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Acropora cervicornis is one of the most important coral species in shallow reefs of the Caribbean as it provides habitat and structural complexity to several species of invertebrates and fish. However, the distribution range of A. cervicornis has shrunk and collapsed considerably in the last five decades, due to a combination of factors including the increase of disease prevalence, storm frequency, and anthropogenic threats. Despite being classed as “Critically Endangered” in the IUCN Red List, information regarding its population status and condition across large Caribbean coralline areas is limited. Herein we conducted the first Marine Protected Area (MPA) scale survey for this species at the Los Roques archipelago, which included visual census across 127 sites to determine the abundance, spatial distribution, habitat type, and patch morphology of A. cervicornis. We selected 11 sites, where this species was predicted and reported to be ubiquitous, to determine live A. cervicornis cover, its recent and old mortality cover, and white band disease prevalence as proxies for coral health. We found Acropora cervicornis in only 29% of the surveyed sites, with dispersed and scattered patches prevailing upon continuous patches. Moreover, the latter were located near the largest human population settlements, and inside the low protection zones of the MPA where fishing and touristic activities are permitted. The photomosaic survey showed that more than 75% A. cervicornis patches showed an average live cover above 27%, low prevalence of white band disease (<7%), and low macroalgal abundance (<10%); suggesting that Los Roques still holds healthy populations. Our results indicate that the persistence of this species urgently requires re-evaluating current MPA zoning, especially following recent evidence of overfishing and inadequate law enforcement. This study provides a baseline of A. cervicornis populations in Los Roques and Southern Caribbean that can be later used for local population management and conservation.
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INTRODUCTION
Coral reefs cover only a small portion of the tropical
ocean’s surface (0.1-0.5%), yet they provide habitat for
thousands of marine species, making these ecosystems one
of the most diverse on Earth (Moberg and Folke 1999;
Roberts et al. 2002; Mora et al. 2011; Fisher et al. 2015).
Scleractinian corals, especially members of the Acropori-
dae family, are foundational species in modern tropical
reefs since they are the major providers of structural com-
plexity (Bellwood et al. 2004; Idjadi and Edmunds 2006;
Wallace 2012; Raza et al. 2015). Their ability to adopt dif-
ferent growth morphologies through environmental gradi-
ents adds spatial heterogeneity to the reef substrate,
allowing many species to coexist (Pratchett et al. 2015).
Before the onset of their population’s collapse during the
late ’70s in the Caribbean, Acropora palmata (elkhorn
coral) and Acropora cervicornis (staghorn coral) formed
dense, monospecific and structurally complex patches that
contributed significantly to calcium carbonate accretion
along the fore reef of many Caribbean coral reefs (Aronson
and Precht 2001; Precht and Aronson 2004; Wapnick et al.
2004). These species also played a vital role in the mainte-
nance of healthy and productive reefs by providing critical
habitat and reef complexity for a large diversity of fish and
other organisms (Rogers et al. 1982; Gates and Ainsworth
2011). Moreover, compiling evidence shows that Caribbean
acroporids played these roles for thousands of years until
their populations were reduced during the last five decades
(Aronson and Precht 2001; Jackson 2001; Pandolfi and
Jackson 2006; Pandolfi and Jackson 2007).
Different studies have shown that the distribution range
of this species has shrunk considerably, with some cases
reporting more than 90% of area loss (Aronson and Precht
2001; Jackson et al. 2014; García Urueña et al. 2020), and
a lack of recovery since its regional decline (Vargas-Angel
et al. 2003; Keck et al. 2005; Busch et al. 2016). The un-
derlying causes of the regional collapse of A. palmata
and A. cervicornis populations have been firmly estab-
lished, and includes a combination of diseases, particularly
white band disease (WBD; Aronson and Precht 2001; Acro-
pora Biological Review Team 2005; Miller et al. 2014), in-
creased storm frequency (Woodley et al. 1981; Lirman and
Fong 1997), and the increase of anthropogenic threats such
as sediment load and overfishing (Bruckner 2002; Precht
et al. 2002; Greer et al. 2009).
