Early to Middle Miocene shallow-water corals from La ...peerj.com/preprints/2507.pdf1" Early to Middle Miocene shallow-water corals from La Guajira, Colombia 2" 3" Paola Flórez1,2,

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

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Paola Flórez1,2, Paula Zapata-Ramírez2,3, Carlos Jaramillo4, James Klaus3 3

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

2 Corporación Geológica ARES, Bogotá, Colombia. 5

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

4 Smithsonian Tropical Research Institute, Panamá, Panamá. 7

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Corresponding Author: 9

Paola Flórez1,2 10

Campus Fuentenueva s/n 18002 Granada, España 11

Email address: [email protected] 12

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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

Abstract 21

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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Introduction 38

39

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

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

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

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

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

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

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

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

48

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

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

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


listed above (from the Oligocene into the Miocene period) the reef building capacity was greatly 52

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

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

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

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

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

58

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

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

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

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

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

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

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

(Fukami et al., 2004). 66

67

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

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

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

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

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

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

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

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

culprits that drive extinction. 76

77

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

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

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

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

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


Oligocene – Miocene transition, therefore giving them a more recent geological record in the 83

Southern Caribbean. 84

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

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

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

regions and their paleo-environmental characteristics. 89

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Geological setting 91

92

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

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

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

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

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

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

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

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

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

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

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

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

105

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

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

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

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

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

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

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

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


present of fossiliferous material (Moreno et al., 2015). According to Moreno et al., (2015) the 114

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

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

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

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Material and Methods 122

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

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

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

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

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

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

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

131


The samples were cleaned and brushed with water to remove the sediment. The taxonomic 132

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

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

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

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

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

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

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

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

141

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

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

station code. 144

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146

147

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

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

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

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

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

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

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

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


156

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

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

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

Paleontological Smithsonian Tropical Research database (http:// 160

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

162

SYSTEMATIC PALEONTOLOGY 163

164

Phylum CNIDARIA Verril, 1865 165

Class ANTHOZOA Ehrenberg, 1834 166

Subclass HEXACORALLIA, Haeckel, 1896 167

Order SCLERACTINIA Bourne, 1900 168

169

Family ACROPORIDAE Verrill, 1902 170

171

Acropora panamensisVaughan, 1919 172

(Pl. 1, Fig. 1) 173

174

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

176

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

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

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

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

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

coenosteum reticulo-costate, constituted by pointed spinules. 182

183

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

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


Formations. In Siamaná Fm. was observed in fringing and patch reefs. In modern reefs, the genus 186

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

188

Acropora sp. 189

(Pl. 1, Fig. 2) 190

191

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

193

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

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

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

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

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

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

and coenosteum reticulo-costate. 200

201

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

panamensis. 203

204

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

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

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

208

Family AGATHIPHYLLIIDAE Vaughan y Wells, 1943 209

210

Agathiphyllia antiguensis Duncan, 1863 211

(Pl. 1, Fig. 3) 212

213


215


Description.---Corallum massive and plocoid. Corallites rounded to oval in shape, 7-10 mm in 216

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

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

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

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

wall synapticulothecal. Extratentacular budding. Coenosteum costate. 221

222

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

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

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

Siamaná Fm. in fringing reefs. 226

227

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

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

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

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

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

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

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

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

Neil-Champagne, 2001). 236

237

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

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

synapticulothecal wall. 240

241

Agathiphyllia tenuis Duncan 1863 242

(Pl. 1, Fig. 4) 243

244


Material.--- MUN-STRI-17275, MUN-STRI-43509, MUN-STRI-43513, MUN-STRI-43518, 245

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

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

248

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

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

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

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

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

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

255

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

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

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

and patch reefs. 259

260

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

number of cycles and size of the corallites. 262

263

Family ASCTROCOENIIDAE Koby, 1890 264

265

Astrocoenia decaturensis Vaughan, 1919 266

(Plate 1, Fig. 5) 267

268



37905. 271

272

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

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

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


The most of the calices bear 16 septa in octameral arrangement, of which 8 reach the styliform 276

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

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

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

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

281

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

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

284

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

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

group of septa, and morphology of the colonies. 287

288

Astrocoenia portoricensis Vaughan, 1919 289

(Plate 1, Fig. 6) 290

291

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

293

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

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

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

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

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

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

shallow. 300

301

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

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

and fringing reefs. 304

305


Remarks.---Samples are poor preserved, consisting of broken branches. A. portoricensis differs of 306

