Unraveling a resilient reef: structure and composition of

Varadero, an imperiled coral reef in the Colombian Caribbean

Valeria Pizarro Corresp., 1 , Sara C Rodríguez 2 , Mateo López-Victoria 2 , Fernando A Zapata 3
, Sven Zea 4
, Claudia
T Galindo-Martínez 5 , Roberto Iglesias-Prieto 5 , Joseph Pollock 5 , Monica Medina 5

1
Ecomares NGO, Cali, Valle, Colombia
2
Department of Natural Sciences and Mathematics, Pontifica Universidad Javeriana, Cali, Valle, Colombia
3
Department of Biology, Universidad del Valle, Cali, Valle, Colombia
4
Centro de Estudios en Ciencias del Mar - CECIMAR, Universidad Nacional de Colombia - Sede Caribe, Santa Marta, Magdalena, Colombia
5
Department of Biology, Pennsylvania State University, State College, Pennsylvania, United States

Corresponding Author: Valeria Pizarro

Email address: [email protected]

Coral reefs supply millions of people with ecosystem goods and services, especially those
living along tropical coastlines. Unfortunately, these ecosystems are disappearing at an
alarming pace. In the Caribbean, the rate of coral loss is high (5.5 – 9.2% per year) and
constant. In 2013, a healthy coral reef was discovered in one of the least expected places
within the Colombian Caribbean: at the entrance of Cartagena Bay, a highly-polluted
system that receives industrial and sewage waste, as well as high sediment and
freshwater loads from an outlet of the Magdalena River (the longest and most populated
river basin in Colombia). Here we provide the first characterization of Varadero Reef’s
geomorphology and biological diversity. We also compare these characteristics with those
of a nearby reference reef, Barú Reef, located in an area much less influenced by the
described polluted system. Below the murky waters, we found high coral cover of 45.1%
(± 3.9; up to 80% in some sectors), three species of lobster, eight of sea urchin, a fish
community composed by 61 species from 24 families, and the typical zonation of a
Caribbean fringing reef. All attributes found correspond to a reef that, according to current
standards should be considered in "good condition". Current plans to dredge part of
Varadero threaten the survival of this reef and could hinder efforts to uncover the
underpinnings of this reef’s remarkable resilience. There is, therefore, an urgent need to
describe the location and characteristics of Varadero as a first step towards gaining
acknowledgement of its existence and garnering inherent legal and environmental
protections.

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 1 Unraveling a resilient reef: structure and composition of
2 Varadero, an imperiled coral reef in the Colombian
3 Caribbean
4 Valeria Pizarro1, Sara Catalina Rodríguez2, Mateo López-Victoria2, Fernando A. Zapata3, Sven
5 Zea4, Claudia Tatiana Galindo-Martínez5, Roberto Iglesias-Prieto5, F. Joseph Pollock5, Mónica
6 Medina5
7 1 Fundación Ecomares, Cali, Valle, Colombia
8 2 Department of Natural Sciences and Mathematics, Pontificia Universidad
9 Javeriana Seccional Cali, Valle, Colombia
10 3 Department of Biology, Universidad del Valle, Cali, Valle, Colombia

11 4 Departament of Biology, Universidad Nacional de Colombia, Santa Marta, Magdalena,

12 Colombia, [email protected]
13 5 Department of Biology, Pennsylvania State University, State College, Pennsylvania, USA

14 Corresponding Author:

15 Valeria Pizarro1
16
17 Calle 5B # 4-139, Santa Marta, Magdalena, Colombia
18
19 Email address: [email protected]

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20 Abstract

21 Coral reefs supply millions of people with ecosystem goods and services, especially those living
22 along tropical coastlines. Unfortunately, these ecosystems are disappearing at an alarming pace.
23 In the Caribbean, the rate of coral loss is high (5.5 – 9.2% per year) and constant. In 2013, a
24 healthy coral reef was discovered in one of the least expected places within the Colombian
25 Caribbean: at the entrance of Cartagena Bay, a highly-polluted system that receives industrial
26 and sewage waste, as well as high sediment and freshwater loads from an outlet of the
27 Magdalena River (the longest and most populated river basin in Colombia). Here we provide the
28 first characterization of Varadero Reef’s geomorphology and biological diversity. We also
29 compare these characteristics with those of a nearby reference reef, Barú Reef, located in an area
30 much less influenced by the described polluted system. Below the murky waters, we found high
31 coral cover of 45.1% (± 3.9; up to 80% in some sectors), three species of lobster, eight of sea
32 urchin, a fish community composed by 61 species from 24 families, and the typical zonation of a
33 Caribbean fringing reef. All attributes found correspond to a reef that, according to current
34 standards should be considered in "good condition". Current plans to dredge part of Varadero
35 threaten the survival of this reef and could hinder efforts to uncover the underpinnings of this
36 reef’s remarkable resilience. There is, therefore, an urgent need to describe the location and
37 characteristics of Varadero as a first step towards gaining acknowledgement of its existence and
38 garnering inherent legal and environmental protections.