Reduction in the populations of these two species led
to significant and unprecedented changes in the structure
and function of Caribbean coral reef ecosystems (Pandolfi
and Jackson 2006; Pandolfi and Jackson 2007). Increasing
ARTICLE
Distribution, abundance, and health indicators of the critically endangered coral
species Acropora cervicornis in Los Roques National Park, 2014
Stephanie J. Martinez1,2, Francoise Cavada-Blanco1,3,4, José Cappelletto5,6, Esteban Agudo-Adriani1,7, Aldo Cróquer1,8
1Experimental Ecology Laboratory, Department of Environmental Studies, University Simón Bolívar, Caracas, Venezuela; 2Department
of Freshwater and Marine Ecology, Institute for Biodiversity and Ecosystem Dynamics, University of Amsterdam, The Netherlands;
3Institute of Marine Science, School of Biological Sciences, University of Portsmouth, UK; 4EDGE of Existence Programme, Conser-
vation and Policy, Zoological Society of London, UK; 5I+D Group in Mechatronics, University Simón Bolívar, Caracas, Venezuela;
6Maritime Robotics Laboratory, Southampton Marine and Maritime Institute, University of Southampton, UK; 7Department of Biology,
The University of North Carolina, Chapel Hill, USA; 8Marine Innovation Center, The Nature Conservancy, Punta Cana, República
Dominicana.
ABSTRACT
Acropora cervicornis is one of the most important coral species in shallow reefs of the Caribbean as it provides habitat and
structural complexity to several species of invertebrates and fish. However, the distribution range of A. cervicornis has shrunk and
collapsed considerably in the last five decades, due to a combination of factors including the increase of disease prevalence, storm
frequency, and anthropogenic threats. Despite being classed as “Critically Endangered” in the IUCN Red List, information regarding
its population status and condition across large Caribbean coralline areas is limited. Herein we conducted the first Marine Protected
Area (MPA) scale survey for this species at the Los Roques archipelago, which included visual census across 127 sites to determine
the abundance, spatial distribution, habitat type, and patch morphology of A. cervicornis. We selected 11 sites, where this species
was predicted and reported to be ubiquitous, to determine live A. cervicornis cover, its recent and old mortality cover, and white
band disease prevalence as proxies for coral health. We found Acropora cervicornis in only 29% of the surveyed sites, with dispersed
and scattered patches prevailing upon continuous patches. Moreover, the latter were located near the largest human population set-
tlements, and inside the low protection zones of the MPA where fishing and touristic activities are permitted. The photomosaic
survey showed that more than 75% A. cervicornis patches showed an average live cover above 27%, low prevalence of white band
disease (<7%), and low macroalgal abundance (<10%); suggesting that Los Roques still holds healthy populations. Our results in-
dicate that the persistence of this species urgently requires re-evaluating current MPA zoning, especially following recent evidence
of overfishing and inadequate law enforcement. This study provides a baseline of A. cervicornis populations in Los Roques and
Southern Caribbean that can be later used for local population management and conservation.
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S.J. Martinez et al.
100
erosion and bioerosion rates (Edinger et al. 2000), species
replacement (Aronson et al. 1998), and loss of spatial het-
erogeneity and biodiversity are amongst the most dra-
matic effects reported in the literature (Bruckner 2002;
Acropora Biological Review Team 2005; Alvarez-Filip et
al. 2011). Because of the sudden decline of Acropora
palmata and A. cervicornis, combined with their current
critical status, both species were listed as threatened under
the United States Endangered Species Act, and classed as
Critically Endangered on the International Union for Con-
servation of Nature (IUCN) Red List of Threatened
Species (Aronson et al. 2008).
Acropora palmata and Acropora cervicornis are broad-
casting species, and both are known to be highly vulnerable
to natural and anthropogenic disturbances (Vargas-Ángel
et al. 2006; Schopmeyer et al. 2011; Miller et al. 2014;
Mercado-Molina et al. 2015). However, A. cervicornis is
capable of fast growth through asexual reproduction via
fragmented branches, and thus it has the potential of quick
recovery by forming monotypic patches/thickets within just
a few years (Bruckner 2002; Acropora Biological Review
Team 2005; Lucas and Weil 2016). Similar to Acropora
palmata, There are examples in a few locations where A.
cervicornis has indeed managed to persist after its regional
collapse (Vargas-Angel et al. 2003; Acropora Biological
Review Team 2005; Walker et al. 2012). However, the in-
creased frequency and intensity of natural and human dis-
turbances have decreased the survival rates and reduced the
probability of broken fragments attaching to suitable and
stable substrates, jeopardizing a good prognosis of recovery
(Lirman and Fong 1997; Goergen et al. 2019). Thus, de-
veloping standard restoration methods to help this species
come back has become a priority for conservation and local
legislations in the region (Schopmeyer et al. 2017).
