Saga – – – – ARCO – – – • SW 1 107 m
Family Plexauridae
Astrogorgia sp. Amsp GUA – TAY – – – – – NE 1 73–152 m
Eunicea sp.+
Eunic GUA – – – – – – – • NE 1 10 m
Hypnogorgia pendula Duchassaing and Michelotti, 1864 Hpen – – – – ARCO – – – • Bioherm 1 160 m
Lytreia plana (Deichmann, 1936) Lplan – – – – ARCO – – – • Bioherm 1 98 m
Muricea elongata (Lamouroux, 1821)+
Melo – – TAY – – – – – • NE 1 35 m
Paracis sp. Parsp GUA PAL – – ARCO – – – Wide 1 50–160 m
Placogorgia atlantica Wright and Studer, 1889 Patl – – TAY – – – – – • NE 1 72.3–200 m
Placogorgia tenuis (Verrill, 1883) Pten – – TAY – ARCO – – – Bioherm 1 155–200 m
Scleracis guadaloupensis Duchassaing and Michelotti, 1860 Sgua – – TAY – ARCO – – – Bioherm 1 113–200 m
Scleracis pumila Reiss, 1919 Spum – – – – ARCO – – – • Bioherm 1 113–160 m
Scleracis sp. Scler GUA PAL – – ARCO – – – Wide 1 50–127 m
18 Mar Biodiv (2013) 43:7–22

