1 Early to Middle Miocene shallow-water corals from La Guajira, Colombia

2 1,2 2,3 4 3 3 Paola Flórez , Paula Zapata-Ramírez , Carlos Jaramillo , James Klaus

1 4 Departamento de Estratigrafía y Paleontología, Universidad de Granada.

2 5 Corporación Geológica ARES, Bogotá, Colombia.

3 6 Department of Geological Sciences, University of Miami, USA.

4 7 Smithsonian Tropical Research Institute, Panamá, Panamá.

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9 Corresponding Author:

1,2 10 Paola Flórez

11 Campus Fuentenueva s/n 18002 Granada, España

12 Email address: [email protected]

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PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 21 Abstract

22

23 Here we describe and illustrate 31 Miocene corals species from the Siamaná and Jimol

24 Formations that were collected over two expeditions in the Guajira basin, Colombia during 2011

25 and 2014. Corals include 25 species, derived from 15 different genera and 12 families. Six of

26 them remain with open nomenclature. From the 25 species found in the study area, 88% are

27 extinct and the remaining under endanger status. Most of the species are hermatypic components

28 of the Scleractinian order, with the exception of a member of the Milleporidae family. The corals

29 described are composed of typical taxa from the Oligocene-Miocene transition, during which

30 they were important components in building fringing and patch reefs in the circum-

31 Caribbean/Gulf of Mexico region. The presence of typical Oligocene coral taxa such as

32 Agathiphyllia spp., Antiguastrea sp., and Diploastrea spp. from La Guajira extend the

33 distribution of these genera into the Miocene, adding a more recent geological presence in the

34 Southern Caribbean. Coral assemblages suggest a development in clear, calm and shallow waters,

35 under oligotrophic conditions and only moderate physical disturbance. Our descriptions represent

36 the first effort to characterize the taxonomy of fossilized corals in Colombia.

37

38 Introduction

39

40 Several paleontological works have been conducted to understand the macro-evolutionary

41 patterns of scleractinian corals around the world (Pandolfi, 2011; Pandolfi and Jackson, 2001),

42 and to interpret the environmental conditions that control their growth and distribution (Geister,

43 1977; Jackson et al., 1996; Pandolfi and Jackson, 2006; Novak et al., 2013). Understanding the

44 evolution of scleractinian corals on geological timescales is useful to face the present

45 environmental changes that are driving global extinction of reef-building coral species (Pandolfi,

46 2011). Moreover, coral fossil studies provide exceptional perspective into the long-term

47 maintenance of biological diversity for our future (López-Pérez, 2016).

48

49 Three important events of coral faunal turnover and speciation have occurred over three

50 transitions within the Cenozoic: the Eocene-Oligocene (ca. 34 Ma), the Oligocene-Miocene (ca.

51 23 Ma), and the Pleiocene-Pleistocene (ca. 2.6 Ma) (Budd, 2000). During the second transition

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 52 listed above (from the Oligocene into the Miocene period) the reef building capacity was greatly

53 reduced due to the loss of an estimated 50% of zooxanthellate corals (Edinger and Risk, 1994;

54 1995; Budd, 2000; Johnson et al., 2009; Budd et al., 2011). As a result, several studies have been

55 focused on fossil studies from this interval (e.g. Budd et al., 1994; Johnson, 2007; Johnson et al.,

56 2009). These works sought to understand the environmental patterns that control the distribution,

57 presence and demise of these communities in time and space.

58

59 The most representative works started with taxonomical descriptions by Vaughan (1919) and

60 later Frost and Langenheim (1974). Subsequently, more detailed descriptions of the Poritidae,

61 Astrocoeniidae and Faviidae families were provided by Budd (1986, 1987, 1991); Budd and

62 Johnson (1999), and Budd et al. (1992). However, since the paleontological works mentioned

63 above, several changes in the taxonomy and phylogeny of the cnidarians have been revealed,

64 particularly regarding the Scleractinian order (Budd and Stolarski, 2011; Budd et al., 2012)

65 highlighting important phylogenetic implications between the Pacific and the Atlantic faunas

66 (Fukami et al., 2004).

67

68 Several works have contributed paleo-environmental and paleo-oceanographic information and

69 updated the coral taxonomy of different assemblages along the circum-Caribbean/Gulf of Mexico

70 region (e.g. Geister 1975, 1983, 1992; Budd, 1980, 1987, 2000; Budd et al., 1995; 1996, 2011;

71 Johnson, 2001, 2007; Klaus and Budd, 2003; Stemann, 2004; Johnson et al., 2008, 2009 and

72 Klaus et al., 2012). These studies credited tectonic events (Roth et al., 2000; Mutti et al., 2005;

73 Newkirk and Martin, 2009), changes in ocean circulation due to the closing or narrowing of

74 gateways (e.g. the emergence of the Isthmus of Panama) (von der Heydt and Dijkstra, 2005),

75 variations in sea level (Iturralde –Vinent, 2006) as well as temperature (Mutti et al., 2005) as the

76 culprits that drive extinction.

77

78 Although these studies have provided insight into the coral diversity during important peaks of

79 speciation and extinction, few of them have been focused in the Southern Caribbean – Northern

80 South America region (e.g. Johnson et al., 2009), which our research seeks to amend.

81 Consequently, our research focused on new coral collections from Miocene paleo-reefs of the

82 Guajira Basin, bearing fossils of coral species previously thought to be extinct during the

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 83 Oligocene – Miocene transition, therefore giving them a more recent geological record in the

84 Southern Caribbean.

85

86 The purpose is to provide a taxonomical guide that will serve as a baseline for future work on the

87 Cenozoic corals of Colombia. In particular, we studied the taxonomic composition of the corals

88 and provided a description of each one of the species found, along with the occurrences in other

89 regions and their paleo-environmental characteristics.

90

91 Geological setting

92

93 The coral specimens studied were collected in the Cocinetas Basin, from the Siamaná and Jimol

94 Formations in northeast Colombia’s Guajira Peninsula (Fig. 1). Siamaná is a diachronic

95 Formation from Late Oligocene, with unities that reached the Early Miocene (Teatin, 1991;

96 Duque-Caro and Reyes, 1999). The Formation is exposed at the northeastern foothills of the

97 Serranía de Cocinas, the south of the Serranía de Jarara and the west of the Serranía de Macuira

98 (Fig. 1.2). The Early Miocene deposits are characterized by shallow reefal limestones onlapping

99 this paleohighs (Renz, 1960; Rollins, 1965; Lockwood, 1965; Macellari, 1995). Regarding the

100 thickness of Siamaná, it is highly variable ranking from 247 m just north of the Cuiza fault, to

101 over 750 m of lower Oligocene sediments, as pointed out by Duque-Caro and Reyes, (1999).

102 Rollins (1965) measured a thickness of 342 meters near to the Uitpa Formation, which overlying

103 the Siamaná. The contact between the two is discordant, especially around the edges of the basin,

104 but it tends to be transitional in the center (Rollins, 1965).

105

106 The Formation is overlain by the Early Miocene Uitpa Formation, which in turn is overlain by

107 the Middle Miocene Jimol Formation (Moreno et al., 2015). The Uitpa Formation corresponds to

108 a deep marine depositional environment and is composed of silt, selenitic clays and shales, with

109 abundant microfauna (Hendy et al., 2015; Moreno et al., 2015). Fine grained, calcareous

110 sandstone interbeds are common in the lower and upper parts of this formation (Thomas, 1972).

111 Conformably overlying the Uitpa Formation is the Jimol Formation. Jimol is dominated by

112 coarse detritic and calcareous lithologies with fewer interbedded muddy levels (Hendy et al.,

113 2015; Moreno et al., 2015) composed mainly of lithic sandstones and mudstones with high

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 114 present of fossiliferous material (Moreno et al., 2015). According to Moreno et al., (2015) the

115 Formation was deposited in a shallow marine environment at the inner shelf depth (< 50m) and

116 contains shallow marine deposits with presence of hermatypic zooxanthellate corals.

117

118

119 120 Figure 1. Study area map with the location of the stations studied.

121

122 Material and Methods

123

124 Coral fossil samples were collected in two expeditions carried out in 2010 and 2014, at five

125 localities of Cocinas Basin: Arroyo Uitpa, Arroyo Ekieps, SW Ekieps, Flor de la Guajira and

126 Punta Espada (Fig. 1, Table 1). Samples were collected manually along 10 m lateral transects,

127 located randomly in the different visited outcrops. In addition, some specimens were obtained

128 outside of lineal transect, in order to increase the taxonomical list as suggested by Johnson and

129 Kirby (2006). The lithology and the stratigraphy of each section as well as the

130 palaeoenvironmental interpretation are presented in Zapata-Ramirez et al. (in prep.).