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

308

Astrocoenia sp. 309

(Plate 1, Fig. 7) 310

311

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

313

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

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

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

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

318

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

reefs. 320

321

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

323

Family CARYOPHYLLIIDAE Dana, 1846 324

325

Caryophylliidae 326

(Plate 1, Fig. 8) 327

328


330

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

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

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

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

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

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


absent or poorly developed, composed by a single blunt element. Costae present. ?Colour of 337

corallum purple. 338

339

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

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

342

Remarks.---Samples recrystallized. 343

344

Family DIPLOASTRAEIDAE Chevalier y Beauvais, 1987 345

346

Diploastrea crassolamellata Duncan, 1863 347

(Plate 1, Fig. 9) 348

349



17187. 352

353

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

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

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

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

width. 358

359

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

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

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

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

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

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

366


Remarks.--- Diploastrea crassolamellata has a wide morphologic variation, and could be 367

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

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

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

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

372

Diploastrea magnifica Duncan, 1863 373

(Plate 1, Fig. 10) 374

375


MUN-STRI-17182. 377

378

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

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

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

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

383

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

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

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

waters (Veron, 2000). 387

388

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

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

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

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

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

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

395

Family MERULINIDAE Verrill, 1865 396

397


Antiguastrea cellulosa Duncan, 1863 398

(Plate 1, Fig. 11) 399

400



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

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



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

408

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

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

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

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

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

extracalicular. 414

415

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

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

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

419

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

421

?Goniastrea canalis Vaughan, 1919 422

(Plate 1, Fig. 12) 423

424


426

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

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


arranged in three cycles. Septa primary and secondary reach the columella, depending upon 429

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

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

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

calicular diameter. Budding intercalicular. 433

434

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

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

Fm. in fringing reef environment. 437

438

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

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

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

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

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

444

Family MONTASTRAEIDAE Yabe y Sugiyama, 1941 445

446

Montastraea canalis Vaughan, 1919 447

(Plate 2, Fig. 1) 448

449



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

453

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

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

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

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

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

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


developed, principally in S1, S2 and S3. Dissepiments endothecal and exothecal developed. 460

Reproduction by extracalicular budding. 461

462

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

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

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

466

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

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

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

Agathiphyllia antiguensis remarks. 470

471

Montastraea endothecatha Duncan, 1863 472

(Plate 2, Fig. 2) 473

474


MUN-STRI-37926. 476

477

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

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

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

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

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

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

granules. Dissepiments endothecal and exothecal present. 484

485

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

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

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

489

Remarks.---Samples well preserved. 490


491

Montastraea imperatoris Vaughan, 1919 492

(Plate 2, Fig. 3) 493

494

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





17350, MUN-STRI-17351. 500

501

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

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

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

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

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

budding. 507

508

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

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

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

Andrés Fms. in patch reefs. 512

513

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

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

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

well the number of cycles is characteristic. 517

518

Montastraea limbata Duncan, 1863 519

(Plate 2, Fig. 4) 520

521



523

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

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

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

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

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

corresponding to all. 529

530

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

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

533

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

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

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

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

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

539

Orbicella cavernosa (Linnaeus, 1767) 540

(Plate 2, Fig. 5) 541

542



17192, MUN-STRI-17193. 545

546

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

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

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

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

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

552


Occurrence and palaeoenvironment.---From Late Oligocene to Present. Caribbean from Antigua, 553

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

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

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

557

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

559

Family MUSSIDAE Ortmann, 1890 560

561

Colpophyllia willoughbiensis (Vaughan, 1919) 562

(Plate 2, Fig. 6) 563

564



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

568

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

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

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

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

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

dissepiments developed and abundant. 574

575

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

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

and patch reefs. 578

579

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

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

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

583


Family POCILLOPORIDAE Gray, 1840 584

585

Pocillopora sp. B 586

(Plate 2, Fig. 7) 587

588


590

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

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

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

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

595

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

597

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

crystallized. 599

600

Stylophora affinis Duncan, 1863 601

(Plate 2, Fig. 8) 602

603

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

MUN-STRI- 37873. 605

606

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

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

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

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

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

whit granules. 612

613


Occurrence and palaeoenvironment.---Oligocene to Quaternary. Caribbean from Agua Clara, 614