References
Adjeroud M (1997) Factors influencing spatial patterns on coral reefs
around Moorea, French Polynesia. Mar Ecol Prog Ser 159:105–119
Alonso D, Ramírez LF, Segura-Quintero C, Castillo-Torres P,
Walschburger T, Arango N (2008a) Hacia la construcción de un
subsistema nacional de áreas marinas protegidas en Colombia.
Instituto de Investigaciones Marinas y Costeras—INVEMAR,
UAESPNN, TNC, Santa Marta
Alonso D, Segura-Quintero C, Castillo-Torres P, Gerhantz-Muro J
(2008b) Avances en el diseño de una red de áreas marinas prote-
gidas: estrategia de conservación para el norte del Caribe conti-
nental colombiano. Bol Invest Mar Cost 37:129–156
Andrade CA, Barton ED (2000) Eddy development and motion in the
Caribbean Sea. J Geophys Res 105:26191–26201
Andrade CA, Barton ED (2005) The Guajira upwelling system. Cont
Shelf Res 25:1003–1022
Andrade CA, Barton ED, Mooers CNK (2003) Evidence for an east-
ward flow along the Central and South American Caribbean
Coast. J Geophys Res 108:1–11
Ardila N, Navas GR, Reyes J (2002) Libro rojo de invertebrados marinos
de Colombia. Serie Libros Rojos de Especies Amenazadas de
Colombia. Instituto de Investigación de Recursos Biológicos Alex-
ander von Humboldt y Ministerio del Medio Ambiente. Bogotá
Bayer FM (1961) The shallow water Octocorallia of the West Indian
region. Stud Fauna Curaçao 12:1–373
Bayer FM, Grasshoff M (1995) Two new species of the gorgonacean
genus Ctenocella (Coelenterata: Anthozoa, Octocorallia) from
deep reefs in the western Atlantic. Bull Mar Sci 56:625–652
Bayer FM, Grasshoff M, Verseveldt J (1983) An illustrated trilingual
glossary of morphological and anatomical terms applied to Octo-
corallia. Brill, Leiden
Benavides-Serrato M, Borrero-Pérez GH, Diaz-Sanchez CM (2011)
Equinodermos del caribe colombiano I: crinoidea, asteroidea y
ophiuoridea. INVEMAR Ser Publ Esp 22:1–384
Bongaerts P, Ridgway T, Sampayo EM, Hoegh-Guldberg O (2010)
Assessing the ‘deep reef refugia’ hypothesis: focus on Caribbean
reefs. Coral Reefs 29:309–327
Borrero-Pérez GH, Benavides-Serrato M (2004) New record of Ophiosy-
zygus disacanthus Clark, 1911 (Echinodermata: Ophiuroidea: Ophio-
myxidae) in the Caribbean Sea. Proc Biol Soc Wash 117:541–544
Borrero-Pérez GH, Benavides-Serrato M, Solano O, Navas G (2002a)
Equinoideos (Echinodermata: Echinoidea) colectados en la franja
superior del talud continental del Caribe colombiano. Bol Inst Inv
Mar Cost 31:133–166
Borrero-Pérez GH, Solano OD, Benavides M (2002b) Lista revisada de
los erizos (Echinodermata: Echinoidea) del mar Caribe colom-
biano. Biota Colomb 3:137–144
Botero L (1987) Zonación de octocorales gorgonáceos en el área de
Santa Marta y Parque Nacional Natural Tayrona, Costa Caribe
colombiana. An Inst Inv Mar Punta Betín 17:61–80
Bottjer DJ, Jablonski D (1988) Paleoenvironmental patterns in the evolution
of post-Paleozoic benthic marine invertebrates. Palaios 3:540–560
Brook G (1889) The Voyage of H.M.S. Challenger. Zoology. Report on
the Antipatharia collected by H.M.S. Challenger during the years
1873–76. Zoology 32(80):1–222, pls 1–15
Brown JH, Lomolino MV (1998) Biogeography, 2nd edn. Sinauer,
Sunderland
Buddemeier RW, Fautin DG (1996) Saturation state and the evolution
and biogeography of symbiotic calcification. Bull Inst Oceánogr
Monaco 14:23–32
Buddemeier RW, Smith SV (1999) Coral adaptation and acclimatiza-
tion: a most ingenious paradox. Am Zool 39:1–9
Cairns SD (1979) The deep-water Scleractinia of the Caribbean Sea
and adjacent waters. Stud Fauna Curaçao Caribb Is 180:1–341
Cairns SD (1999) Cnidaria Anthozoa: deep-water azooxanthellate
Scleractinia from Vanuatu, and Wallis and Futuna Islands. Mem
Mus Natl Hist 180:31–167
Cairns SD (2000) A revision of the shallow-water azooxanthellate
Scleractinia of the Western Atlantic. Stud Nat Hist Caribb Reg
75:1–231
Table 2 (continued)
Swiftia exserta Duchassaing and Michelotti, 1864 Sexe – PAL – – – – DAR – Wide 1 20–70 m
Thesea bicolor Deichamnn, 1936 Tbic – PAL – – – – – – • NE 1 70 m
Thesea nutans (Duchassaing and Michelotti, 1864) Cnut – – – – ARCO – – – • Bioherm 1 98–160 m
Thesea parviflora Deichmann, 1936 Tpar GUA PAL TAY – – – – – NE 2 20–300 m
Thesea solitaria Pourtalès, 1868 Tsol – – TAY – – – – – • Bioherm 1 200 m
Thesea sp. These GUA PAL TAY – – – – – NE 1 20–154 m
Villogorgia nigrescens Duchassaing and Michelotti, 1860 Vnig – – TAY – – – – – • Bioherm 1 200 m
Villogorgia sp. Villog – PAL – – – – – – • NE 3 300 m
Family Primnoidae
Callogorgia sp. Callo – – TAY – – – – – • Bioherm 1 200 m
Order Pennatulacea
Family Kophobelemnidae
Sclerobelemnon theseus Bayer, 1959 Sthe GUA PAL TAY MAG ARCO – – – Wide 1 20–153 m
Family Renillidae
Renilla muelleri Kölliker, 1872 Rmue GUA PAL TAY MAG – – – – NE 1 10–76 m
Renilla reniformis (Pallas, 1766) Rren – PAL – MAG – – DAR – Wide 1 10–70.4 m
Family Virgulariidae
Acanthoptilum sp. Almsp GUA – TAY – ARCO – – – Wide 1 50–151 m
Stylatula diadema Bayer, 1959*
Stdia – – – – ARCO – – – • SW 1 20 m
a
Specimens were identified as belonging to the genera Ctenocella and Verrucella according to Bayer and Grasshoff (1995)
Mar Biodiv (2013) 43:7–22 19