39 Introduction
40 Coral reefs provide important ecosystem services (Moberg & Folke, 1999), but many currently
41 face unprecedented pressure from multiple natural and anthropogenic factors (Wilkinson, 2008).
42 Caribbean reefs have been particularly impacted, with coral cover decreasing from an average of
43 50% to 10% in just four decades (Jackson et al., 2014). Coral cover loss has resulted in a phase
44 shift from coral to macroalgal domination with a concurrent increase in sponge abundance (e.g.,
45 Rose and Risk, 1985; Szmant, 2002; Ward-Paige et al., 2005; Chaves-Fonnegra et al., 2007;
46 Maliao et al., 2008; Jackson et al., 2014).

47 Coral reef ecosystems, built mainly by scleractinian corals, typically thrive within a narrow
48 range of environmental conditions characterized by low sedimentation rates, low nutrient

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49 availability (i.e., oligotrophic waters), high light penetration, warm waters (e.g., around 28 oC)
50 and salinity between 33 and 36 psu (Kleypas, McManus & Meñez, 1999; Díaz et al., 2000;
51 Sheppard, Davy & Pilling, 2009). Although reefs can be found outside these ranges in “extreme”
52 environmental conditions, such reefs are typically dominated by a low number of resistant
53 specialist species. Some examples include reefs under higher water temperatures in the Persian
54 Gulf and Hawaii (Oliver & Palumbi, 2009; Riegl & Purkis, 2012), reefs under low pH waters in
55 Japan and Papua New Guinea (Fabricius et al., 2011; Inoue et al., 2013), and reefs under high
56 salinity such as those at the Arabian Sea were salinity can exceed 45 psu and temperatures
57 regularly top 34 oC (Rezai et al., 2004).

58 In 2013, a reef was discovered under unexpected conditions below a thick layer of highly turbid
59 water at the mouth of Cartagena Bay, Colombia (López-Victoria et al., 2015). This reef, known
60 as Varadero, is located south of Tierra Bomba Island, at the mouth of the highly-polluted Bay.
61 The man-made “Canal del Dique” dumps industrial and sewage waste as well as discharges of
62 sediment from the Magdalena River into the vicinity of Varadero. With a drainage basin
63 covering 24% of Colombia’s surface area (27.3 million hectares), the Magdalena River feeds
64 approximately 144 x 106 tons of suspended solids into Cartagena Bay each year. This enormous
65 sediment load has contributed to the demise of the Bay’s once vibrant coral reefs (Restrepo et al.,
66 2006). Paradoxically, Varadero Reef has not only survived, but thrived with up to 80% coral
67 cover dominated by large Orbicella spp. colonies, the major reef building corals in the Caribbean
68 (López-Victoria et al. 2015).

69 Despite its close proximity to the city of Cartagena, Colombia (> 1 million inhabitants),
70 Varadero Reef remained concealed due to perception that local environmental conditions were
71 incompatible with reef growth. High levels of sedimentation and turbidity have previously been
72 shown to drive coral bleaching and disease that can ultimately lead to coral death (Bruno et al.,
73 2003; Harvell et al., 2007; Pollock et al., 2014). Here we provide a preliminary characterization
74 of Varadero Reef, including its geomorphology (i.e., size, shape and location) and biological
75 diversity (i.e., coral, fish and sponge community composition). We also compare these
76 characteristics with those of a nearby reference reef, Barú Reef, located 4.5 km south of
77 Varadero, in a location much less influenced by runoff from the Canal del Dique and the city of
78 Cartagena.

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 79 Current plans to dredge part of Varadero threaten the survival of this reef and could hinder
80 researchers’ ability to gain insights into the factors that have allowed corals to thrive under such
81 unusual conditions. There is, therefore, an urgent need to describe the location and characteristics
82 of Varadero as a first step towards gaining acknowledgement of its existence and garnering
83 inherent legal and environmental protections.