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

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

617

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

619

Stylophora sp. 620

(Plate 2, Fig. 9) 621

622


624

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

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

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

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

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

630

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

environments. 632

633

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

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

corallites the sample could be Stylophora sp. 636

637

Family PORITIDAE Gray, 1840 638

639

Alveopora tampae Weisbord 1973 640

(Plate 2, Fig. 10) 641

642


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


645

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

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

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

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

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

651

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

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

fringing reefs. 654

655

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

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

658

Goniopora hilli Vaughan 1919 659

(Plate 2, Fig. 11) 660

661

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

663

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

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

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

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

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

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

extracalicular budding. 670

671

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

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

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

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


676

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

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

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

680

Porites anguillensis Vaughan, 1919 681

(Plate 2, Fig. 12) 682

683

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




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

689

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

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

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

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

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

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

Coenosteum reticulate. 696

697

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

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

700

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

good preserved. 702

703

Porites baracoaensis Vaughan, 1919 704

(Plate 3, Fig. 1) 705

706




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

710

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

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

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

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

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

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

717

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

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

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

environments. 721

722

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

by their thin calicular wall. 724

725

Porites portoricensis Vaughan, 1919 726

(Plate 3, Fig. 2) 727

728




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

733

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

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

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

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


Columella trabecular developed, formed by a single trabecular blunt, at the same level of the 738

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

trabecular rings. Coenosteum reticulate. 740

741


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

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

745

Porites waylandi Foster, 1986 746

(Plate 3, Fig. 3) 747

748







STRI-17348, MUN-STRI-17349. 755

756

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

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

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

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

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

reticulate. 762

763


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

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

patch reefs environments. 767

768


Porites sp. 769

(Pl. 3, Fig. 4) 770

771

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

773

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

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

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

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

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

779

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

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

782

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

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

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

slightly exert. 786

787

Family SIDERASTREIDAE Vaughan y Wells, 1943 788

789

Siderastrea conferta Duncan, 1863 790

(Plate 3, Fig. 5-6) 791

792

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

794

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

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

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

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

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


present, S5 fuse to adjacent S4 at close to the calicular wall. Septal margins bear acute teeth, 6-7 800

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

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

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

the columella. Wall is synapticulothecal. 804

805

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

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

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

809

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

colonies. 811

812

Siderastrea siderea (Ellis y Solander, 1786) 813

(Plate 3, Fig. 7-8) 814

815


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

818

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

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

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

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

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

patterns. Septal margins bear acute teeth. 824

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

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

Reproduction by extracalicular budding. 827

828


Occurrence and palaeoenvironment.--- At present this species inhabit the paths reef in La Guajira 829

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

Caribbean region. 831

832

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

colonies remain preserved. 834

835

Orden ANTHOATHECATA Cornelius, 1992 836

837

Family MILLEPORIDAE Fleming, 1828 838

839

Millepora alcicornis Linnaeus, 1758 840

(Plate 3, Fig. 9) 841

842


844

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

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

Corallum surface reticulate, composed of a of rods meshwork. Pores rounded with density 847

between 25 and 37 per cm2. Gastropore of 0.4 mm in diameter, 1 or 2 gastropore per cm2. 848

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

cyclosystems are not distinguished. 850

851

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

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

854

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

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

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

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


encrusting to hemispheric colonies (Amaral et al., 2008), whereby could be found similar forms 859

in the fossil record. 860

861

Discussion 862

863

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

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

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

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

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

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

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

871

872

873

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

Siamaná and Jimol Formations. 875

876


In particular, Siderastrea Siderea was reported for the first time also in Colombia at the San 877

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

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

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

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

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

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

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

2000; Reyes et al., 2010). 885

886

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

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

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

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

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

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

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

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

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

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

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

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

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

900

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

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

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

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

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

906


Table 2. Species list and summary of the principal taxonomic classification characters of the 907

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

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

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

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

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

showing the Formations were found it. 913

914

915

916

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

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

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


species such as Antiguastrea cellulosa, Colpophyllia willoughbiensis, Diploastrea 920

crassolamellata, Montastrea endothecata, Agathiphyllia tenuis and Siderastrea conferta 921

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

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

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

925

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

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

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

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

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

931

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

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

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

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

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

937

Conclusions 938

939

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

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

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

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

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

945

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

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

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

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

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


951

Acknowledgements 952

953

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

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

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

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

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

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

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

de Granada) for providing helpful comments. 961

962

963

964

965

966

967

968

969

970

971

972

973

974

975

976

977

978

979

980

981


982

983

984

985

986

987

988

Plate 1 989

990

General view of the colonies. 991

992

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

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

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

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

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

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

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

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

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

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

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

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

1005

1006

1007

1008

1009

1010

1011

1012


1013

1014

Plate 1 1015

1016

1017

1018

1019


1020

1021

1022

1023

1024

1025

Plate 2 1026

1027


1029

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

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

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

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

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

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

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

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

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

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

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

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

1042

1043

1044

1045

1046

1047

1048

1049

1050


1051

1052

1053

Plate 2 1054

1055

1056

1057


1058

1059

1060

1061

1062

1063

Plate 3 1064

1065


1067

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

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

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

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

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

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

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

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

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

1077


1078

1079

Plate 3 1080

1081

References 1082

1083

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