Cairns SD (2007) Deep-water corals: an overview with special refer-
ence to diversity and distribution of deep-water scleractinian
corals. Bull Mar Sci 81:311–322
Cairns SD, Kitahara MV (2012) An illustrated key to the genera and
subgenera of the Recent azooxanthellate scleractinia (Cnidaria:
Anthozoa), with an attached glossary. ZooKeys 227:1–47
Cairns SD, Zibrowius H (1997) Cnidaria Anthozoa: Azooxanthellate
Scleractinia from the Philippines and Indonesian regions. Mem
Mus Natl Hist 172:27–243
Cairns SD, Hoeksema BW, van der Land J (1999) Appendix: list of
extant stony corals. Atoll Res Bull 459:13–46
Campos NH, Navas G, Bermúdez A, Cruz A (2004) Los crustáceos
decápodos de la franja superior del talud continental (300–500 m)
del Mar Caribe Colombiano. Monogr Fauna Colombia. 2. Instituto
de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá
Carpenter KE, Abrar M, Aeby G et al (2008) One-third of reef-building
corals face elevated extinction risk from climate change and local
impacts. Science 321:560–563
Chacón-Gómez IC, Santodomingo N, Reyes J (2008) First record of
Tobagogorgia hardyi (Octocorallia: Gorgoniidae) from the
Colombian Caribbean. Bol Inv Mar Cost 37:209–214
Chacón-Gómez IC, Santodomingo N, Reyes J, Andrade Sossa CE
(2010) Chrysogorgiidae (Octocorallia) recolectados durante las
expediciones Macrofauna I y II en el Caribe colombiano. Bol
Inv Mar Cost 39:427–445
Chacón-Gómez IC, Reyes J, Santodomingo N (2012) Deep-water
octocorals (Cnidaria: Anthozoa) collected from the Colombian
Caribbean during the Macrofauna explorations 1998–2002. Bol
Clarke KR, Gorley RN (2006) PRIMER v6: user manual/tutorial.
PRIMER-E. Plymouth Marine Laboratory, Plymouth
Collins L, Budd AF, Coates AG (1996) Earliest evolution associated
with closure of the Tropical American Seaway. Proc Natl Acad
Sci USA 93:6069–6072
Cruz N, Bermúdez A, Campos NH, Navas GR (2002) Los camarones
de la Familia Crangonidae del talud continental entre 200 y 500 m
del Mar Caribe colombiano. Bol Inv Mar Cost 31:183–203
Dawson JP (2002) Biogeography of azooxanthellate corals in the
Caribbean and surrounding areas. Coral Reefs 21:27–40
Dayton PK (1998) Reversal of the burden of proof in fisheries man-
agement. Science 279:821–822
Deichman E (1936) The Alcyonaria of the western part of the Atlantic
Ocean. Mem Mus Comp Zool Harvard Univ 53:1–317
Díaz JM, Acero A (2003) Marine biodiversity in Colombia: achieve-
ments, status of knowledge, and challenges. Gayana 67:261–274
Díaz JM, Barrios LM, Cendales MH, Garzón-Ferreira J, Geister J,
López-Victoria M, Ospina GH, Parra-Valencia F, Pinzón J,
Vargas-Angel B, Zapata FA, Zea S (2000) Áreas coralinas de
Colombia. INVEMAR Ser Publ Esp 5:1–176
Durham JW, Barnard JL (1952) Stony corals of the eastern Pacific
collected by the Velero III and Velero IV. Allan Hancock Pacific
Expeditions 1(16):1–110
Fabricius K, Aldersdale P (2001) Soft corals and sea fans: acompre-
hensive guide to the tropical shallow water genera of the central-
west Pacific, the Indian Ocean and the Red Sea. Australian
Institute of Marine Science, Townsville
Flórez-Romero P, Montoya-Cadavid E, Reyes J, Santodomingo N
(2007) Briozoos cheilostomados del Caribe Colombiano. Bol
Friedlander A, Sladek-Nowlis J, Sánchez JA, Appeldoorn R, Usseglio
P, Mccormick C, Bejarano S, Mitchell-Chui A (2003) Designing
effective marine protected areas in Seaflower Biosphere Reserve,
Colombia, based on biological and sociological information. Con-
serv Biol 17:1769–1784
García CB, Perez D, Duarte LO, Manjarres L (2008) Experimental
results with a reducing device for juvenile fishes in a tropical
shrimp fishery: impact on the invertebrate bycatch. Pan-American
J Aqua Sci 3:275–281