To success in an effective population restoration, the
paucity of geographically extended demographic and eco-
logical data of Acropora cervicornis needs to be addressed
since it limits the planning for proper and coordinated
conservation actions (Bruckner 2002; Precht et al. 2002;
Mercado-Molina et al. 2015). Therefore, the identification
of locations where A. cervicornis populations of this
species still exist as shallow reef-builders, the character-
ization of these habitats, and the proper evaluation of po-
tential local threats are all critical to improve the impact
of local and regional conservation efforts (Bruckner et al.
2002; Aronson et al. 2008).
Within the Southern Caribbean, Archipelago Los
Roques National Park (“Los Roques”) has been highlighted
as one of the healthiest reef ecosystems due to its coral
cover remaining above the regional average (Villamizar et
al. 2003; Jackson et al. 2014; Debrot et al. 2019; Miyazawa
et al. 2020). Furthermore, different studies have identified
this location as a potential stronghold for Acropora palmata
(Zubillaga et al. 2008; Croquer et al. 2016). However, the
available reports on Acropora cervicornis for the MPA are
scarce and outdated, mostly collected during the mid-
’80s (Sandía and Medina 1987) and originated from studies
focused on characterizing benthic communities across the
archipelago rather than specifically assessing this
species (Villamizar et al. 2003; Weil 2003). This study
aimed to conduct the first systematic assessment on the sta-
tus of A. cervicornis at Los Roques, and to produce a base-
line on the species’ local distribution, abundance, and
health. Even though the data presented here does not rep-
resent the current status of the species, it enhances local and
regional knowledge while filling gaps about the spatial dis-
tribution of this critically endangered coral species.
METHODS
Study area
Archipelago Los Roques National Park is an oceanic
coral reef system located 170 km north of the Venezuelan
coast (REGVEN/UTM 19N 721011-7671071324721-
1297746; Figure 1). The reef system includes more than 50
coralline cays with fringing reefs, patch reefs, over 200
sandbanks, and extensive mangrove forests and seagrass
beds (Weil 2003; Croquer et al. 2016). The MPA zoning en-
compasses nine different use zones, including four coastal-
marine habitats, making Los Roques a multi-use
MPA (Croquer et al. 2016). The MPA zones range from high
protection (i.e., authorized scientific research or managed
non-extractive activities) to low protection (i.e., artisanal
fishing and recreational activities (Croquer et al. 2016;
Cavada-Blanco et al. 2021). According to this zoning,
human activities are mostly concentrated within the north-
east main island, (Gran Roque) and nearby cays (Figure 1).
Abundance, distribution, and habitat
To determine the distribution and abundance of Acro-
pora cervicornis in the MPA, visual censuses were con-
ducted between April and November 2014, encompassing
127 sites across the archipelago. These sites were selected
to cover the vast majority of potential and confirmed A.
cervicornis habitats within the MPA. Several criteria were
used during the selection, including (1) personal expertise
and knowledge of the MPA, (2) anecdotal information
gathered from local stakeholders (e.g., diving operators,
fishers, and homestay owners), and (3) observation of po-
tential habitats from raster satellite images. With these cri-
teria, the surveys included a suite of different habitats
including windward (exposed) and leeward (protected)
cays, fringing and barrier reefs, reef patches, mixed sea-
grasses, and sand habitats within the lagoon (Figure 1).
From this data, we produced a distribution map for A. cer-
vicornis at the scale of the entire MPA.
At each site, five observers conducted the visual sur-
veys through free-diving following the reef contour along
shallow to intermediate habitats (1–15 m depth). Twenty-
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Distribution of Acropora cervicornis in Los Roques 101
meter-wide belt-transects were surveyed ranging from 500
to 800 m in length following the procedures outlined in
(Croquer et al. 2016). At each belt-transect, the start and
endpoints were geo-referenced with a Garmin 60S GPS (lo-
cation accuracy within 3-15m) and the presence/absence
of Acropora cervicornis along each transect with basic de-
scriptive variables of their habitat (e.g., depth, level of wave
exposure) was recorded.
Colony density and ramification within patches made
colony differentiation too inaccurate to estimate abundance
through the direct count of discrete colonies. Consequently,
we used a qualitative approach to estimate abundance and
health status following the IUCN Red List of threatened
species guidelines for modular organisms (IUCN Standards
and Petitions Committee 2019). We classified A. cervicor-
nis patches into four different morphologies (Figure 2a-d):
(1) continuous patches or thickets (i.e., fields extending
over 100 m), (2) dispersed patches (i.e., fragmented patches
separated to each other by less than 2 m and extending for
10 m), (3) scattered patches (i.e., mixed patches composed
of isolated and mingled colonies), and (4) isolated colonies
(i.e., patches composed of individual colonies smaller than
2m wide, and at least 5 m apart from each other). Each mor-
phology type was then categorized according to their fre-
quency of occurrence as abundant (76-100%), common
(51-75%), uncommon (26-50%), and rare (0-25%). Patches
distribution and abundance were later mapped using QGIS
v3.16.4 (QGIS Development Team 2021).