131

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 132 The samples were cleaned and brushed with water to remove the sediment. The taxonomic

133 classification were performed following the works of Vaughan (1919); Wells (1956); Frost and

134 Langenheim (1974); Budd (1980; 1986; 1987; 1991), Johnson (2001; 2007); Johnson et al.

135 (2009); Budd et al. (1992; 2012) and Wallace (2012), principally. The classification was

136 performed by characters macro and micro structural related to the colony shape, septum

137 development, corallite diameter, number of pali, and degree of development of the columella.

138 The observations were performed with an optical equipment of 2X and 4 X magnifications, and

139 the principal measurements, described and compiled in the Table 2, were taken with a digital

140 caliper. The illustrations of the colonies and details were performed with a digital camera.

141

142 Table 1. Coordinates of the stations studied. (Station), Nomenclature of number station follow

143 the STRI projects parameters; two first characters correspond to collector, and the following to

144 station code.

145

146 147

148 The systematic paleontology of the samples is presented in Family and genera alphabetic order.

149 For each one of the species we provide a description of the characters and important remarks of

150 their classification. These results are summarized in the Table 2. Paleoenvironmental

151 assumptions are applied under the principle of uniformitarianism, which implies that, the

152 environmental conditions of modern communities or species can be employed to infer the

153 paleoenvironmental in which ancient populations inhabited (Frost and Langenheim, 1974;

154 Bosence and Allison, 1995). All coral occurrences and depositional ages presented here along

155 with the coral taxonomical description were provided by Zapata-Ramirez et al., (in prep.).

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 156

157 The reference collection has been deposited at the Mapuka Museum of Universidad del Norte

158 (MUN-STRI)(http:// www.uninorte.edu.co/web/mapuka) and voucher specimens are stored in

159 Universidad de Los Andes. All metadata of the samples and localities are available in the

160 Paleontological Smithsonian Tropical Research database (http://

161 biogeodb.stri.si.edu/jaramillo/fossildb).

162

163 SYSTEMATIC PALEONTOLOGY

164

165 Phylum CNIDARIA Verril, 1865

166 Class ANTHOZOA Ehrenberg, 1834

167 Subclass HEXACORALLIA, Haeckel, 1896

168 Order SCLERACTINIA Bourne, 1900

169

170 Family ACROPORIDAE Verrill, 1902

171

172 Acropora panamensisVaughan, 1919

173 (Pl. 1, Fig. 1)

174

175 Material.---MUN-STRI-17331, MUN-STRI-17325, MUN-STRI-17327, MUN-STRI-37928.

176

177 Description.---The corallum is plocoid, with cuneiforme to ?caespitose shape. Branches thick

178 with blunt to acuminate tips, diameter between 1 to 2.5 cm. Corallites exerts on direction to the

179 apex, with a diameter around 3mm. Calices rounded with a diameter calicular of 1-1.4 mm and

180 1.2-2 mm of intercalicular space, wall thickens of 1 mm. Septa hexamerally arranged in 2 cycles,

181 S1 reach the center of the corallite, and S2 rudimentary. Columella absent. Corallites wall and

182 coenosteum reticulo-costate, constituted by pointed spinules.

183

184 Occurrence and palaeoenvironment.--- In the Caribbean A. panamensis is recorded of Oligocene

185 to Pleistocene from Anahuac, Culebra, Valiente, La Quinta, Moneague, Lares and Seroe domi

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 186 Formations. In Siamaná Fm. was observed in fringing and patch reefs. In modern reefs, the genus

187 is a typical builder, and is found in front reefs, crest and lagoon zones (Wallace, 1999; 2012).

188

189 Acropora sp.

190 (Pl. 1, Fig. 2)

191

192 Material.--- MUN-STRI-43531, MUN-STRI-43532, MUN-STRI-43533.

193

194 Description.--- Corallum plocoid, probably arborescent or corymbose-caespitose. Branches terete

195 to slightly flattened with 6.6 to 13 mm in diameter. Corallites exerts on direction to the apex.

196 Sometimes the corallites are vertically lined, spaced apart by 2.4 to 4.2 mm, but this pattern is

197 variable and is intercalate laterally. Calices rounded of 0.9 to 1 mm in diameter. Septa

198 hexamerally arranged in 2 complete cycles. Principal septa reach the center of corallites, while

199 the secondary septa slender than S1 and, often half its width. Columella absent. Corallites wall

200 and coenosteum reticulo-costate.

201

202 Occurrence and palaeoenvironment.---In Siamaná Fm. are in beds in the fringing reefs. See A.

203 panamensis.

204

205 Remarks.---The samples are principally broken fragments, most of them without tips, and with a

206 poor preservation. Nevertheless is assigned to genus Acropora by the morphology protuberant of

207 the corallites, the absence of columella and, the spinose and costate pattern of the coenosteum.

208

209 Family AGATHIPHYLLIIDAE Vaughan y Wells, 1943

210

211 Agathiphyllia antiguensis Duncan, 1863

212 (Pl. 1, Fig. 3)

213

214 Material.--- MUN-STRI-17304, MUN-STRI-17309, MUN-STRI-17328.

215

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 216 Description.---Corallum massive and plocoid. Corallites rounded to oval in shape, 7-10 mm in

217 diameter. Bear 38-40 septa, hexamerally arranged in 4 cycles with the fourth cycle rarely

218 complete. S1, S2 reach the columella, while S3 not always, S4 ¼ o the length of S1-2. Primary

219 and secondary septa having paliform lobes only observed in transverse section. Columella

220 trabecular of 1.3-3.21 mm in diameter, usually 1/3 of corallite length. Fossa shallow. Corallite

221 wall synapticulothecal. Extratentacular budding. Coenosteum costate.

222

223 Occurrence and palaeoenvironment.---Caribbean of Oligocene to Miocene from Antigua,

224 Castillo, Rancho Berlin, San Luis and Lares Fms. A. antiguensis is indicator of shallow waters

225 and common in the building reefs, the genera is globally extinct (Budd, 2000). Colombia of

226 Siamaná Fm. in fringing reefs.

227

228 Remarks.--- The Montastraea, Agathiphyllia and Antiguastrea genera shows several similar

229 external morphological characters, whereby often are confused, especially if the samples are

230 poorly preserved by effects of the diagenetic processes (Neil-Champagne, 2011). Although the

231 Agathiphyllids are characterized by have rounded corallites, synapticulothecal wall, trabecular

232 columella and paliform lobes. Unlike to Antiguastrea which have circular to poligonal corallites,

233 parathecal wall, a lamellar columella developed, as well paliform lobes absent. By other hand,

234 although Montastraea have circular corallites, these are biggest and exert than Agathiphyllia, as

235 well the columella usually is trabecular to spongy and biggest also (Frost and Langenheim, 1974;

236 Neil-Champagne, 2001).

237

238 The reviewed samples of A. antiguensis from Siamaná Fm. are poor preserved and, not conserve

239 the paliform crown, as well the complete septa. Nevertheless was classified by the

240 synapticulothecal wall.

241

242 Agathiphyllia tenuis Duncan 1863

243 (Pl. 1, Fig. 4)

244

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 245 Material.--- MUN-STRI-17275, MUN-STRI-43509, MUN-STRI-43513, MUN-STRI-43518,

246 MUN-STRI-37877, MUN-STRI-37890, MUN-STRI-37893, MUN-STRI-37894, MUN-STRI-

247 37900, MUN-STRI-37901, MUN-STRI-37903.

248

249 Description.--- Corallum massive and plocoid. Corallites rounded to slightly compress in shape,

250 3-5 mm in diameter. Bear 20-31 septa, hexamerally arranged in 3 cycles. S1, S2 reach the

251 columella, while S3 extend to 1/3 of the total length of S1-2. Pali front S1-2, forming ?two

252 circular crowns encircling columella trabecular. Faces of septa, costae and pali finely spinose.

253 Fossa is shallow to moderately deep. Corallites with synapticulothecal wall. Budding is

254 extratentacular. Costae are thick and converge with the adjacent calices.

255

256 Occurrence and palaeoenvironment.--- Caribbean of Oligocene to Miocene from Antigua,

257 Castillo, Moneague, San Luis, Baitoa, and Lares Fms. A. antiguensis is a common in the building

258 reefs, the genera is globally extinct (Budd, 2000). From Colombia of Siamaná Fm. in fringing

259 and patch reefs.