84 Materials & Methods

85 In order to supplement the brief, general description of Varadero Reef reported by López-
86 Victoria et al. (2015), detailed geomorphological and biological surveys were performed
87 between 2014 (March) and 2015 (March and October). During the March 2015 field trip, Reef’s
88 geographic extent was assessed by two researchers diving along the border of the Reef with a
89 GPS, recording in tracking mode, attached to an accompanying buoy. This information was
90 downloaded using GIS software (Garmin BaseCamp). Subsequently a map of the Reef was
91 produced. The Reef’s coral diversity was characterized by two coral experts performing three
92 replicate profiles starting in the deepest zone (in direction to open sea) towards Cartagena Bay
93 (shallowest zone). The methods of Geister (1977) were employed, which include annotations of
94 coral community composition at multiple depths. All profiles were compared and compiled to
95 obtain a detailed profile of the Reef’s coral community structure and diversity.

96 The vertical attenuation coefficients (Kd) were determined at both sites using the cosine corrected
97 sensor of a diving pulse modulated fluorometer (PAM) (Waltz, Germany). The PAM sensor was
98 calibrated against a traceable reference sensor LiCor (USA). A diver operating the PAM
99 maintained the instrument in a horizontal position and triggering the data collection system of the
100 fluorometer at different depths. The maximum excitation pressure over photosystem II (Qm) was
101 calculated in both sites using the effective quantum yield of photosystem II at apparent noon
102 (F/Fm’) and the maximum quantum yield of charge separation at dusk (Fv/Fm) (Iglesias-Prieto
103 et al., 2004).

104 A detailed benthic community assessment was also conducted to evaluate sessile and mobile
105 species composition, fish diversity and abundance, and sponge richness. To allow comparison of
106 Varadero with a nearby reef that reflected typical Caribbean reef environmental conditions, a
107 reef on the Barú Peninsular (from now on Barú Reef) was also surveyed. At each reef, five

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108 stations were established and two 30 m transects were deployed in the same landscape unit (i.e.,
109 reef type and depth). Quadrats (50 by 50 cm) were placed every three meters on each side of the
110 transect and photographed for a total of 20 photo-quadrats per transect. Each photograph was
111 analyzed using Coral Point Count 4.1 software (Kohler & Gill, 2006), which randomly places 50
112 points within the quadrat for a randomly stratified methodology (Kohler & Gill, 2006; Dumas et
113 al., 2009; Andersen et al., 2012). The benthic component below each point was identified and
114 categorized as coral (identified to coral species), sponge (identified to sponge species), algal
115 overgrown dead coral, sand/rubble or other invertebrates (e.g., tunicates, gorgonians or
116 zoanthids). Mobile reef invertebrates were also assessed using the same benthic transects. A
117 visual census was preformed of all sea urchins, conchs, and lobsters within a 1 m band of the
118 transect. Macroalgal communities were characterized by randomly selecting five photoquadrats
119 per transect, randomly placing 10 points within each quadrat (using Coral Point Count 4.1), and
120 categorizing any observed macroalgae as fleshy, coralline or turf. Fleshy algae were identified to
121 genus level. To compare Varadero and Barú Reefs, species richness, abundance and composition
122 were tested for normal distributions (Shapiro-Wilk's test) then compared using a two sample
123 Student’s t-test in the software PAST version 3.14 (Hammer, Harper, & Ryan, 2001).

124 During exploratory dives, sponges were visually identified while swimming over the reef.
125 Photographs and small samples were also taken for later spicule examination in cases when
126 sponges could not be readily distinguished in the field. Species lists were made for both
127 Varadero and Barú Reefs, separately for the upper terrace (down to 10 – 13 m) and slope (below
128 10 – 13 m) zones. Sponge species present within each of the 2 x 30 m band transects in the
129 shallow terrace zone of Varadero (n = 7 transects) and Barú (n = 4 transects) were also recorded.
130 This sampling scheme permitted calculation of gross abundances as percent frequency of
131 occurrence (number of transects in which a sponge was present/total transects) and species
132 richness per transect. Data on total coral and sponge cover obtained in 10 phototransects
133 (covering 5 m2 each, see above) in the upper terrace of each locality were also analyzed for
134 trends in cover of sponges vs. corals vs. available substratum using simple correlation analysis.
135 For sponge identification in the laboratory, small fragments of each collected sponge were
136 digested in commercial bleach to obtain free spicules, which were observed under a light
137 microscope. Species were identified using specialized literature and extensive local
138 knowledge/experience (see Zea, 1987; Zea et al., 2014).