Cover and health status
Using the distribution map of Acropora
cervicornis obtained from the previous section, a total of
11 sites were randomly selected to describe the benthic
community associated with these patches. In order to have
a representation of the benthic cover of these patches
across the entire archipelago, at least three sites were in-
cluded in major geographic sectors of the MPA: (1)
North-East (Madrizquí, Bajo de Medio 1 and Bajo del
Medio 2), (2) Central-East (Noronquí, La Venada, and
Rabusquí), and (3) Central-Southwest (Laguna de Espen-
quí, Isla Felipa, Isla Larga, Herradura Dos Mosquises, and
Los Canquises). On each site, four 25 m-long transects
were deployed systematically at depths that ranged be-
Figure 1. Archipelago Los Roques National Park (Los Roques) map showing survey locations and coastal-marine use zones.
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S.J. Martinez et al.
102
tween 2-10 meters. Along each transect, high-resolution
photos were taken ensuring an overlap of at least 30% to
produce 25 m2photomosaics (Figure 2e). The photomo-
saics were built using Hugin (D’Angelo 2010), following
the same methodology as in Agudo-Adriani et al. (2019)
in a process that entailed three main steps. First, the iden-
tification of matching control points between images to
estimate the relative position of an image in a sequence.
Secondly, photo alignment and optimization of movement
(tilt and balance), and position axes (x, y, z). Lastly, the
stitching and blending of images using a rectilinear pro-
jection together with brightness and colour exposure cor-
rections to produce a unique 25m-long photomosaic.
On each photomosaic, live cover of Acropora cervicor-
nis, other biotic (e.g., sponges, octocorals, macroalgae, other
coral species; see S.M.1), and abiotic substrates (e.g., dead
coral, sand, and coral rubble) were determined from 100
randomly overlayed points per mosaic using the software
CPCe (Kohler and Gill 2006). We used 100 points based on
species richness accumulation curves estimated from 20 ran-
dom transects (see S.M.2). Disease frequency was deter-
mined by counting the number of branches bearing signs of
white band disease (WBD) (Weil and Hooten 2008) in re-
lation to the total number of branches overlapping with the
100 points. The frequency of old (i.e., exposed skeleton cov-
ered by opportunistic organisms such as algae) and recent
mortality (i.e., bared coral skeletons) was also determined
(see S.M.3). The criteria for determining the previous vari-
Figure 2. Photographs showing the four types of Acropora cervicornis patches morphology: (a) Continuous patch, (b) dispersed patches, (c)
scattered patches, (d) isolated colonies. (d) Example of a reconstructed 25m long photomosaic, where the red line follows the transect line.
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Distribution of Acropora cervicornis in Los Roques 103
ables were based in the AGRRA protocol (Lang et al. 2010)
and field standardization.
Statistical analysis
We aimed at identifying the variables that better ex-
plained differences in live cover of Acropora cervicornis
across sites. For this, a distance-based linear model (Dis-
tLM; Legendre and Anderson 1999) was performed using A.
cervicornis live cover as the response variable, using the fol-
lowing as predictor variables: distance to the biggest per-
manent human settlement and least protected zone in the
MPA (Gran Roque Island), latitude and longitude coordi-
nates to control for spatial correlations, patch morphology,
habitat wave exposure (leeward, windward), and depth.
To test whether changes in the community structure as-
sociated with Acropora cervicornis patches and prevalence
of WBD varied across locations and sites, two-way Analy-
sis of Variance based on Permutations (PERMANOVA)
were done using the vegan package (Anderson 2005; Ok-
sanen et al. 2019). For these, similarity matrices were built
using Bray-Curtis dissimilarities and Euclidean distances
respectively, with geographic sector as a fixed factor (with
three levels: North-East, Central-East, and Central-South-
west), and sites as a random factor nested within sectors. A
SIMPER analysis was later performed to determine the
variables that contributed to the most dissimilarity between
geographical sectors and sites. Plots were made using the
package ggplot2 (Wickham 2009), implemented in R (R
Core Team 2020; RStudio Team 2020). Source code mate-
rial, and data matrices available at https://github.com/
Stephanocoenia/cervicornisLR.