260

261 Remarks.--- See remarks of A. antiguensis. A. tenuis is easily differs of A. antiguensis by the

262 number of cycles and size of the corallites.

263

264 Family ASCTROCOENIIDAE Koby, 1890

265

266 Astrocoenia decaturensis Vaughan, 1919

267 (Plate 1, Fig. 5)

268

269 Material.--- MUN-STRI-17294, MUN-STRI-37858, MUN-STRI-37863, MUN-STRI-37869,

270 MUN-STRI-37876, MUN-STRI-37878, MUN-STRI-37880, MUN-STRI-37881, MUN-STRI-

271 37905.

272

273 Description.---Corallum cerioid, massive to encrusting, or columnar in shape. Columns ovals in

274 transversal section of 5 x 4 cm in diameter, which could be cover by encrusting layers. Corallites

275 generally pentagonal or hexagonal with fine blunts in the calicular edge, 1.5-1.9 mm in diameter.

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 276 The most of the calices bear 16 septa in octameral arrangement, of which 8 reach the styliform

277 columella, shows a thickening close to the columella, forming a ?palar crown. The second group

278 of septa extends 1/3 to ½ of the distance of the first cycle. Septal edges with beaded teeth, 5-6 in

279 septa of 0.6 mm. The septa that reach the columella shows and thickness on the last teeth of the

280 ornate septal, forming a ?palar crown. Fossa shallow.

281

282 Occurrence and palaeoenvironment.---Caribbean of Oligocene from Antigua, and Lares Fms. In

283 Colombia of Miocene from Siamaná Fm. in fringing and patch reefs.

284

285 Remarks.---Surface of the samples poor preserved, however the knots in the septa and calicular

286 edge could be distinguished. A. decaturensis differs of A. portoricensis by develop of secondary

287 group of septa, and morphology of the colonies.

288

289 Astrocoenia portoricensis Vaughan, 1919

290 (Plate 1, Fig. 6)

291

292 Material.--- MUN-STRI-17628, MUN-STRI-17311.

293

294 Description.---Corallum cerioid and branching. Branches circular to oval in shape, 1.5 to 2 cm in

295 diameter. Corallites polygonal in shape, 1.5-2 mm in diameter. Calicular edges with blunts. The

296 regular calices bear 16 septa in octameral arrangement, 8 of them extended to the columella and,

297 the rest poor developed or rudimentary. Sporadically, corallites biggest are present, 2.52 mm in

298 diameter, with ?15-16 septa that reach the columella and other ?15-16 rudimentary. Septal edges

299 with beaded teeth, the last one forming a ?palar crown, circling the columella stylform. Fossa

300 shallow.

301

302 Occurrence and palaeoenvironment.--- Caribbean of Oligocene to Miocene from Antigua,

303 Castillo, Culebra, Moneague, Rancho Berlin and Lares. From Colombia of Siamaná Fm. in patch

304 and fringing reefs.

305

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 306 Remarks.---Samples are poor preserved, consisting of broken branches. A. portoricensis differs of

307 A. decaturensis by the morphology of the colony and the presence of giant corallites.

308

309 Astrocoenia sp.

310 (Plate 1, Fig. 7)

311

312 Material.--- MUN-STRI-43497.

313

314 Description.--- Corallum plocoid, massive and ?encrusting. Corallites circular to oval in shape, 2-

315 3 mm in diameter, spaced apart by 1-2 mm. 20 septa in decameral arrangement, 10 of them reach

316 the columella, and the rest extend to the middle or more of the total length of the first group.

317 Morphology of the columella no determinate. Calicular edges and coenosteum with blunts.

318

319 Occurrence and palaeoenvironment.--- In Colombia of Miocene from Siamaná Fm. in patch

320 reefs.

321

322 Remarks.---Sample is a fragment of colony, poor preserved.

323

324 Family CARYOPHYLLIIDAE Dana, 1846

325

326 Caryophylliidae

327 (Plate 1, Fig. 8)

328

329 Material.--- MUN-STRI-17305, MUN-STRI-43525, MUN-STRI-43528.

330

331 Description.--- Corallum faceloid, trocoid. Extretentacular budding, pedicel rises of outer

332 margins of parent corallites. Calice oval in shape, 6-8 mm in the greatest calicular diameter, and

333 4.5-5.6 mm in the minor diameter. Around 48 septa hexamerally arranged in four cycles,

334 sometimes with additional S5. Septa primary and secondary extended to the calicular center, S3

335 more of a half of the total distance of S1-2, S4 almost equal of S3 and, when S5 is present, are

336 poorly developed. Septal face bear beaded teeth. Pali and paliform lobes absent. Columella

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 337 absent or poorly developed, composed by a single blunt element. Costae present. ?Colour of

338 corallum purple.

339

340 Occurrence and palaeoenvironment.--- In Colombia of Miocene from Siamaná Fm. in fringing

341 reefs. Is not a builder coral but contributes in the fill of cavities.

342

343 Remarks.---Samples recrystallized.

344

345 Family DIPLOASTRAEIDAE Chevalier y Beauvais, 1987

346

347 Diploastrea crassolamellata Duncan, 1863

348 (Plate 1, Fig. 9)

349

350 Material.--- MUN-STRI-43488, MUN-STRI-17614, MUN-STRI-17617, MUN-STRI-17631,

351 MUN-STRI-17634, MUN-STRI-17635, MUN-STRI-17638, MUN-STRI-43499, MUN-STRI-

352 17187.

353

354 Description.---Corallum plocoid and massive. Calices slightly exerts and circular in shape, 5-7

355 mm in diameter, distance apart 1-2 mm. Calices bear 18-21 septa hexamerally arranges in three

356 cycles, all septa are exerts, thickened close to the calicular edge and reach the columella.

357 Coenosteum costate. Columella trabecular and wide, 1-2 mm in diameter, occupying 1/3 of calice

358 width.

359

360 Occurrence and palaeoenvironment.--- Caribbean, Oligocene, Early and Middle Miocene from

361 Antigua, Castillo, La Quinta, Moneague, Rancho Berlin, San Luis and Lares Fms. Colombia in

362 Siamaná and Jimol Fms. D. crassolamellata is actually extinct, and was considered a common

363 species of the American Oligocene (Frost and Langenheim, 1974; Johnson, 2007; Johnson et al.,

364 2009), nevertheless some stratigraphic unities has been re-dated into Early Miocene, e.g. Castillo

365 and San Luis Fms. (Rincón et al., 2014; Albert-Villanueva, 2016).

366

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 367 Remarks.--- Diploastrea crassolamellata has a wide morphologic variation, and could be

368 confused with samples of Montastraeidae, but could be differenced by the presence of a wall

369 synapticulothecate at the calices plane (Frost and Langenheim, 1974). The samples from Siamaná

370 Fm. are poorly preserved, however D. crassolamellata differs to D. magnifica by the calicular

371 size, which is usually minor, and the thick of the septocostae.

372

373 Diploastrea magnifica Duncan, 1863

374 (Plate 1, Fig. 10)

375

376 Material.--- MUN-STRI-17616, MUN-STRI-17618, MUN-STRI-43496, MUN-STRI-17322,

377 MUN-STRI-17182.

378

379 Description.--- Corallum plocoid and massive. Calices circular in shape, 7-10 mm in diameter,

380 distance apart 3-5 mm. Septa 42-48 hexamerally arranged in four incomplete cycles, which

381 extended to the columella. Septocostae thickened close to calicular edge. Columella trabecular, 3-

382 4 mm in diameter, extended around 1/3 of calicular width.

383

384 Occurrence and palaeoenvironment.--- Caribbean, Oligocene and Early Miocene from Antigua

385 and San Luis Fms. Colombia in Siamaná and Jimol Fms., in patch and fringing shallow reefs. D.

386 magnifica is globally extinct. The only specie living of the genera is D. heliopora of Indo-Pacific

387 waters (Veron, 2000).

388

389 Remarks.--- D. crassolamellata var. magnifica was described by Duncan (1863), and adopted by

390 Vaughan (1919), base in a major size of the corallite, less exert calices, as well a few thickness of

391 the septocostae in the wall. However Frost (1974) simonized the variety to D. crassolamelata.

392 But subsequently, Johnson and collaborators (2009) use the name Diploastrea magnifica. The

393 Colombian samples are poor preserved and do not have the calicular external structures, however

394 are classified by the low thickness of the septa in the wall and the corallite larger.