Gaston KJ (2000) Global patterns in biodiversity. Nature 405:220–228
Gering JC, Crist TO, Veech JA (2003) Additive partitioning of species
diversity across multiple spatial scales: implications for regional
conservation of biodiversity. Conserv Biol 17:488–499
Gracia A, Ardila NE, Díaz JM (2002) Cefalópodos (Mollusca: Ceph-
alopoda) del talud superior del Caribe colombiano. Bol Inst Inv
Mar Cost 31:219–238
Gracia A, Ardila NE, Díaz JM (2004) Gastropods collected along the
continental slope of the Colombian Caribean during the
INVEMAR-macrofauna campains (1998–2001). Iberus 22:43–75
Gracia A, Rangel-Buitrago N, Sellanes J (2012) Methane seep mol-
luscs from the Sinu-San Jacinto fold belt in the Caribbean Sea of
Colombia. J Mar Biol Ass UK 92:1367–1377
Grassle JF (1991) Deep-sea benthic biodiversity. Bioscience 41:464–
469
Guinotte JM, Orr J, Cairns S, Freiwald A, Morgan L, George R (2006) Will
human induced changes in seawater chemistry alter the distribution of
deep-sea scleractinian corals? Front Ecol Environ 4:141–146
Herazo D, Torres A, Olsen E (2006) Análisis de la composición y
abundancia de la ictiofauna presente en la pesca del camarón
rosado (Penaeus notialis) en el Golfo de Morrosquillo, Caribe
Colombiano. Rev MVZ Cordoba 11(Suppl 1):47–61
Hoeksema BW (2007) Delineation of the Indo-Malayan centre of
maximum marine biodiversity: the Coral Triangle. In: Renema
W (ed) Biogeography, time and place: distributions, barriers and
islands. Springer, Dordrecht, pp 117–178
Hoeksema BW, Best MB (1991) New observations on scleractinian
corals from Indonesia: 2. Sipunculan-associated species belong-
ing to the genera Heterocyathus and Heteropsammia. Zool Meded
65:221–245
Hoeksema BW, van Ofwegen LP (2008) Oceanic distribution ranges
and conservation status of extant soft and hard reef coral genera.
In: Leewis RJ, Janse M (eds) Advances in coral husbandry in
public Aquariums. Burger’s Zoo, Arnhem, pp 427–438
Hoeksema BW, van der Land J, van der Meij SET, van Ofwegen LP,
Reijnen BT, van Soest RWM, de Voogd NJ (2011) Unforeseen
importance of historical collections as baselines to determine biotic
change of coral reefs: the Saba Bank case. Mar Ecol 32:135–141
Houston M (1985) Patterns of species diversity in relation to depth at
Discovery Bay, Jamaica. Bull Mar Sci 3:928–935
INVEMAR (2000) Programa Nacional de Investigaciones en Biodi-
versidad Marina y Costera-PNIBM. INVEMAR-FONADE,
MMA, Santa Marta
Janiszewska K, Stolarski J, Benzerara K, Meibom A, Mazur M, Kitahara
M, Cairns SD (2011) A unique skeletal microstructure of the deep-
sea micrabaciid scleractinian corals. J Morphol 272:191–203
Kahng SE, Maragos JE (2006) The deepest, zooxanthellate scleracti-
nian corals in the world? Coral Reefs 25:254
Kahng SE, Garcia-Sais JR, Spalding HL, Brokovich E, Wagner D,
Weil E, Hinderstein L, Toonen RJ (2010) Community ecology of
mesophotic coral reef ecosystems. Coral Reefs 29:255–275
Kitahara MV (2007) Species richness and distribution of azooxanthel-
late Scleractinia in Brazil. Bull Mar Sci 81:497–518
Knowlton N, Weigt LA (1998) New dates and new rates for divergence
across the Isthmus of Panama. Proc R Soc Lond B 265:2257–2263
Lakewood CO (1999) Boundary development on the outer continental
shelf. OCS Report MMS 99-0006. US Department of the Interior
Minerals Management Service Mapping and Boundary Branch
Lattig P, Cairns S (2000) A new species of Tethocyathus (Cnidaria:
Anthozoa: Scleractinia: Caryophylliidae), a trans-isthmian azoox-
anthellate species. Proc Biol Soc Wash 113:590–595
Lattig P, Reyes J (2001) Nueve primeros registros de corales azoox-
anthellados (Anthozoa: Scleractinia) del Caribe colombiano (200–
500 m). Bol Inv Mar Cost 30:19–38
20 Mar Biodiv (2013) 43:7–22