RESULTS
Abundance, distribution, and habitat
Out of the 127 sites surveyed, we found Acropora cer-
vicornis in 37 (29%) sites; indicating that this species has
a narrow and very restricted distribution in Los Roques
(Figure 3a). The distribution of the four morphologies var-
ied across the archipelago, with dispersed patches being
the most frequent type (76% of sites were A. cervicornis
was found). Isolated colonies and scattered patches were
recorded in 70% and 43% of the sites, respectively. In
terms of abundance, dispersed and scattered patches were
common in most of the sites where these morphology
types occurred (32% and 22% of occurrence sites, respec-
tively; Figure 3a). Continuous and large patches were only
observed in two sites (La Venada and Madrizquí; 5% of
all surveyed sites); both within the least protection levels
of the MPA zoning (recreation area; Figure 1).
As for habitat features, we found the majority of A.
cervicornis patches were twice as frequent in leeward
reefs regardless of patch morphology (Figure 3b), and
more than 40% of the patches were frequent in shallow
depths (0-5 meters; Figure 3c). The only two continuous
patches were found in deeper reef zones
(Figure 3c). Combined, these results indicate that larger
patches of A. cervicornis are currently limited to a few
sites in Los Roques; either because this species always
had limited spatial distribution within the archipelago, or
because its populations have declined in the past.
The DistLM analysis (Table 1) showed that only two
(i.e., patch morphology and depth) out of the seven vari-
ables included in the model, significantly explained live
cover variations of Acropora cervicornis across sites.
However, these two variables combined explained 49%
of the variability in live coral cover recorded across sites.
Community structure within
Acropora cervicornis patches
The average live cover of Acropora cervicornis across
the surveyed (via photomosaics) sites was 26.9±14.2 %
(Figure 4), with values above this recorded at La Venada
(54.5±25.9%), Madrizquí (37.7±25.5%), Noronquí
(35.7±10.5%), Isla Felipa (34.7±10.1%), Isla Larga
(34.7±5.9%) and Bajo del Medio 1 (32.5±9.1%). Seven
scleractinian species were also part of the coral commu-
nity, but their live cover never exceeded 3% (i.e., Orbi-
cella annularis, Orbicella faveolata, Siderastrea siderea,
Eusmilia fastigiata, Diploria strigosa; Figure 5). Other
Table 1. Distance-based multivariate analysis for a linear model (DistLM) using Acropora cervicornis cover as the response variable.
Predictor variables were site coordinates (latitude and longitude), distance of each location to Gran Roque, thicket morphology, habitat
wave exposure, depth, and nearest neighbour relation. * Indicates p-values <0.05
Variable Sum Squares Pseudo-F p-value Prop
Longitude 490.730 1.3935 0.244 0.0321
Latitude 360.430 1.0146 0.325 0.0235
Distance to GR 359.970 1.0133 0.340 0.0235
Thicket morphology 5166.000 21.5400 0.001* 0.3380
Habitat 79.401 0.2194 0.622 0.0519
Depth 2288.900 7.3994 0.008* 0.1497
Nearest neighbour 600.210 1.7171 1.171 0.0393
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S.J. Martinez et al.
104
organisms such as octocorals (e.g., Briareum as-
bestinum, Plexaura homomalla, and Pseudoplexaura
porosa), hydrocorals (i.e., Millepora complanata), and
sponges were ubiquitously found in these patches and oc-
cupying less than 2% of the substrate (Figure 4). The
algae community was dominated by macroalgae, but
never exceeded 10% of cover (Figure 4). Highest cover
for abiotic substrates corresponded to sand (6.4±2.7-
Figure 3. Acropora cervicornis patches abundance and distribution map in Los Roques (a), frequency of patches according to habitat
exposure (b), and depth profiles (c).
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Distribution of Acropora cervicornis in Los Roques 105
52.3±16.1 %, Figure 4) and coral rubble (8.1±3.7-
32.7±8.6%, Figure 4). The benthic community of Acrop-
ora cervicornis patches significantly varied at the scale
of sites within sectors of the MPA, encompassing 52% of
the total variation (Table 2). To a lesser extent, the geo-
graphical sector explained only 13% of this benthic vari-
ation (Table 2). Where 70% of this variability was due to
differences in the cover of live A. cervicornis, macroalgae,
Pseudoplexaura porosa tissue, as well as old coral mor-
tality, sand, and rubble.