395

396 Family MERULINIDAE Verrill, 1865

397

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 398 Antiguastrea cellulosa Duncan, 1863

399 (Plate 1, Fig. 11)

400

401 Material.--- MUN-STRI-17603, MUN-STRI-43490, MUN-STRI-43493, MUN-STRI-17610,

402 MUN-STRI-17615, MUN-STRI-43494, MUN-STRI-17619, MUN-STRI-17620, MUN-STRI-

403 17622, MUN-STRI-17625, MUN-STRI-17629, MUN-STRI-17637, MUN-STRI-17640, MUN-

404 STRI-17600, MUN-STRI-17602, MUN-STRI-43498, MUN-STRI-17197, MUN-STRI-17199,

405 MUN-STRI-43500, MUN-STRI-17201, MUN-STRI-43501, MUN-STRI-17202, MUN-STRI-

406 17203, MUN-STRI-17230, MUN-STRI-17224, MUN-STRI-17287, MUN-STRI-17261, MUN-

407 STRI-17296, MUN-STRI-37886, MUN-STRI-37902, MUN-STRI-37906, MUN-STRI-37922.

408

409 Description.---Corallum subplocoid and massive. Corallites rounded to polygonal in shape, 3-4

410 mm in diameter, spaced apart by a furrow of 0.5-1 mm. Calices bear around 48 septa hexamerally

411 arranged in four complete cycles, septa primary and secondary thick and reaching the columella,

412 S3 extended about half of the total length of S1-2. S4 extended ½ of S3, or not extended away

413 from calicular wall. Columella lamellar and thin rises from a shallow fossa. Budding

414 extracalicular.

415

416 Occurrence and palaeoenvironment.---Caribbean of Oligocene to Miocene from Anahuac,

417 Anguilla Antigua Castillo Chipola, La Quinta, Moneague, Rancho Berlin, San Luis, Tampa, and

418 Lares Fms. Colombia from Siamaná in patch and fringing reefs. A. cellullosa is globally extinct.

419

420 Remarks.---Samples moderately preserved, generally recover by red algae.

421

422 ?Goniastrea canalis Vaughan, 1919

423 (Plate 1, Fig. 12)

424

425 Material.--- MUN-STRI-17332.

426

427 Description.---Corallum cerioid and massive. Calices highly irregular in shape, polygonal to

428 oval, 2.2-5.6 mm in diameter, spaced apart by 08-1.5 mm. Calices bear 28-33 septa hexamerally

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 429 arranged in three cycles. Septa primary and secondary reach the columella, depending upon

430 development of S2, and tertiary extended half or 2/3 of the length of S2, sometimes fused to

431 them. Septal faces finely granulate with small rounded granules. Wall formed by synapticulae.

432 Paliform lobes developed in ?S1-2. Columella trabecular and wide occupying around 1/3 of

433 calicular diameter. Budding intercalicular.

434

435 Occurrence and palaeoenvironment.---Caribbean of Oligocene to Miocene from Antigua,

436 Castillo, Culebra, La Quinta, Rancho Berlin, Tampa and Lares Fms. Colombia form Siamaná

437 Fm. in fringing reef environment.

438

439 Remarks.--- The identification remains uncertain because the sample is a single fragment of

440 colony, recrystallized and poorly preserved. However, the wall synapticulothecal suggests that is

441 G. canalis. In the Caribbean fossil record, Goniastrea could be confused with Favites spp., but

442 differs by the presence of abortive septa in Goniastrea, and double wall or fused walls in Favites

443 (Frost and Langenheim, 1974; Huang et al., 2014).

444

445 Family MONTASTRAEIDAE Yabe y Sugiyama, 1941

446

447 Montastraea canalis Vaughan, 1919

448 (Plate 2, Fig. 1)

449

450 Material.--- MUN-STRI-17243, MUN-STRI-17283, MUN-STRI-17290, MUN-STRI-17307,

451 MUN-STRI-17293, MUN-STRI-17298, MUN-STRI-43529, MUN-STRI-37866, MUN-STRI-

452 37874, MUN-STRI-37923, MUN-STRI-37925.

453

454 Description.---Corallum plocoid and massive. Corallites circular to slightly oval in shape,

455 moderately raised, 4-8 mm in diameter, spaced apart by 3-6 mm. Calices bear 42-49 septa,

456 hexamerally arranged in four cycles, generally complete. Primary, secondary and some tertiary

457 septa reach the columella, S4 is projecting half of S3 or less. Septal faces granulate with spaced

458 pointed granules. Paliform lobes front of S1-2, and S3 when reach the columella. Columella

459 trabecular, wide and raised extended around 1/3 of the total corallite distance. Costae well

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 460 developed, principally in S1, S2 and S3. Dissepiments endothecal and exothecal developed.

461 Reproduction by extracalicular budding.

462

463 Occurrence and palaeoenvironment.---Antigua, Castillo, Culebra, La Quinta, Rancho Berlin,

464 Tampa and Lares Fms. Colombia in Siamaná Fm. from fringing and patch reefs environments.

465 Specie is common in the buildup reefs. M. canalis is globally extinct.

466

467 Remarks.---Colonies poor preserved could be confused with Antiguastrea spp., and Agathiphyllia

468 spp., however the corallites in Montastraea spp. could be differentiated by the size of calices and

469 columella, which are biggest in Montastraea, as well as the corallites usually are more exert. See

470 Agathiphyllia antiguensis remarks.

471

472 Montastraea endothecatha Duncan, 1863

473 (Plate 2, Fig. 2)

474

475 Material.--- MUN-STRI-17229, MUN-STRI-17225, MUN-STRI-17284, MUN-STRI-17303,

476 MUN-STRI-37926.

477

478 Description.--- Corallum plocoid and massive. Corallites circular to oval in shape, moderately

479 raised, 5.3-10 mm in diameter, spaced apart by 1.4-6 mm. Calices bear around 48 septa,

480 hexamerally arranged in four cycles. Primary, secondary and tertiary septa reach the columella,

481 S4 is thin and extend 1/3 of S3 length or less. Septal faces granulate with spaced rounded or

482 pointed granules arrangement irregularly. Columella trabecular and wide, 2-3 mm in diameter.

483 Theca is septothecal. Costae dentate well developed in S1, S2 and S3, ornate with pointed

484 granules. Dissepiments endothecal and exothecal present.

485

486 Occurrence and palaeoenvironment.---Oligocene and Lower Miocene of Caribbean from

487 Anahuac, Anguilla, Antigua, Chipola, Culebra, La Quinta, Moneague, Rancho Berlin, Santa Ana,

488 Tamana, Valiente, Lares and Seroe Domi Fms. Colombia from Siamaná Fm. in fringing reefs.

489

490 Remarks.---Samples well preserved.

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 491

492 Montastraea imperatoris Vaughan, 1919

493 (Plate 2, Fig. 3)

494

495 Material.--- MUN-STRI- 43534, MUN-STRI-17246, MUN-STRI-17247, MUN-STRI-17252,

496 MUN-STRI-17253, MUN-STRI-43536, MUN-STRI-43537, MUN-STRI-17255, MUN-STRI-

497 17337, MUN-STRI-17338, MUN-STRI-17339, MUN-STRI-43538, MUN-STRI-17340, MUN-

498 STRI-17341, MUN-STRI-17342, MUN-STRI-17343, MUN-STRI-17344, MUN-STRI-43539,

499 MUN-STRI-17346, MUN-STRI-43540, MUN-STRI-43541, MUN-STRI-17347, MUN-STRI-

500 17350, MUN-STRI-17351.

501

502 Description.--- Corallum plocoid and massive. Corallites moderately raised, and circular in

503 shape, 3.2-4 mm in diameter, spaced apart by 1.7-4.2 mm. Calices bear 24 septa, hexamerally

504 arranged in 3 cycles, sometimes incomplete. Primary septa reach the columella. Columella

505 ?trabecular, formed by the union of S1. Costae well developed corresponding to all or, almost all

506 cycles. Dissepiments endothecal and exothecal well developed. Reproduction by extracalicular

507 budding.