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139 Overall fish diversity and community composition were visually assessed. In order to compile
140 fish species lists for each reef, a team of three divers recorded all fishes observed while exploring
141 the general reef areas of Varadero and Barú during a total of 8 dives on each reef (approximately
142 1-hour per dive), in 2014 and 2015. In 2015, 22 visual censuses were performed along 30 x 2 m
143 belt-transects (n = 15 at Varadero and n = 7 at Barú) to characterize fish community
144 composition. All individuals observed within each belt transect were counted and these counts
145 were used to estimate mean species richness, diversity (Shannon’s H’), dominance (Simpson’s
146 D) and evenness (Pielou’s J’). These community variables were compared between Varadero and
147 Barú using a two sample Student’s t-test, after establishing that the data met assumptions of
148 normality and homoscedasticity with Shapiro-Wilk’s and F tests, respectively. All tests were
149 performed using PAST 3.14 (Hammer, Harper & Ryan, 2001).

150 To assess species abundance differences between sites, a regression analysis of mean species
151 abundance was performed along with paired Student’s t-tests. Given the different sampling
152 efforts between the two localities, a sample-based rarefaction procedure was carried out to
153 compare fish species richness between Varadero and Barú. Finally, a non-Metric
154 Multidimensional Scaling (nMDS) analysis was carried out using Jaccard’s similarity index
155 (based on species occurrence) and the Bray-Curtis similarity index [based on the log (x + 1)
156 transformed abundance data] to examine differences in assemblage structure between the two
157 localities based on species composition and abundance, respectively. The nMDS analysis was
158 complemented with analyses of similarity (ANOSIM) based on either Jaccard or Bray-Curtis
159 similarity. All statistical analyses and calculation of community indices were performed using
160 the software PAST 3.11 (Hammer, Harper & Ryan, 2001).

161 Results
162 Geomorphology and optical properties

163 Located between the Bocachica navigation channel and the island of Barú, Varadero Reef has an
164 area of approximately 1.12 km2 (Figure 1). The Reef has two contrasting zones, the first (0.44
165 km2) is a well-developed reef where scleractinian coral colonies dominate the substratum. The
166 second (0.68 km2) is a carbonated terrace with scattered corals, octocorals, a few other benthic
167 species and sand patches with seagrasses (Figure 1c, Figure 2). The largest seagrass beds were

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168 observed near the islands of Draga and Abanico (Figure 1c). Analyses of the vertical attenuation
169 coefficients of the water in both sites indicate significant vertical stratification. We identify an
170 upper layer with high attenuation values located between the surface and 3-5 m depth.
171 Comparisons between the attenuation coefficients of the first layer at both sampling sites indicate
172 significantly (p<0.001 ANOVA) higher attenuation values for Varadero Reef (0.336 ± 0.050 m-1,
173 average ± SE, n = 32) relative to Barú Reef (0.243 ± 0.053 m-1, n = 11). In some cases, we
174 identify a second layer with Kd values ranging between 0.193 and 0.051 m-1 at depths above the
175 limit of the first layer between three to five meters (Figure 3). Depending on the depth profile of
176 the reef, some corals were completely contained within the first optical layer (Figures 2-3). We
177 recorded the maximum excitation pressure of photosystem II for Orbicella faveolata colonies
178 growing in the shallow parts of both reefs. In both cases corals were exposed to irradiances high
179 enough to induce significant levels of photoprotection at noon with Qm values of 0.208 ± 0.109,
180 average ± SE, n = 25 at 4.5 m depth and 0.249 ± 0.052, n = 25 at 6.0 m depth for Varadero and
181 Barú Reefs respectively.

182 Coral and benthic community

183 In total, 42 scleractinian coral and four fire coral species (Families Milleporidae and
184 Stylasteridae) were identified at Varadero (Table S1). These species include several threatened
185 species such as the acroporids (Acropora cervicornis and A. palmata). Depth profiles indicate
186 that Varadero Reef's calcareous matrix starts at around 27 to 35 m depth (Figure 2). At greater
187 depths, moving towards open sea, the sand bottom has small patches of sponges and black corals
188 (Anthipatharia). Coral cover from 27 to 35 m until approximately 10 to 12 m is relatively low (1
189 to 5%) and the reef slope is around 45o. Coral communities at this depth range are dominated by
190 Agaricia spp. (A. lamarcki, A. grahamae), Madracis spp. and Helioceris cucullata. At 25 m and
191 shallower, small plate-like growth forms of Siderastrea siderea, Montastraea cavernosa and
192 Mycetophyllia aliciae were observed. Besides corals, tube and branching sponges, encrusting
193 algae and cyanobacteria are present. Beginning at 18 m and shallower, small patches of Undaria
194 tenuifolia start to appear, becoming more abundant until they dominate the landscape between 12
195 and 10 m. Between 12 and 10 m, live coral cover increases to 40 – 45%, the slope reduces to 25
196 – 30o and other scleractinian species are present, including Porites astreoides, H. cucullata,
197 Colpophyllia natans, Madracis auretenra, Scolymia cubensis, and O. faveolata becomes more