Lessios HA (2008) The great American schism: divergence of marine
organisms after the rise of the Central American isthmus. Annu
Rev Ecol Evol Syst 39:63–91
Lindner A, Cairns SD, Cunningham CW (2008) From offshore to
onshore: multiple origins of shallow-water corals from deep-sea
ancestors. PLoS ONE 3(6):e2429
Lomolino MV (2004) Conservation biogeography. In: Lomolino MV,
Heaney LR (eds) Frontiers of biogeography: new directions in the
geography of nature. Sinauer, Sunderland, pp 293–296
Ludwig D, Hilborn R, Walters C (1993) Uncertainty, resource exploi-
tation, and conservation: lessons from history. Science 260:17–18
Matsumoto AK, Iwase F, Imahara Y, Namikawa H (2007) Bathymetric
distribution and biodiversity of cold-water octocorals (Coelenter-
ata: Octocorallia) in Sagami Bay and adjacent waters of Japan.
Bull Mar Sci 81(Suppl 1):231–251
MAVDT (2005) Resolución 0679 del 31 de mayo de 2005, por medio
de la cual se declara el Área Marina Protegida de los Archipiéla-
gos del Rosario y de San Bernardo, se adopta su zonificación
interna y se dictan otras disposiciones. MAVDT, Bogotá
Miloslavich P, Díaz JM, Klein E et al (2010) Marine biodiversity in the
Caribbean: regional estimates and distribution patterns. PLoS
ONE 5(8):e11916
Mittermeier RA, Gil PR, Mittermeier CG (1997) Megadiversity:
Earth’s biologically wealthiest nations. Conservation Internation-
al, Washington DC
Mok H, Saavedra-Díaz LM, Acero A (2001) Two new species of
Eptatretus and Quadratus (Myxinidae, Myxiniformes) from the
Caribbean coast of Colombia. Copeia 4:1026–1033
Opresko DM (2001) Revision of the Antipatharia (Cnidaria: Antho-
zoa). Part I. Establishment of a new family, Myriopathidae. Zool
Meded 75:147–174
zoa). Part II. Schizopathidae. Zool Meded 76:411–442
zoa). Part III. Cladopathidae. Zool Meded 77:495–536
zoa). Part IV. Establishment of a new family, Aphanipathidae.
Zool Meded 78:209–240
zoa). Part V. Establishment of a new family, Stylopathidae. Zool
Meded 80:109–138
Opresko DM, Sánchez JA (1997) A new species of antipatharian coral
(Cnidaria: Anthozoa) from the Caribbean coast of Colombia.
Caribb J Sci 33:75–81
Opresko DM, Sánchez JA (2005) Caribbean shallow-water black cor-
als (Cnidaria: Anthozoa: Antipatharia). Caribb J Sci 41:492–507
Pandolfi JM, Jackson JBC, Baron N, Bradbury RH, Guzman HM,
Hughes TP, Kappel CV, Micheli F, Ogden JC, Possingham HP,
Sala E (2005) Are U.S. coral reefs on the slippery slope to slime?
Science 307:1725–1726
Restrepo JD, Zapata P, Díaz JM, Garzón-Ferreira J, García CB (2006)
Fluvial fluxes into the Caribbean Sea and their impact on coastal
ecosystems: the Magdalena river, Colombia. Glob Planet Change
50:33–49
Reyes J (2000) Lista de los Corales (Cnidaria: Anthozoa: Scleractinia)
de Colombia. Biota Colomb 1:164–176
Reyes J, Santodomingo N (2002) Manual de identificación CITES de
invertebrados marinos de Colombia. Serie manuales de identifi-
cación CITES de Colombia. INVEMAR, Santa Marta
Reyes J, Santodomingo N, Gracia A, Borrero G, Navas G, Mejía-
Ladino L, Bermúdez A, Benavides M (2005) Southern Caribbean
azooxanthellate coral communities off Colombia. In: Freiwald A,
Roberts JM (eds) Cold-water corals and ecosystems. Springer,
Berlin, pp 309–330
Reyes J, Santodomingo N, Cairns S (2009) Caryophylliidae
(Scleractinia) from the Colombian Caribbean. Zootaxa 2262:1–39
Reyes J, Santodomingo N, Flórez P (2010) Corales escleractinios de
Colombia. INVEMAR Ser Publ Esp 14:1–246
Reyes-Bonilla H, Cruz-Piñón G (2000) Biogeography of the aherma-