508

509 Occurrence and palaeoenvironment.--- Caribbean of Oligocene to Miocene from Agua Clara,

510 Anahuac, Anguilla, Castillo, Culebra, Pedregoso, Tampa, Valiente, Lares and Seroe domi Fms.

511 Common specie from the middle Miocene (Budd et al., 1992). Colombia from Jimol and San

512 Andrés Fms. in patch reefs.

513

514 Remarks.---Colonies poor preserved and highly crystallized. Many characters of M. imperatoris

515 are not observed, such as the morphology of columella, the presence of paliform lobes, as well

516 the extension of primary and secondary septa. However the size, shape and raised of corallites, as

517 well the number of cycles is characteristic.

518

519 Montastraea limbata Duncan, 1863

520 (Plate 2, Fig. 4)

521

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 522 Material.--- MUN-STRI-17185.

523

524 Description.--- Corallum plocoid and massive. Corallites circular in shape, 3.5-4.2 mm in

525 diameter, spaced apart by 0.4-1.3 mm. Calices bear 24 septa, hexamerally arranged in three

526 complete cycles. Primary and secondary septa reach the columella; S3 is projecting half of S2 or

527 little more, occasionally reaching the columella. Paliform lobes front of S1-2, and S3 when reach

528 the columella. Columella trabecular extended 0.7-1 mm in diameter. Costae well developed

529 corresponding to all.

530

531 Occurrence and palaeoenvironment.---Caribbean from Miocene of Agua Clara, Pedregoso, San

532 Luis, Tamana and Seroe Domi Fms. Colombia from Jimol in patch reefs.

533

534 Remarks.--- The sample is a single fragment of colony, highly recrystallized. Because of

535 characters such as costae and dissepiments endothecal and exothecal was not possible to observe,

536 which are present in the species description. Despite to similitude between M. limbata and M.

537 imperatoris by bear three cycles and similar size of corallite, the samples differs by developing of

538 the septa, the morphology of columella, and the space apart between corallites.

539

540 Orbicella cavernosa (Linnaeus, 1767)

541 (Plate 2, Fig. 5)

542

543 Material.--- MUN-STRI-43489, MUN-STRI-43491, MUN-STRI-17607, MUN-STRI-17306,

544 MUN-STRI-17295, MUN-STRI-17329, MUN-STRI-37907, MUN-STRI-17190, MUN-STRI-

545 17192, MUN-STRI-17193.

546

547 Description.--- Corallum plocoid and massive. Corallites circular to oval in shape, 6-7 mm in

548 diameter, spaced apart by 2.5-4.5 mm. Calices bear 38-48 septa, hexamerally arranged in four

549 cycles. Primary, secondary and tertiary septa reach the columella, S4 is thin and extend 1/4 of S3

550 length or less. Columella trabecular and wide, 1-2 mm in diameter. Paliform lobes are absent.

551 Costae developed in all cycles. Dissepiments endothecal and exothecal present.

552

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 553 Occurrence and palaeoenvironment.---From Late Oligocene to Present. Caribbean from Antigua,

554 Castillo, Chipola, Baitoa, Valiente and Seroe domi. In Colombia from San Andres, Siamaná and

555 Jimol Fms. in fringing and patch reefs. Actually M. cavernosa is common specie in the

556 Caribbean, Bahamas and Florida, inhabit until 90 m depth, but usually in a range of 12-30 m.

557

558 Remarks.---The samples are poor preserved, recrystallized.

559

560 Family MUSSIDAE Ortmann, 1890

561

562 Colpophyllia willoughbiensis (Vaughan, 1919)

563 (Plate 2, Fig. 6)

564

565 Material.--- MUN-STRI-17276, MUN-STRI-17301, MUN-STRI-43515, MUN-STRI-17310,

566 MUN-STRI-17314, MUN-STRI-17318, MUN-STRI-17320, MUN-STRI-17300, MUN-STRI-

567 43526, MUN-STRI-37864, MUN-STRI-37867, MUN-STRI-37924, MUN-STRI-37927.

568

569 Description.---Corallum meandroid, massive and relatively flattened. Corallum attached to the

570 substrata by a central peduncle. Valleys large and sinuous of 10 mm width and 0.5-10 mm in

571 height. Walls usually single, but series could be separated by a furrow. The colline bear 12-13

572 septa per centimeter. Septa equal thick, 0.37-0.4 in width, septal face finely granulate. Columella

573 trabecular discontinue poorly developed or absent. Budding intracalicular. Endothecal

574 dissepiments developed and abundant.

575

576 Occurrence and palaeoenvironment.---Caribbean Eocene, Miocene in Antigua, Castillo, La

577 Quinta, Moneague, Rancho Berlin, Santa Ana y Lares Fms. Colombia in Siamaná Fm. in fringing

578 and patch reefs.

579

580 Remarks.---The samples are moderately preserved, the lower surface do not conserve the

581 epitheca. Samples from different localities have wide morphologic variation of arrangement of

582 valleys and crest (Frost and Langenheim, 1974).

583

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 584 Family POCILLOPORIDAE Gray, 1840

585

586 Pocillopora sp. B

587 (Plate 2, Fig. 7)

588

589 Material.--- MUN-STRI-17345, MUN-STRI-43542.

590

591 Description.--- Corallum plocoid and branching. Branches are thick and flattened in shape, with

592 2.8x9 cm. Corallites circular to oval, 1-2 mm in diameter, spaced between them by 0.5-1 mm.

593 Calices bear around ?12 septa. Calicular fossa is moderately deep. Columella is loose.

594 Coenosteum covered by granules and circular perforations around 0.2 mm in diameter.

595

596 Occurrence and palaeoenvironment.---Colombia from Jimol Fm. in patch reefs environments.

597

598 Remarks.---Samples poor preserved, most of the superficial structures losses. Inside is highly

599 crystallized.

600

601 Stylophora affinis Duncan, 1863

602 (Plate 2, Fig. 8)

603

604 Material.--- MUN-STRI-17608, MUN-STRI-17609, MUN-STRI-37921, MUN-STRI- 37932,

605 MUN-STRI- 37873.

606

607 Description.--- Corallum plocoid and branching. Branches robust, terete to slightly compressed in

608 shape, with 3-3.5 cm in diameter. Branches tips blunt or flattened. Corallites circular, 1-1.2 mm

609 in diameter, spaced between them by 0.4-0.8 mm. Calices bear around 12 septa, hexamerally

610 arranged in two cycles. Primary septa reach the columella, while the secondary do not apart far

611 from calicular wall. Calicular fossa shallow contains a columella styliform. Coenosteum covered

612 whit granules.

613

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 614 Occurrence and palaeoenvironment.---Oligocene to Quaternary. Caribbean from Agua Clara,

615 Castillo, Culebra, Moneague, Pedregoso, Tamana and Seroe domi Fms. Colombia from Siamaná

616 Fm. In shallow patch reefs environments. S.affinis is globally extinct.

617

618 Remarks.---Colonies poor preserved, highly crystallized.

619

620 Stylophora sp.

621 (Plate 2, Fig. 9)

622

623 Material.--- MUN-STRI-43535.

624

625 Description.--- Corallum plocoid and branching. Branches thin, terete to slightly compressed in

626 shape, with 7.4-14 mm in diameter. Branches tips loss. Corallites circular to slightly irregular of

627 variable size 0.1- 0.5 mm in diameter, arrangement irregularly in the coenosteum, spaced

628 between them by 0.4-0.8 mm. Septa and columella loss. Coenosteum covered with granules or

629 spines, and frequently with circular perforations of 0.2-0.42 mm in diameter.

630

631 Occurrence and palaeoenvironment.---In Colombia from Siamaná Fm. in patch reefs

632 environments.

633

634 Remarks.---The samples are fragments of colonies poor preserved, with many characters missing.

635 However the by the general pattern of the corallum and coenosteum, as well as the size of the

636 corallites the sample could be Stylophora sp.

637

638 Family PORITIDAE Gray, 1840

639

640 Alveopora tampae Weisbord 1973

641 (Plate 2, Fig. 10)

642

643 Material.--- MUN-STRI-43504, MUN-STRI-17268, MUN-STRI-43508, MUN-STRI-17274,

644 MUN-STRI-43517, MUN-STRI-43524, MUN-STRI-17323, MUN-STRI-37892.

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 645

646 Description.--- Corallum plocoid and columniform. Columns thick, slightly compressed

647 transversally with blunt tips, diameter between 3 to 3.5 x 2.5 cm. Corallites circular to polygonal,

648 1.9-2.9 mm in diameter, spaced apart by a calicular wall conformed by 11-12 rods of 0.3-0.5-

649 mm thick. Sinaptyculae present linked the rods of the wall. Septal spines thin, irregularly

650 arrangement in different levels, sometimes fused in the axis of the corallite.