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198 common. At 10 – 12 m depth, growth morphologies of typically massive species are plate like
199 and small (~ 10 – 40 cm maximum diameter). M. auretenra also form scattered monospecific
200 patches in this area.

201 At approximately 8 m, the slope decreases to 10 – 15o, corals are more abundant and larger (up
202 to 2 – 3 m diameter), but the main coral matrix is still dominated by U. tenuifolia and in some
203 areas is mixed with P. divaricata. The morphology of typically massive coral species is a mix of
204 massive and plate. The most common species are O. faveolata, O. annularis, Meandrina
205 meandrites, Pseudodiploria strigosa, M. ferox, M. cavernosa and S. siderea. At this depth, it is
206 possible to find A. cervicornis. At 6 m, coral cover increases to 50 – 60%, massive corals
207 become dominant (especially Orbicella spp.), and patches of U. tenuifolia and P. divaricata can
208 be found in sand patches. Between 5 and 3 m, massive corals dominate the reefscape, O.
209 faveolata and O. annularis colonies with diameters exceeding 5 m are common and the slope
210 decreases to almost 0o. Other common coral species include U. agaricites, C. natans, U.
211 tenuifolia, P. strigosa, P. astreoides, P. divaricata, S. cubensis, S. siderea, M. aliciae, Millepora
212 complanata, M. alcicornis, and M. striata. Live coral cover is higher than 50% and colonies of A.
213 cervicornis, A. palmata and A. prolifera are found scattered throughout the Reef. This area of
214 high coral cover which is dominated by large colonies of Orbicella spp. continues until around 3
215 m. At this depth, coral colony size and abundance decreases. Common coral species, between 3
216 and 2 m depth include P. divaricata, O. faveolata, A. tenuifolia, P. astreoides, Favia fragum, P.
217 strigosa, P. clivosa, M. cavernosa, S. siderea, A. fragilis, as well as the milleporids Millepora
218 complanata and M. striata. Most of the massive coral species’ growth morphologies change to
219 crustose, and the reef slope is less than 10o. Calcareous terraces appear at 2 m. In this area,
220 dispersed corals (P. clivosa, S. radians and S. siderea), octocorals, and sand patches are
221 common. Towards the Bay, close to the islands of Abanico and Draga, seagrasses (i.e., Thalassia
222 testudinum and Halodule wrightii) are common.

223 Varadero Reef’s benthos between 3 and 15 m is dominated by live coral (45.1 ± 3.9%) and
224 algae-overgrown dead coral (47.5 ± 4.0%; average ± SE). Sand and rubble (4.6 ± 0.6%), sponges
225 (0.7 ± 0.1%) and other invertebrates (gorgonians, zoanthids, etc.) (1.8 ± 0.9%) are also observed.
226 38 coral species (scleractinian and fire corals) were identified. The most abundant species are O.
227 faveolata (38.1%), U. agaricites (28.8%), O. annularis (14.4%) and U. tenuifolia (12.2%) (Table

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228 S1). Similar to Varadero, the most common benthic components at Barú Reef are algae-
229 overgrown dead coral (56.9 ± 2.7%) and live coral (38.1 ± 3.2%). The other benthic categories
230 assessed show low percentage cover of sand and rubble (3.4 ± 1.6%), sponges (0.8 ± 0.2%) and
231 other invertebrates (0.9 ± 0.3%). In total, 35 coral species were identified, and, similar to
232 Varadero, the most common were O. faveolata (25.6%), U. agaricites (11.3%), O. annularis
233 (10.4%) and U. tenuifolia (4.5%) (Table S1).