Health status
Dead branches were consistently found at every sur-
veyed patch; however, old mortality was on average 10-
Table 2. PERMANOVA based using a Bray-Curtis dissimilarity matrix of benthic substrates across two factors: geographical sectors
and sites. * Indicates p-values <0.05
Variation source df Sum of Squares Mean Squares F-Model p-value R2
Sector 2 0.51 0.26 5.96 0.001* 0.13
Sector: site 8 2.07 0.26 6.02 0.001* 0.52
Residuals 33 1.42 0.04 0.35
Total 43 4.01 1.00
Figure 4. Average cover of biotic benthic community groups and abiotic substrates associated to Acropora cervicornis patches per lo-
cation, and geographic sectors. Sponges, octocoral and scleractinian species were grouped into major categories. HDMS: Herradura
Dos Mosquises, IF: Isla Felipa, CAN: Los Canquises, IL: Isla Larga, ESP: Laguna de Espenquí, NOR: Noronquí, VEN: La Venada,
RAB: Rabusquí, MAD: Madrizquí, BM1: Bajo de Medio 1, BM2: Bajo del Medio 2.
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S.J. Martinez et al.
106
fold higher compared to the frequency of recent mortality
which seldom exceeded 2% across all sites (Figure 5a).
The highest average of old mortality was recorded at the
largest and more continuous patches, ranging from
27.7±7.9 in La Venada to 26.8±27.1 in Madrisquí. In Los
Canquises, the farthest site from any recreational activity
and human settlement point in the MPA, partial mortality
was 12-fold lower compared to the previous sites (Fig-
ure 5a). Moreover, only two sites (Bajo del Medio 1, Bajo
del Medio) presented higher mortality than live tissue.
WBD prevalence ranged between 0.1% and 22%, signif-
icantly varying at the scale of sites and explaining 43%
of the total variation among sites (Figure 5b, Table 3).
Furthermore, prevalence above the average was only
recorded at Boca del Medio 2 (19.0±16.0%), La Venada
(17.5±11.9%), and Espenquí (12.2±11.3 %; Figure 5b).
Our results indicate that surveyed patches in the MPA
showed a live cover of A. cervicornis above 10%, with low
macroalgal abundance (<5%), predominance of old mortality
(≤25%), and a WBD prevalence that never exceeded 10%.
DISCUSSION
This study represents the first systematic and compre-
hensive survey of Acropora cervicornis populations across
Los Roques. Prior to our study, all available information was
Figure 5. Health status of Acropora cervicornis in Los Roques. Live cover tissue versus old and recent mortality (a), and white band
disease (WBD) prevalence (b) per location. HDMS: Herradura Dos Mosquises, IF: Isla Felipa, CAN: Los Canquises, IL: Isla Larga,
ESP: Laguna de Espenquí, NOR: Noronquí, VEN: La Venada, RAB: Rabusquí, MAD: Madrizquí, BM1: Bajo del Medio 1, BM2: Bajo
del Medio 2.
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Distribution of Acropora cervicornis in Los Roques 107
limited to a single site (Sandía and Medina 1987) or aimed
to describe the benthic community structure at three points
in time, as opposed to assessing the status of A. cervicornis
(Villamizar et al. 2003; Weil 2003; Miyazawa et al. 2020).
We found that A. cervicornis has a restricted distribution in
Los Roques, where it was typically located on shallow and
protected reefs, and mostly forming scatter and/or dispersed
patches. Only two continuous and abundant patches of A.
cervicornis were spotted in two surveyed sites located
within the low protection zones of the MPA. Moreover, the
prevalence of WBD and recent mortality was low (<10%)
across the survey sites, and the cover of recently dead tissue
did not surpass old mortality and live tissue.
The continuous Acropora cervicornis patches showing
the highest live cover recorded within the MPA were only
found in La Venada and Madrizquí (53.3±24.4 and
50.2%±3.1% respectively), both locations inside the lowest
level protection zones. These islands are close to the biggest
and most heavily populated island of Gran Roque Island,
where various tourism and diving activities are permitted.
In addition, the proximity to Gran Roque Island also in-
creases the proximity to anthropogenic stressors such as
pollution and sewage discharges (Croquer et al. 2016).
Within the MPA, Acropora cervicornis occupies habi-
tats with highly specific features, consisting of leeward
sandy bottoms protected from strong wave energy. The
presence of sand as dominant substrate contrast with pre-
vious habitat descriptions from other parts of the species
distribution range in the Florida Keys (Miller et al. 2008;
D’Antonio et al. 2016) and other U.S. Caribbean territories
(Wirt et al. 2015) where they report that this species occurs
mostly on consolidated hardbottom, and rubble zones.