651

652 Occurrence and palaeoenvironment.---Oligocene to Miocene from Caribbean of Antigua,

653 Castillo, Moneague, San Luis, Baitoa. From Colombia in Siamaná Fm. inhabit in patch and

654 fringing reefs.

655

656 Remarks.---Samples recrystallized and poor preserved. However is easily distinguishable for the

657 thick of the columns, and the typical spine septa and columella absence.

658

659 Goniopora hilli Vaughan 1919

660 (Plate 2, Fig. 11)

661

662 Material.--- MUN-STRI-43511, MUN-STRI-17312, MUN-STRI-17297, MUN-STRI-43521.

663

664 Description.--- Corallum subplocoid, massive, branching, columnar or contorted plates.

665 Corallites hexagonal in shape, sometimes compressed, 2.9-3.8 mm in diameter. Calices bear 24

666 septa, hexamerally arranged in three cycles. Septa dorsal and ventral are free. Septa primary and

667 secondary reach the columella, while S3 fuse to adjacent S2 close to the columella. Septal

668 margins and faces with denticles. Columella is trabecular, matted and wide, around 1 mm in

669 diameter. Wall is synapticulothecal and prominent. Fossa is moderately deep. Reproduction by

670 extracalicular budding.

671

672 Occurrence and palaeoenvironment.---Late Oligocene to Early Pleistocene Caribbean from

673 Anguilla, Culebra, La Quinta, Rancho Berlin, Tampa, Tamana and Lares Fms. Colombia in

674 Siamaná Fm. The genus is an important builder during the Cenozoic Tethys, today inhabit in the

675 Red Sea and Indo-Pacific waters. However G. hilli are extinct.

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 676

677 Remarks.---Observations of Frost and Langenheim (1974) describe a crown of 6 paliform knots

678 circling the columella, at the fused at the end of S1-2. This character does not observe in the

679 samples from Siamaná, by the preservation of the samples.

680

681 Porites anguillensis Vaughan, 1919

682 (Plate 2, Fig. 12)

683

684 Material.---MUN-STRI-43507, MUN-STRI-17279, MUN-STRI-17308, MUN-STRI-17237,

685 MUN-STRI-17239, MUN-STRI-17240, MUN-STRI-17241, MUN-STRI-17244, MUN-STRI-

686 17256, MUN-STRI-43506, MUN-STRI-17271, MUN-STRI-17277, MUN-STRI-17278, MUN-

687 STRI-17285, MUN-STRI-17288, MUN-STRI-17289, MUN-STRI-17313, MUN-STRI-43520,

688 MUN-STRI-17315, MUN-STRI-17316, MUN-STRI-43523.

689

690 Description.---Corallum subplocoid, encrusting and multilaminar with knobs. Laminae undulate

691 to flattened of 1-2.5 mm thick, spaced apart and filled with sediment or cryptic fauna. Corallites

692 circular to polygonal in shape, 1.5-2 mm in diameter, spaced apart by 0.3-0.5 mm. Corallites bear

693 12 septa arrangement in a dorsal directive free, ventral triplet fused, and four lateral pairs.

694 Columella trabecular well developed, formed by a single trabecular blunt, at the same level of the

695 palar crown. Palar crown of 5 or 6 pali. Wall conformed by one or two trabecular rings.

696 Coenosteum reticulate.

697

698 Occurrence and palaeoenvironment.---Oligocene to Miocene of Caribbean from Anguilla, La

699 Quinta, Culebra, Moneague Fms. In Colombia from Siamaná Fm. in fringing reef environment.

700

701 Remarks.---The Porites samples often confused with each other, especially if the material are not

702 good preserved.

703

704 Porites baracoaensis Vaughan, 1919

705 (Plate 3, Fig. 1)

706

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 707 Material.--- MUN-STRI-43505, MUN-STRI-43510, MUN-STRI-43514, MUN-STRI-17302,

708 MUN-STRI-43516, MUN-STRI-43519, MUN-STRI-17299, MUN-STRI-43522, MUN-STRI-

709 17324, MUN-STRI-43527, MUN-STRI-17326, MUN-STRI-43530.

710

711 Description.--- Corallum cerioid and branching. Branches are thin, circular to flattened. Branch

712 circular of 5.7-8.9 cm in diameter and flat branches 11-17 mm in wide, by 5-6 mm thick.

713 Corallites polygonal in shape, 1.3-1.4 mm in diameter, spaced apart by 0.3-0.5 mm. Corallites

714 bear 12 septa arrangement in a dorsal directive free, ventral triplet fused, and four lateral pairs.

715 Columella poor developed or absent, when is present are constitute by a small trabecula. Palar

716 crown prominent, intermediate in width, bearing 5 pali. Wall conformed by one trabecular ring.

717

718 Occurrence and palaeoenvironment.--- Oligocene to Miocene of Caribbean from Agua Clara,

719 Anahuac, Anguilla, Antigua, Castillo, Culebra, Moneague, Santa Ana, Tamana, Vliente, Lares

720 and Seroe Domi Fms. In Colombia from San Andres and Siamaná Fm. in fringing reefs

721 environments.

722

723 Remarks.---P. baracoaensis could be differentiate from others species of the branching Porites

724 by their thin calicular wall.

725

726 Porites portoricensis Vaughan, 1919

727 (Plate 3, Fig. 2)

728

729 Material.--- MUN-STRI-17200, MUN-STRI-43485, MUN-STRI-17226, MUN-STRI-17220,

730 MUN-STRI-17223, MUN-STRI-17272, MUN-STRI-17273, MUN-STRI-43484, MUN-STRI-

731 43486, MUN-STRI-17258, MUN-STRI-17259, MUN-STRI-43487, MUN-STRI-37857, MUN-

732 STRI-37862, MUN-STRI-37868, MUN-STRI-37880, MUN-STRI-37898, MUN-STRI-37899.

733

734 Description.--- Corallum cerioid, branching to columnar. Branches are thick oval to flattened.

735 Branch oval of 1.5-2.5 cm in major diameter, flat branches 2.5-3 cm in wide, by 1-2 cm thick.

736 Corallites polygonal in shape, 1.3-2 mm in diameter, spaced apart by 0.5-0.9 mm. Corallites bear

737 12 septa arrangement in a dorsal directive free, ventral triplet fused, and four lateral pairs.

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 738 Columella trabecular developed, formed by a single trabecular blunt, at the same level of the

739 palar crown. Palar crown intermediate in width of 6 pali. Wall conformed by one or two

740 trabecular rings. Coenosteum reticulate.

741

742 Occurrence and palaeoenvironment.--- Oligocene to Miocene of Caribbean from Agua Clara,

743 Anguilla, Antigua, Castillo, Culebra, La Quinta, Moneague, Baitoa, Tampa, Valiente, Lares y

744 Seroe Domi Fms. In Colombia from Siamaná Fm. in fringing and patch reefs environments.

745

746 Porites waylandi Foster, 1986

747 (Plate 3, Fig. 3)

748

749 Material.--- MUN-STRI-17604, MUN-STRI-43492, MUN-STRI-43495, MUN-STRI-17639,

750 MUN-STRI-17601, MUN-STRI-43502, MUN-STRI-43503, MUN-STRI-17242, MUN-STRI-

751 17245, MUN-STRI-17221, MUN-STRI-17222, MUN-STRI-17317, MUN-STRI-17319, MUN-

752 STRI-37857, MUN-STRI-37860, MUN-STRI-37861, MUN-STRI-37871, MUN-STRI-37872,

753 MUN-STRI-37889, MUN-STRI-37891, MUN-STRI-37896, MUN-STRI-17183, MUN-STRI-

754 17184, MUN-STRI-17186, MUN-STRI-17248, MUN-STRI-17249, MUN-STRI-17336, MUN-

755 STRI-17348, MUN-STRI-17349.

756

757 Description.--- Corallum subplocoid, columnar to massive with knobs. Corallites polygonal in

758 shape, 1.3-1.8 mm in diameter, spaced apart by 0.5 mm. Corallites bear 12 septa arrangement in a

759 dorsal directive free, ventral triplet fused, and four lateral pairs. When are present, the columella

760 is trabecular, formed by a single trabecular blunt, at the same level of the palar crown or lowest.

761 Palar crown wide of 4-5 pali. Wall conformed by one or two trabecular rings. Coenosteum

762 reticulate.