typic corals (Scleractinia) of the Mexican Pacific. Cienc Mar
26:511–531
Roa-Varón A, Saavedra-Díaz L, Acero A, Mejía LS, Navas G (2003)
Nuevos registros de peces óseos para el Caribe colombiano de los
órdenes Beryciformes, Zeiformes, Perciformes y Tetraodonti-
formes. Bol Inst Inv Mar Cost 32:3–24
Roberts JM, Wheeler AJ, Freiwald A (2006) Reefs of the deep: the
biology and geology of cold-water coral ecosystems. Science
213:543–547
Roberts JM, Wheeler AJ, Freiwald A, Cairns S (2009) Cold-water
corals: The biology and geology of deep-sea coral habitats. Cam-
bridge University Press, Cambridge
Saavedra-Díaz L, Acero A, Navas A (2000) Lenguados de la familia
Paralichthyidae (Pisces: Pleuronectiformes) conocidos del Caribe
colombiano. Rev Acad Colomb Cienc Exact Fís Nat 24:295–310
Sánchez JA (1994) Presencia de los Octocorales Stylatula diadema
Bayer (Pennatulacea) y Carijoa riisei (Duchassaing y Michelotti)
/Telestacea) en la costa Caribe colombiana. An Inst Inv Mar Punta
Betín 23:137–147
Sánchez JA (1995) Benthic communities and geomorphology of the
Tesoro Island reefs, Colombian Caribbean. An Inst Inv Mar Punta
Betín 24:55–77
Sánchez JA (1998) Sistemática filogenética del género Eunicea Lamoroux,
1816 (Octocorallia: Gorgonacea: Plexauridae) con aspectos sobre la
historia natural de algunas especies en el Caribe colombiano. MSc
Thesis Universidad Nacional de Colombia, Bogotá
Sánchez JA (1999) Black coral-octocoral distribution patterns on
Imelda Bank, a deep-water reef, Colombia, Caribbean Sea. Bull
Mar Sci 65:215–225
Sánchez JA (2001) Systematics of the southwestern Caribbean Mur-
iceopsis Aurivillius (Cnidaria: Octocorallia), with the description
of a new species. Bull Biol Soc Wash 10:160–180
Sánchez JA, Wirshing H (2005) A field key to the identification of
tropical western Atlantic zooxanthellate octocorals (Octocorallia:
Cnidaria). Caribb J Sci 41:508–522
Sánchez JA, Díaz JM, Zea S (1997) Gorgonian communities in two
contrasting environments on oceanic atolls of the southwestern
Caribbean. Bull Mar Sci 61:453–465
Sánchez JA, Zea S, Díaz JM (1998) Patterns of octocoral and black
coral distribution in the oceanic barrier reef-complex of Providen-
cia Island, Southwestern Caribbean. Caribb J Sci 34:250–264
Santodomingo N, Reyes J, Gracia A, Martínez A, Ojeda G, García C
(2007) Azooxanthellate Madracis coral communities off San
Bernardo Islands (Colombian Caribbean). Bull Mar Sci 81(Suppl
1):273–287
Spalding MD, Fox HE, Allen GR, Davidson N, Ferdaña ZA, Finlayson
M, Halpern BS, Jorge MA, Lombana A, Lourie SA, Martin KD,
McManus E, Molnar J, Recchia CA, Robertson J (2007) Marine
ecoregions of the world: a bioregionalization of coastal and shelf
areas. Bioscience 57:573–583
Stewart R (2008) Introduction to physical oceanography. Department
of Oceanography, Texas A&M University
Stolarski J, Zibrowius H, Löser H (2001) Antiquity of the scleractinian-
sipunculan symbiosis. Acta Palaeontol Pol 46:309–330
Veron JEN (2000) Corals of the world. Australian Institute of Marine
Science, Townsville
Verseveldt J, Bayer FM (1988) Revision of the genera Bellonella,
Eleutherobia, Nidalia and Nidaliopsis (Octocorallia: Alcyoniidae
and Nidaliidae), with descriptions of two new genera. Zool Verh
245:1–131
Viaña, J, Medina A, Barrios M, Manjares L (2002) Evaluación de la
ictiofauna demersal extraída por la pesquería industrial de arrastre
en el área norte del Caribe colombiano (Enero 2000–Junio 2001).
Mar Biodiv (2013) 43:7–22 21