234 Sponge community

235 In total, fifty sponge species were observed at Varadero (38 species) and Barú (31 species)
236 Reefs. The upper shallower terraces (between 3 and 10 m depth), which were more thoroughly
237 documented through more dives and transects, hosted a total of 43 species. Upper terraces were
238 more sponge species rich at Varadero (36 in total) than Barú (25 in total), although the number of
239 species per transect were not significantly different (t-Student test, p = 0.86), 10.0 ± 1.23 species
240 per transect (mean ± 1 standard error, n = 7 transects) for Varadero, and 10.5 ± 2.36 for Barú (n
241 = 4 transects) (Table S2). Eight species were observed in greater than 50% of terrace transects on
242 both reefs, Mycale laevis, Niphates erecta, Ircinia felix, Monanchora arbuscula, Lissodendoryx
243 colombiensis, Haliclona wallentinae, Cliona laticavicola and Scopalina ruetzleri. None of these
244 common species were exclusive to either reef, and when reef-specific species were observed,
245 they were typically comprised of single occurrences. Visually, sponge abundance was similarly
246 low in both Varadero and Barú Reef terraces though there were sponge patches growing on dead
247 coral. Mean coral cover estimated from phototransects was slightly but not significantly higher in
248 Varadero than in Barú (45.1 ± 14.3% vs. 38.1 ± 12.0% respectively, t-Student test, p = 0.18, n=
249 10 transects per site, Figure 4). Sponge cover was equally low and similar between the two
250 localities (0.66 ± 0.21% and 0.80 ± 0.25% respectively, t-Student test, p = 0.52). Moreover,
251 correlations between per-transect total coral and sponge cover, although negative as expected,
252 were not significant (Varadero, r = -0.42, p = 0.22; Barú, r = -0.06, p = 0.86). Mean sponge cover
253 was also not significantly correlated with the availability of dead coral substratum (covered with
254 turf and macroalgae, Varadero, r = 0.42, p = 0.23; Barú, r = 0.36, p = 0.30), which was higher in
255 Barú (56.9 ± 18.0%) than in Varadero (51.4 ± 16.3%).

256 Fish community

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257 A total of 61 fish species from 24 families was observed at Varadero Reef compared to 44
258 species from 22 families observed at Barú. While a total of 67 species were observed at both
259 sites combined, 38 species were common to both. Twenty four species were observed at
260 Varadero only, while six species were observed exclusively at Barú. Overall, Jaccard’s
261 coefficient of similarity considering the full fish species list of each site was 0.57 (Table S3). The
262 number of species per family was similar between Varadero and Barú (r = 0.90, p << 0.001, n =
263 26 families) and at both sites damselfishes (Pomacentridae) were the most species rich (8 and 7
264 species at Varadero and Barú, respectively), followed by wrasses (Labridae; 5 species at each
265 site), groupers (Serranidae; 5 and 4 species, respectively) and parrotfishes (Scaridae; 4 species at
266 each site; Table S3).

267 Considering only data from visual censuses, a total of 834 individuals belonging to 36 species
268 were observed at Varadero, while only 519 individuals of 32 species were observed at Barú.
269 Correcting for differences in sampling effort, sample-based rarefaction indicated that, for the
270 same number of samples, species richness was slightly greater at Barú than at Varadero (Figure
271 S1). Nonetheless, mean species richness within transects at Varadero did not differ significantly
272 from mean species richness at Barú (Table 1). Except for the total number of individuals per
273 transect, which was on average significantly greater at Barú than at Varadero, none of the other
274 community parameters (Simpson’s Dominance D, Shannon’s Diversity H’, and Pielou’s
275 Evenness J’) differ significantly between Varadero and Barú (p > 0.05) (Table 1). Even though
276 there was a highly significant positive correlation between the abundance of species common to
277 both sites (considering only species observed in transects at both sites; r = 0.95, p << 0.001, n =
278 26 species), a paired Student’s t-test indicated that mean abundance was significantly greater at
279 Barú than at Varadero (mean difference = 0.78, t = -2.51, p = 0.019).

280 Results of the nMDS analysis showed that there was a great deal of overlap in fish assemblage
281 structure between Varadero and Barú considering either species composition alone (based on
282 Jaccard’s similarity; Figure 5a) or species abundance and composition (based on Bray Curtis’s
283 similarity; Figure 5b). ANOSIMs based on these two similarity measures indicated that the fish
284 assemblage at Varadero did not differ significantly from that at Barú (Jaccard-based ANOSIM, R
285 = 0.03, p = 0.37; Bray-Curtis-based ANOSIM, R = -0.06, p = 0.69).