763

764 Occurrence and palaeoenvironment.--- Oligocene to Miocene of Caribbean from Agua Clara,

765 Anahuac, Anguilla, Castillo, Culebra, La Quinta, Moneague, San Luis, Tampa, Tamana,

766 Valiente, Lares and Seroe Domi Fms. Colombia from Siamaná and Jimol Fms. in fringing and

767 patch reefs environments.

768

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 769 Porites sp.

770 (Pl. 3, Fig. 4)

771

772 Material.---MUN-STRI-17254.

773

774 Description.---Corallum branching, plocoid to subplocoid. Branches much compressed, of 23-29

775 mm in thick and 37-110 mm in length, whit evidence of anastomosis in the grow pattern.

776 Corallites rounded to slightly compressed, 1.3 to 2 mm in diameter. Distance apart of the calices

777 of 0.6 to 0.8 mm. Septa arranged in two cycles completes. Columella not evident. Coenosteum

778 with circular perforations, 0.3-0.4 mm in diameter. Fossa deep.

779

780 Occurrence and palaeoenvironment.---Early and Middle Miocene from Culebra, Castillo and

781 Lares Fms. Colombia of Jimol Fm. in a patch reef.

782

783 Remarks.--- Preservation is too poor. The sample consists in two recrystallized broken branches,

784 whereby several characters not was observed, such as coenosteum costate, presence of tertiary

785 septa, as well the difference between S1 and S2, columella poorly developed, and calicular rims

786 slightly exert.

787

788 Family SIDERASTREIDAE Vaughan y Wells, 1943

789

790 Siderastrea conferta Duncan, 1863

791 (Plate 3, Fig. 5-6)

792

793 Material.--- MUN-STRI-17265, MUN-STRI-17270, MUN-STRI-43512, MUN-STRI-17291.

794

795 Description.--- Corallum cerioid and massive. Corallites tetra, penta or hexagonal in shape, 4-10

796 mm in diameter. Calices bear 54-67 septa, which could be confluent or not with adjacent

797 corallites. Septa hexamerally arranged in five cycles always incomplete. Septa uniformly spaced,

798 primary cycle reach the columella free, while the rest are fused to adjacent systems. S3 fuse to

799 adjacent S2 close to the columella, S4 fuse to S3 at half or ¾ of the total length of S1, and when

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 800 present, S5 fuse to adjacent S4 at close to the calicular wall. Septal margins bear acute teeth, 6-7

801 by millimeter, and the septal faces granulate with thick trabeculae, generally fused to the adjacent

802 septa. Paliform lobes are absent. Columella is trabecular with few components or weekly

803 developed. Calices are shallow, septa equally exert forming a convex surface, that fall soft toward

804 the columella. Wall is synapticulothecal.

805

806 Occurrence and palaeoenvironment.---Agua Clara, Anahuac, Anguilla, Antigua, Castillo,

807 Culebra, La Quinta, Moneague, Rancho Berlín, San Luis and Lares Fms. In Colombia from San

808 Andrés, and Siamaná Fms. in fringing reef. Common in the build of the reef and, lagoon zones.

809

810 Remarks.---Colonies well preserved, easily distinguish of S. siderea by the morphology of the

811 colonies.

812

813 Siderastrea siderea (Ellis y Solander, 1786)

814 (Plate 3, Fig. 7-8)

815

816 Material.--- MUN-STRI-17260, MUN-STRI-17269, MUN-STRI-17263, MUN-STRI-17292,

817 MUN-STRI-17250, MUN-STRI-17251.

818

819 Description.--- Corallum cerioid and massive. Corallites penta or hexagonal in shape, 3.5-5 mm

820 in diameter. Calices bear 45-50 septa, which could be confluent or not with adjacent corallites.

821 Septa hexamerally arranged in four cycles. Septa uniformly spaced, which go down into the fossa

822 in a softly slope, generally the primary cycle reach the columella free, rest of them are fused to

823 adjacent systems while reaching the columella, generally S4 to S3 and S3 to S2, forming trident

824 patterns. Septal margins bear acute teeth.

825 Septal faces granulate with thick trabeculae, sometimes fused to the adjacent septa. Paliform

826 lobes are absents. Columella is trabecular with several components. Wall is synapticulothecal.

827 Reproduction by extracalicular budding.

828

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 829 Occurrence and palaeoenvironment.--- At present this species inhabit the paths reef in La Guajira

830 Peninsula (Díaz et al., 2010; Reyes et al., 2010), and remains an important reefal buildup in the

831 Caribbean region.

832

833 Remarks.---Despite the crystallization intern of the most of the corallum, the surface of the

834 colonies remain preserved.

835

836 Orden ANTHOATHECATA Cornelius, 1992

837

838 Family MILLEPORIDAE Fleming, 1828

839

840 Millepora alcicornis Linnaeus, 1758

841 (Plate 3, Fig. 9)

842

843 Material.--- MUN-STRI-17218, MUN-STRI-17286.

844

845 Description.---Corallum ramose. Branches terete to flattened, with anastomosis, 8-16 mm in

846 diameter at the mid of the branch. Branches tips rounded and bifurcated, 6-9 mm in diameter.

847 Corallum surface reticulate, composed of a of rods meshwork. Pores rounded with density 2 2 848 between 25 and 37 per cm . Gastropore of 0.4 mm in diameter, 1 or 2 gastropore per cm .

849 Dactylopore with diameter ranges between 0.29-0.3 mm. Ampullae and arrangements of

850 cyclosystems are not distinguished.

851

852 Occurrence and palaeoenvironment.--- Caribbean from Eocene to present day. Colombia from

853 Siamaná Fm., in fringing reefs. Species common in the buildup reef, and lagoon zones.

854

855 Remarks.---Samples poor preserved could be confused with samples of the scleractinian Porites

856 spp., or the octocoral Heliopora sp. because the similarity of the coenosteum surface, which is

857 meshwork. But M. alcicornis could be differenced by the skeleton growth form, as well details of

858 the pore. In modern samples, the morphology of M. alcicornis is highly variable, from branching,

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 859 encrusting to hemispheric colonies (Amaral et al., 2008), whereby could be found similar forms

860 in the fossil record.

861

862 Discussion

863

864 A total of 270 samples were collected in the study area. From this total, 80% were identified to

865 species level, 4% to genera level and the remaining 8%, to family level (Table 2). In general, the

866 species assemblages reported in this study are taxa commonly found between the late Oligocene

867 and Early-Middle Miocene in the Caribbean region (Fig. 2). In some cases, some of these species

868 (12 species) reach the Quaternary with the exception of Siderastrea siderea and Montastraea

869 limbata whose first occurrences were reported from the Early Miocene (Jung, 1971; Frost and

870 Langenheim, 1974; Geister, 1975; Johnson et al., 2009).

871

872 873

874 Figure 2. Range chart of first and last occurrence in the Great Caribbean for the species found in

875 Siamaná and Jimol Formations.

876

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 877 In particular, Siderastrea Siderea was reported for the first time also in Colombia at the San

878 Andres Formation (Geister, 1975). Besides, 48% of the species found became extinct during the

879 Middle Miocene, 40% at the end of Miocene or during the Quaternary, and finally the remaining

880 12% of the species continue to inhabit the reefs today (Fig. 2) (Frost and Langenhelm, 1974;

881 Budd et al., 1995; Budd, 2000; Johnson, 2007; Johnson et al., 2009). These species are the

882 scleractinians Siderastrea siderea, Orbicella cavernos and the hydrocoral Millepora alcicornis.

883 They represent the winners after the turnover events and are presented in the study region since

884 the early Miocene till today characterizing patch communities in protected Bays (Díaz et al.,

885 2000; Reyes et al., 2010).

886

887 The Siamaná and Jimol Formations are also characterized by a high richness of Porites spp. and

888 Montastraea spp. This richness indicates the resilience of corals to maintain the largest spatial

889 distribution even in areas such la Guajira with strong upwelling effects, high sedimentation and

890 fluctuant salinity (Díaz et al., 2000; Reyes et al., 2010). The presence of these two genera in the

891 Siamaná Formation suggest protected areas in shallow waters. These assemblages were also

892 accompanied by species of massive shapes of Agathyphyllia, Antiguastrea, Astrocoenia,

893 Colpophyllia and Acropora sp genera, supporting as well low-energy wave regime characteristic

894 of lagoon environments. Nevertheless, nowadays Acropora sp is also a genera that characterize

895 high environments such the reef crest (Schuster, 2000), representing an exception of the

896 uniformitarianism theory. Outside of the Guajira Peninsula, the Siamaná Formation has most taxa

897 in common with the Late Oligocene Antigua Formation of Antigua (36 species) and the Lares

898 Formation (41 species). Other unites with high similarity include the Early Miocene Castillo (21

899 species) and San Luis Formations (11 species) of the Falcon Basin in Venezuela.