In: INPA-COLCIENCIAS. Informe técnico final Proyecto Eval-
uación de las pesquerías demersales del área norte del Caribe
colombiano y parámetros ecológicos, biológico-pesqueros y
poblacionales del recurso pargo (Código: 3135-09-550-98)
von Prahl H, Erhardt H (1985) Colombia, corales y Arrecifes coralinos.
FEN-Colombia, Editorial Presencia, Bogotá
von Prahl H, Erhardt H (1989) Lista anotada de corales ahermatípicos
de Colombia. Memorias del VI Seminario Nacional de Ciencias
del Mar, pp 539–556
Werding B, Sánchez H (1989) The coral formations and their distribu-
tional pattern along a wave exposure gradient in the area of Santa
Marta, Colombia. Medio Ambiente 10:61–68
Williams GC (1995) Living genera of sea pens (Coelenterata: Octocor-
allia: Pennatulacea): illustrated key and synopses. Zool J Linn Soc
Lond 113:93–140
Williams GC (1999) Index Pennatulacea: annotated bibliography and
indexes of the Sea Pens (Coelenterata: Octocorallia) of the World
1469–1999. Proc Calif Acad Sci 51:19–103
Zibrowius H (1980) Les Scléractiniaries de la Mediterranée et de
l’Atlantique nord-oriental. Mém Inst Océanogr Monaco 11:1–
248, 107 pls
Zibrowius H (1998) A new type of symbiosis: Heterocyathus japoni-
cus (Cnidaria: Scleractinia) living on Fissidentalium vernedei
(Mollusca: Scaphopoda). Zool Verh 323:319–334
22 Mar Biodiv (2013) 43:7–22

(2013) - Santodomingo - Corals colombia.pdf

Recommended

Recommended

More Related Content

Similar to (2013) - Santodomingo - Corals colombia.pdf

Similar to (2013) - Santodomingo - Corals colombia.pdf (20)

Recently uploaded

Recently uploaded (20)

(2013) - Santodomingo - Corals colombia.pdf