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286 Discussion
287 Caribbean coral reefs are declining rapidly due to anthropogenic activities (e.g., overfishing,
288 pollution, etc.), climate change and the synergies between these factors. Caribbean reefs have
289 experienced declines in coral cover (and increases in macroalgae and sponge cover), and
290 reduction in the abundances of sea turtles, sharks and fish populations since the 1970s (Jackson
291 et al. 2014). Reef deterioration has not been equal throughout the Caribbean with few regions
292 still holding coral cover higher than 30% (Gardner et al., 2003). Most areas with relatively high
293 coral cover are under some conservation/management program and have experienced little
294 anthropogenic influence from land-based pollution and fisheries (Jackson et al., 2014).
295 Moreover, regional and global risk assessments correlate reefs’ vulnerability to their proximity to
296 man-made stressors (Burke et al., 2011). The discovery of an apparently healthy reef in Varadero
297 adjacent to the major population center of Cartagena, Colombia, apparently runs counter to the
298 prevailing dogma.

299 The development of coral reefs under “sub-optimal” conditions (e.g., high sedimentation,
300 nutrients) does not appear to be a widespread phenomenon, though a few disparate cases have
301 been recently reported. These anomalous reef ecosystems can be found in warm waters (Liddell
302 & Ohlhorst, 1987; Spalding & Brown, 2015), upwelling-influenced areas (Bayraktarov et al.,
303 2013), high latitudes (Harriot & Banks, 2002) and naturally turbid waters (Anthony, 2006;
304 Smithers & Larcombe, 2003). Under extreme conditions, corals have adapted and/or
305 acclimatized to the high temperature variance, and heterotrophic feeding is their dominant
306 feeding mode (Teece, et al., 2011; Hughes & Grottoli, 2013). Additionally, the shading from
307 elevated turbidity decreases photo-oxidative damage produced during warm-water stress (Lirman
308 & Fong, 2007; Manzello et al., 2015).

309 Most of the reefs subjected to ongoing or temporal sedimentation have growth constrains due to
310 the limitation on light penetration. Perry and Larcombe (2003) predicted that reef framework
311 development in turbid environments might be restricted or absent, limiting coral distribution to
312 shallow waters. Correspondingly, the portions of Varadero Reef with highest coral cover are
313 currently constrained to the shallower portions of the reef, were they appear to be autotrophic as
314 indicated by their relatively high Qm values. Environmental conditions at Varadero Reef have
315 changed drastically since the Spaniards arrived several centuries ago. As described by Restrepo

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316 et al. (2017), before the opening of the Canal del Dique during the 16th century in the colonial
317 period, and subsequent modifications in the 19th Century, Cartagena Bay had no river inputs and
318 coral reefs and seagrass beds flourished inside the Bay (Martínez et al., 2010). The massive
319 arrival of waters from the Magdalena River via the Canal del Dique, after the three major
320 modifications to the channel in 1925, 1951 and 1984 (Mogollon, 2013), drastically changed
321 conditions within the Bay from clear, warm-waters to a tidal estuarine environment (Restrepo et
322 al., 2017). The dispersion patterns of the turbid plume of the Canal del Dique in the Cartagena
323 Bay are highly variable depending on the hydrodynamic and meteorological conditions (Lonin et
324 al., 2004). In this context, the optical properties of the water at Varadero Reef could experiment
325 dramatic short-term changes depending on the prevailing hydro-meteorological conditions. The
326 description of the variability in the optical properties of the water column is key to understand
327 the energy and calcification balance of the coral community.

328 Varadero Reef is highly influenced by local stressors including eutrophication, agro-chemical
329 runoff, port and industry development, and tourism activities. The main stressor being land-
330 based pollution that flows into the Bay through the Canal del Dique (Mogollón, 2013). In
331 addition to the influx of large volumes of fresh water, sediment loads arriving into the Bay can
332 top 150 million tons per year (Restrepo et al., 2006). Varadero Reef appears to be a relic of the
333 reef formations that dominated Cartagena Bay and adjacent coastal regions during the pre-
334 Columbian period. Despite these challenging environmental conditions, our results on reef
335 structure and species composition demonstrate that Varadero Reef is a functional ecosystem,
336 fully developed and similar to those found on nearby reefs (e.g., Barú and Rosario Archipelago)
337 and Caribbean reefs more broadly (Zea, 2001; Claro & Cantelar-Ramos, 2003; Pattengill-
338 Semmens & Semmens, 2003; Valderrama & Zea, 2003; Alvarado-Chacon, Pizarro, &
339 Sarmiento-Segura, 2011; Kramer, Marks, & Turnbull, 2014).