900

901 Dominant taxa of the Jimol Formation, include five species of the genera Montastraea,

902 Pocillopora, Porites and Siderastrea with predominant massive shapes (Table 2), which also

903 indicate characteristic of shallow waters with moderate physical disturbance. The Jimol

904 Formation has most taxa in common with Middle Miocene Formations of Valiente Formation (22

905 species) in Panamá and with Seroe Domie Formation (29 species) of Curacao.

906

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 907 Table 2. Species list and summary of the principal taxonomic classification characters of the

908 samples reviewed from Siamaná and Jimol Formations. Colony growth: B: branching, M:

909 massive, P: platy; Ph: phaceloid. E: encrusting, K: Knobs. CD: calicular diameter, (* dactilopore

910 in milleporids). ICD: Intercalicular calicular diameter. Coenosteum: Cos: costae presents, Tb:

911 Tubercles, Sp: spongy, Sm: smooth. No. cicles: inc.: incomplete. Kind of columella: St:

912 Styliform, L: lamellar, T: trabecular, A: absent. In all items (-) means not determined. As well as

913 showing the Formations were found it.

914

915 916

917 As depicted in Fig. 2 a gross temporal distribution is presented in the coral species from the late

918 Eocene and in the case of Antiguastrea cellulosa even from the late Cretaceous. The taxa

919 recovered from the Siamaná Formation deposits reveals an extended temporal distribution of

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 920 species such as Antiguastrea cellulosa, Colpophyllia willoughbiensis, Diploastrea

921 crassolamellata, Montastrea endothecata, Agathiphyllia tenuis and Siderastrea conferta

922 previously thought to be exclusively from the Oligocene (Vaughan, 1919; Budd, 2000; Johnson,

923 2007; Johnson et al., 2009). The presence of these species suggest different local/environmental

924 responses that perhaps allowed them to stay longer in the study area.

925

926 Otherwise, and according to Wells (1959) and Frost and Leighenhaim (1974), genera such as

927 Alveopora, Diploastrea, and Antiguastrea are relict fauna of the European pool that start to

928 disappear in the Late Oligocene. However, in our case the presence of these genera was

929 maintained after the early Miocene and just after the middle Miocene they start to disappear and

930 be replaced by new assemblages of the Jimol Formation as outlined above.

931

932 Comparing our assemblages with early Miocene shallow water corals from the Indo-Pacific and

933 the Mediterranean Sea, just few similarities were found to species level. Thus, only Porites

934 baracoensis was a common specie with the Indo-Pacific (Bromfield, 2013) while at genera level

935 Porites spp. and Acropora spp., are both presented in the Mediterranean and at the Indo-Pacific

936 waters (Bromfield, 2013; Santodomingo et al., 2016).

937

938 Conclusions

939

940 This study increases understanding of coral distribution in the southern Caribbean during the

941 Oligocene- Miocene period, through detailed taxonomical analysis of samples collected during

942 field activities in the Guajira Basin. Additionally, insight was gained into the temporal variations

943 of specific species when compared with their first and last occurrences at other regional reef

944 assemblages showing that differences could be related with local/regional environmental events.

945

946 Despite that a better understanding about the Cenozoic history of the Guajira Basin coral fauna

947 was gained and the compilation will serve as a baseline for future works in Colombian reefs. We

948 stress, however, that more data collection is necessary in the study area to fully understand the

949 evolutionary story of corals in the region before firm conclusions about the timing and localities

950 that better represent the turnover events during the Neogene in the Guajira Basin.

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 951

952 Acknowledgements

953

954 This study was supported by Colciencias Grant Agreement No. 7277 569 33195, contract 2013-

955 0217. Project: Links Between Marine Biotic Evolution and Carbonate Platform and Petroleum

956 Reservoir Development in the Guajira Basin, Colombian Caribbean. We acknowledge the

957 support of Ecopetrol, as well as the Smithsonian Tropical Research Institute STRI, University of

958 Zurich, Universidad del Norte and Universidad de Granada. PF is supported by Colciencias

959 scholarship ‘Doctorados en el exterior 2015’. We are also grateful to the Wayúu community for

960 their hospitality and guidance in the field. Special thanks to Dr. Juan Carlos Braga (Universidad

961 de Granada) for providing helpful comments.

962

963

964

965

966

967

968

969

970

971

972

973

974

975

976

977

978

979

980

981

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 982

983

984

985

986

987

988

989 Plate 1

990

991 General view of the colonies.

992

993 1. Acropora panamensis (MUN-STRI-17331), scale bar 3 cm.

994 2. Acropora sp. (MUN-STRI-), scale bar 4 cm.

995 3. Agathiphyllia antiguensis (MUN-STRI-17309), scale bar 3 cm.

996 4. Agathiphyllia tenuis (MUN-STRI-17275), scale bar 1 cm.

997 5. Astrocoenia decaturensis (MUN-STRI-17294), scale bar is 2 cm.

998 6. Astrocoenia portoricensis (MUN-STRI-17311), scale bar 2.5 cm.

999 7. Astrocoenia sp. (MUN-STRI-43497), scale bar 3.5 cm.

1000 8. Caryophyllidae (MUN-STRI-43528), scale bar 4 mm.

1001 9. Diploastrea crassolamellata (MUN-STRI-17635), scale bar is 8 mm.

1002 10. Diploastrea magnifica (MUN-STRI-43496), scale bar is 5 mm.

1003 11. Antiguastrea cellulosa (MUN-STRI-17224), scale bar is 2 cm.

1004 12. ?Goniastrea canalis (MUN-STRI-17332), scale bar is 1.5 mm.

1005

1006

1007

1008

1009

1010

1011

1012

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 1013 1014

1015 Plate 1

1016

1017

1018

1019

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 1020

1021

1022

1023

1024

1025

1026 Plate 2

1027

1028 General view of the colonies.

1029

1030 1. Montastraea canalis (MUN-STRI-17883), scale bar 2 cm.

1031 2. Montastraea endothecatha (MUN-STRI-17229), scale bar 3 cm.

1032 3. Montastraea imperatoris (MUN-STRI-17344), scale bar 1.5 cm.

1033 4. Montastraea limbata (MUN-STRI-17185), scale bar 1.5 cm.

1034 5. Orbicella cavernosa (MUN-STRI-17306), scale bar 2 cm.

1035 6. Colpophyllia willoughbiensis (MUN-STRI-17318), scale bar 2 cm.

1036 7. Pocillopora sp. B. (MUN-STRI-43542), scale bar 2 cm.

1037 8. Stylophora affinis (MUN-STRI-17608), scale bar 2 cm.

1038 9. Stylophora sp. (MUN-STRI-43535), scale bar 2 cm.

1039 10. Alveopora tampae (MUN-STRI-43508), scale bar 2 cm.

1040 11. Goniopora hilli (MUN-STRI-43521), scale bar 2 cm.

1041 12. Porites anguillensis (MUN-STRI-17285), scale bar 2 cm.

1042

1043

1044

1045

1046

1047

1048

1049

1050

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 1051

1052 1053

1054 Plate 2

1055

1056

1057

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 1058

1059

1060

1061

1062

1063

1064 Plate 3

1065

1066 General view of the colonies.

1067

1068 1. Porites baracoaensis (MUN-STRI-43527), scale bar 1.5 cm.

1069 2. Porites portoricensis (MUN-STRI-43486), scale bar 1.8 cm.

1070 3. Porites waylandi (MUN-STRI-17222), scale bar 2 cm.

1071 4. Porites sp. (MUN-STRI-17254), scale bar 3 cm.

1072 5. Siderastrea conferta (MUN-STRI-17270), scale bar 3 cm.

1073 6. Siderastrea conferta (MUN-STRI-17270), detail of the coralite, scale bar 6 mm.

1074 7. Siderastrea siderea (MUN-STRI-17269), scale bar 2.5 cm.

1075 8. Siderastrea siderea (MUN-STRI-17263), detail of the coralite, scale bar 2 mm.

1076 9. Millepora alcicornis (MUN-STRI-17286), scale bar 4 cm.

1077

PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2507v1 | CC BY 4.0 Open Access | rec: 8 Oct 2016, publ: 8 Oct 2016 1078 1079

1080 Plate 3

1081

1082 References

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