However, the presence of most patches in leeward reefs is
consistent with previous studies (Vargas-Angel et al. 2003;
D’Antonio et al. 2016; Weil et al. 2020). Furthermore, we
found that A. cervicornis cover in Los Roques was highly
related to patch morphology and depth, where the species
presence was seldom observed deeper than 5 m. This is
consistent with previous findings indicating that A. cervi-
cornis occurred at depths ranging from 15 to 25 m before
their population collapsed, and now the species is regularly
found in shallower habitats ranging from 5 to 14 m (Var-
gas-Angel et al. 2003; Miller et al. 2008). Even though pre-
vious population data of the species in the MPA is limited
for comparison, the low occupancy, predominance of scat-
ter patches, and in the occurrence in shallow areas could
suggest that A. cervicornis has reduced its distribution habi-
tats across the MPA. This evidence should prompt decision-
makers to design specific actions to restore and/or to foster
population recovery of this species in specific areas within
the MPA. Such efforts are currently absent and have never
been attempted in Los Roques.
The health status of Acropora cervicornis suggests
that factors producing mortality in the archipelago are
widespread across the MPA. The prevalence of WBD was
commonly observed across the MPA distribution of the
species, ranging from 0% to 33.3% (mean 6.8%±9.2% ),
which was similar to other reports across other large and
continuous patches in the Caribbean (Lirman et al. 2010;
Miller et al. 2014; Goergen et al. 2019). Also, the contin-
uous patch in La Venada presented the second highest
prevalence of WBD (mean 17.5%±11.9%) across survey
sites. It has been previously observed that high-density
patches are more susceptible to predation and diseases are
less likely to be to persist through modern disturbances
and conditions (Goergen et al. 2019). Therefore, high-
lighting the increased vulnerability of one of the biggest
patches of A. cervicornis in the MPA.
Different studies have shown that Los Roques harbours
healthy populations of other key and vulnerable scleractin-
ian species such as Acropora palmata and Dendrogyra
cylindrus (Zubillaga et al. 2005; Croquer et al. 2016;
Cavada-Blanco et al. 2020; Cavada-Blanco et al. 2021).
Most likely because the archipelago remains outside the
main impact route of hurricanes and has not been subjected
to coastal development (Zubillaga et al. 2005; Croquer et
al. 2016). However, it is imperative to maintain constant
surveillance across the whole MPA because A. cervicornis
is known to be highly vulnerable to natural and anthro-
pogenic disturbances such as diseases (Gladfelter 1982;
Aronson and Precht 2001; Verde et al. 2016), recurrent epi-
zootic events (Knowlton 1992; Williams and Miller 2005;
Miller et al. 2014; Goergen et al. 2019), increase of storms
frequency and habitat degradation (Hernández-Delgado et
al. 2014; Goergen et al. 2019), and episodes of high thermal
stress that can lead to loss of disease resistance (Quinn and
Kojis 2008; Muller et al. 2018). Even in cases where the
population’s dynamic seems stable and without distur-
bances, the growth rate of patches is below equilibrium
(Mercado-Molina et al. 2015).
Table 3. Univariate PERMANOVA using a Euclidean dissimilarity matrix of white band disease frequency. * Indicates p-values <0.05
Variation source df Sum of Squares Mean Squares F-Model p-value R2
Sector 2 204.14 102.07 1.80 0.216 0.06
Sector: site 8 1548.06 193.51 3.41 0.015* 0.43
Residuals 33 1874.41 56.80 0.52
Total 43 3626.61 1.00
Non-commercial use only
S.J. Martinez et al.
108
The survey shows that Los Roques holds one of the
few healthy and large Acropora cervicornis populations
in the Southern Caribbean. A. cervicornis patches live
cover and disease prevalence comparable to other contin-
uous patches found in the region such as Fort Lauderdale-
Florida (Vargas-Angel et al. 2003; Williams et al. 2008;
Walker et al. 2012), La Parguera-Puerto Rico (Lucas and
Weil 2016; Weil et al. 2020), Coral Gardens-Belize
(Busch et al. 2016), Punta Rusia-Dominican Republic
(Lirman et al. 2010), and Roatan-Honduras (Keck et al.