340 The existence of Varadero, a “paradoxical reef” (López-Victoria et al., 2015), is a call for
341 scientists and managers to start looking in unexpected places for similar reefs. More importantly,
342 Varadero may hold information on reef coral resilience, resistance, and adaptations to high
343 sedimentation and turbidity. In this context, Varadero could serve as a natural laboratory and
344 potentially provide source material for reseeding future reef environments. Current reef
345 degradation challenges the initial goal of restoration ecology, meaning that returning to a pre-

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346 disturbance state might be not possible and/or practical under present climate change (van Oppen
347 et al., 2017). Tolerance to warmer and acidified waters, greater fluctuations in salinity and
348 exposure to nutrients, herbicides and other pollutants are critical coral resilience traits. Our
349 observations and preliminary results of ongoing research indicate that some of these traits can be
350 found at Varadero, but further research is needed.

351 If the dredging for a new shipping channel is authorized by government authorities (Agencia
352 Nacional de Licencias Ambientales – ANLA), we estimate that 25% of the reef will be directly
353 affected and around 50% will be indirectly affected. The environmental impacts of this dredging
354 include sediment stress (suspended and deposited), release of toxic contaminants, noise
355 contamination, and complete destruction of benthic organisms within the dredge path (Rogers,
356 1990; Erftemeijer et al., 2012; Roberts, 2012). Depending on the intensity, duration and
357 frequency of increased turbidity and sedimentation, the impacts on corals may include:
358 smothering and burial, shading, bleaching, disease (Pollock et al. 2014), and decreased survival
359 and recruitment success of coral larvae (Erftemeijer et al., 2012). Additionally, a recent review
360 on the effect of dredging on fish suggests the potential for elevated fish mortality, especially in
361 early life stages (eggs and larvae) (Wegner et al, 2017). The destruction of Varadero Reef would
362 be a loss for the scientific community, for local stakeholders and for Colombia, a country
363 struggling to emerge from a 50-year civil war conflict.

364 Acknowledgements

365 We would like to thank the community of Bocachica, specially the Eight Brothers with whom we
366 did all our fieldwork in Varadero and Barú. This community has welcomed and teach us about
367 their uses of Varadero Reef and other nearby areas. Additionally, to all the people including the
368 crew of Oregon State University from Terra, that have spread the word about Varadero Reef.

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Figure 1
Location and distribution of Varadero Reef.

Figure 1. Location and distribution of Varadero Reef. The reef continues to the South towards

Barú Island.

*Note: Auto Gamma Correction was used for the image. This only affects the reviewing manuscript. See original source image if needed for review.

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Figure 2
Varadero Reef profile

Figure 2. Profile of Varadero Reef showing the typical zonation and coral composition. Top

panels correspond to each sector of the reef and the dominant scleractinian coral

species/genus (Credit: coauthors).

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Figure 3
Varadero Reef optical properties.

Figure 3. Analyses of the variations in the optical properties of the water column in Varadero

Reef (solid circles) indicate the presence of highly stratified water masses. The blue symbols

in the blue shaded area highlight the upper layer with Kd values of 0.488 m-1, the black

symbols indicate transition region with Kd of 0.19 m-1 whereas the orange symbols in the

shaded area indicate the presence of very clear waters with Kd values of 0.041. For

comparison the monotonic vertical attenuation for the Rosario Island is presented (open

circles) with Kd values of 0.165 m-1.

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Figure 4
Varadero and Barú benthic cover.

Figure 4. Average benthic coverage Varadero (blue) and Barú (red) Reefs. Error bars indicate

standard error.

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Figure 5
Fish presence-absence and abundance data for Varadero and Barú Reefs.

Figure 5. Non-metric multidimensional scaling analysis biplots based on A) presence-absence

data (Jaccard's similarity) and B) abundance data (Bray-Curtis's similarity) for fish visual

censuses made at Barú (red) and Varadero (blue) Reefs.

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Table 1(on next page)

Fish assemblage at Varadero and Barú Reefs.

Table 1. Fish assemblage attributes estimated through visual censuses on 30 x 2 m belt

transects made at Varadero and Barú Reefs.

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 Varadero (n = 15) Barú (n = 7)
Community attribute t p
Mean ± SD Mean ± SD
Species richness 12.4 3.0 15.0 2.4 -1.99 0,06
Number of individuals 55.6 15.9 74.1 14.4 -2.62 0,02
Dominance (Simpson's D) 0.18 0.05 0.16 0.04 0.94 0,36
Diversity (Shannon's H') 2.0 0.3 2.2 0.2 -1.36 0,19
Evenness (Pielou's J') 0.81 0.07 0.80 0.04 0.27 0,79
1

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