2005). This shows that even after the critical mass bleach-
ing events of 2005 and 2010 (Villamizar et al. 2008;
Bastidas et al. 2012), A. cervicornis populations were still
found thriving in shallow and vulnerable habitats affected
by sudden increases in water temperature. Thus, the re-
sults presented in this study show that there are still po-
tential refugia for this species in Los Roques.
Multiple studies have shown that there are phylogeo-
graphical barriers (e.g., Mona Passage, Mesoamerican
Barrier Reef) across the extent of the wider Caribbean for
multiple coral species (Baums et al. 2005; Galindo et al.
2006; Vollmer and Palumbi 2007; Zubillaga et al. 2008;
Baums et al. 2010; Foster et al. 2012; Rippe et al. 2017).
Moreover, larvae dispersal and gene flow of Acropora
cervicornis is limited when distances exceed 500 km
(Vollmer and Palumbi 2007). In this sense, the popula-
tions in the MPA might represent reservoirs of genetic
variation for the Southern Caribbean, where sexually pro-
duced larvae can be used to assist the restoration of pop-
ulations that are not thriving well in the neighbouring
areas. However, it is important to first assess the propor-
tion of clones, reproductive success, and the population’s
genetic variability to further understand the population’s
dynamics within the MPA. Therefore, with appropriate
local management, conservation, and protection of local
source populations we can safeguard the future of the
species in the area (Vollmer and Palumbi 2007; Weil et
al. 2020).
Overall local and anthropogenic threats might be min-
imized due to limited access to the area, its MPA status,
and low human populations. However, the latest evidence
of the MPA’s degradation due to parrotfish and other her-
bivores overfishing, changes in governance, and inade-
quate surveillance and law enforcement poses an
imminent threat (Croquer et al. 2016; Agudo-Adriani et
al. 2019; Cavada-Blanco et al. 2020; Cavada-Blanco et
al. 2021). Thus, we suggest the urgent need to revise the
MPA´s zoning and regulations, established in 1991, to
protect these dense and extensive patches of Acropora
cervicornis. Finally, information presented in this paper
could be used to plan future restoration plans for this
species based on sexual and/or asexual propagation as
other Caribbean countries are currently implementing
(Bayraktarov et al. 2020; Sellares-Blasco et al. 2021).
CONCLUSIONS
This paper represents the first baseline study showing
the distribution and status of Acropora cervicornis in Los
Roques. We showed this species is restricted to a limited
number of sites within the MPA with dispersed and scat-
tered patches prevailing upon continuous patches. How-
ever, most patches of A. cervicornis showed average live
cover >30%, low prevalence of WBD, and macroalgal
abundance, further suggesting that Los Roques represents
a stronghold for staghorn corals as it still holds healthy
populations. Nonetheless, the distribution of this species
clearly overlaps with areas with low protection levels
where fishing and tourism activities occur, and the largest
human population is settled. This, along with an outdated
MPA zonation, may severely hamper the persistence of A.
cervicornis in Los Roques in the future.
ACKNOWLEDGEMENTS
This project was partially funded by an EDGE of Exis-
tence Fellowship awarded to Francoise Cavada-Blanco in
2013, and by the Laboratorio de Ecología Experimental
(Universidad Simón Bolívar) through in-kind contributions.
Field studies were authorized by the Ministerio del Poder
Popular de Ecosocialismo, Hábitat y Vivienda (Approval
number: 0323), and Territorio Insular Miranda (Approval
number: 006). We would like to acknowledge our boat
driver, fishers, dive operators and lodges for their time, and
for providing information about the species in Los Roques.
We also thank Luis M. Montilla for helping with data
analysis, and Maria Zalm for proofreading the manuscript.
Corresponding author: Stephanie J. Martinez.
E-mail: jmartinez.sh@gmail.com
Authors’ contributions: All the authors made a substantive intel-
lectual contribution, performed part of the experiments. All the au-
thors have read and approved the final version of the manuscript
and agreed to be accountable for all aspects of the work.
Availability of data and materials: All data generated or analyzed
during this study are included in this published article.
Conflict of interest: The authors declare no potential conflict of in-
terest.
Keywords: Acropora cervicornis, MPA, Los Roques, staghorn
coral.
Received: 5 December 2021.
Accepted: 29 December 2021.
This work is licensed under a Creative Commons Attribution Non-
Commercial 4.0 License (CC BY-NC 4.0).
©Copyright: the Author(s), 2021
Licensee PAGEPress, Italy
Advances in Oceanography and Limnology, 2021; 12:10005
DOI: 10.4081/aiol.2021.10005
Non-commercial use only
Distribution of Acropora cervicornis in Los Roques 109
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