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Early to Middle Miocene shallow-water corals from La Guajira, Colombia 1
2
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
8
Corresponding Author: 9
Paola Flórez1,2 10
Campus Fuentenueva s/n 18002 Granada, España 11
Email address: [email protected] 12
13
14
15
16
17
18
19
20
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Abstract 21
22
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
37
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
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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
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Oligocene – Miocene transition, therefore giving them a more recent geological record in the 83
Southern Caribbean. 84
85
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
90
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
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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
117
118
119
Figure 1. Study area map with the location of the stations studied. 120
121
Material and Methods 122
123
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
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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
145
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
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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
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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
Material.--- MUN-STRI-17304, MUN-STRI-17309, MUN-STRI-17328. 214
215
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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
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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
Material.--- MUN-STRI-17294, MUN-STRI-37858, MUN-STRI-37863, MUN-STRI-37869, 269
MUN-STRI-37876, MUN-STRI-37878, MUN-STRI-37880, MUN-STRI-37881, MUN-STRI-270
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
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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
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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
Material.--- MUN-STRI-17305, MUN-STRI-43525, MUN-STRI-43528. 329
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
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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
Material.--- MUN-STRI-43488, MUN-STRI-17614, MUN-STRI-17617, MUN-STRI-17631, 350
MUN-STRI-17634, MUN-STRI-17635, MUN-STRI-17638, MUN-STRI-43499, MUN-STRI-351
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
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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
Material.--- MUN-STRI-17616, MUN-STRI-17618, MUN-STRI-43496, MUN-STRI-17322, 376
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
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Antiguastrea cellulosa Duncan, 1863 398
(Plate 1, Fig. 11) 399
400
Material.--- MUN-STRI-17603, MUN-STRI-43490, MUN-STRI-43493, MUN-STRI-17610, 401
MUN-STRI-17615, MUN-STRI-43494, MUN-STRI-17619, MUN-STRI-17620, MUN-STRI-402
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
MUN-STRI-43500, MUN-STRI-17201, MUN-STRI-43501, MUN-STRI-17202, MUN-STRI-405
17203, MUN-STRI-17230, MUN-STRI-17224, MUN-STRI-17287, MUN-STRI-17261, MUN-406
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
Material.--- MUN-STRI-17332. 425
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
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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
Material.--- MUN-STRI-17243, MUN-STRI-17283, MUN-STRI-17290, MUN-STRI-17307, 450
MUN-STRI-17293, MUN-STRI-17298, MUN-STRI-43529, MUN-STRI-37866, MUN-STRI-451
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
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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
Material.--- MUN-STRI-17229, MUN-STRI-17225, MUN-STRI-17284, MUN-STRI-17303, 475
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
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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
MUN-STRI-17253, MUN-STRI-43536, MUN-STRI-43537, MUN-STRI-17255, MUN-STRI-496
17337, MUN-STRI-17338, MUN-STRI-17339, MUN-STRI-43538, MUN-STRI-17340, MUN-497
STRI-17341, MUN-STRI-17342, MUN-STRI-17343, MUN-STRI-17344, MUN-STRI-43539, 498
MUN-STRI-17346, MUN-STRI-43540, MUN-STRI-43541, MUN-STRI-17347, MUN-STRI-499
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
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Material.--- MUN-STRI-17185. 522
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
Material.--- MUN-STRI-43489, MUN-STRI-43491, MUN-STRI-17607, MUN-STRI-17306, 543
MUN-STRI-17295, MUN-STRI-17329, MUN-STRI-37907, MUN-STRI-17190, MUN-STRI-544
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
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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
Material.--- MUN-STRI-17276, MUN-STRI-17301, MUN-STRI-43515, MUN-STRI-17310, 565
MUN-STRI-17314, MUN-STRI-17318, MUN-STRI-17320, MUN-STRI-17300, MUN-STRI-566
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
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Family POCILLOPORIDAE Gray, 1840 584
585
Pocillopora sp. B 586
(Plate 2, Fig. 7) 587
588
Material.--- MUN-STRI-17345, MUN-STRI-43542. 589
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
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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
Material.--- MUN-STRI-43535. 623
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
Material.--- MUN-STRI-43504, MUN-STRI-17268, MUN-STRI-43508, MUN-STRI-17274, 643
MUN-STRI-43517, MUN-STRI-43524, MUN-STRI-17323, MUN-STRI-37892. 644
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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
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Page 24
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-17239, MUN-STRI-17240, MUN-STRI-17241, MUN-STRI-17244, MUN-STRI-685
17256, MUN-STRI-43506, MUN-STRI-17271, MUN-STRI-17277, MUN-STRI-17278, MUN-686
STRI-17285, MUN-STRI-17288, MUN-STRI-17289, MUN-STRI-17313, MUN-STRI-43520, 687
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
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Page 25
Material.--- MUN-STRI-43505, MUN-STRI-43510, MUN-STRI-43514, MUN-STRI-17302, 707
MUN-STRI-43516, MUN-STRI-43519, MUN-STRI-17299, MUN-STRI-43522, MUN-STRI-708
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
Material.--- MUN-STRI-17200, MUN-STRI-43485, MUN-STRI-17226, MUN-STRI-17220, 729
MUN-STRI-17223, MUN-STRI-17272, MUN-STRI-17273, MUN-STRI-43484, MUN-STRI-730
43486, MUN-STRI-17258, MUN-STRI-17259, MUN-STRI-43487, MUN-STRI-37857, MUN-731
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
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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
Occurrence and palaeoenvironment.--- Oligocene to Miocene of Caribbean from Agua Clara, 742
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
Material.--- MUN-STRI-17604, MUN-STRI-43492, MUN-STRI-43495, MUN-STRI-17639, 749
MUN-STRI-17601, MUN-STRI-43502, MUN-STRI-43503, MUN-STRI-17242, MUN-STRI-750
17245, MUN-STRI-17221, MUN-STRI-17222, MUN-STRI-17317, MUN-STRI-17319, MUN-751
STRI-37857, MUN-STRI-37860, MUN-STRI-37861, MUN-STRI-37871, MUN-STRI-37872, 752
MUN-STRI-37889, MUN-STRI-37891, MUN-STRI-37896, MUN-STRI-17183, MUN-STRI-753
17184, MUN-STRI-17186, MUN-STRI-17248, MUN-STRI-17249, MUN-STRI-17336, MUN-754
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
Occurrence and palaeoenvironment.--- Oligocene to Miocene of Caribbean from Agua Clara, 764
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
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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
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Page 28
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
Material.--- MUN-STRI-17260, MUN-STRI-17269, MUN-STRI-17263, MUN-STRI-17292, 816
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
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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
Material.--- MUN-STRI-17218, MUN-STRI-17286. 843
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
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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
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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
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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
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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
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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
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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
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1013
1014
Plate 1 1015
1016
1017
1018
1019
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1020
1021
1022
1023
1024
1025
Plate 2 1026
1027
General view of the colonies. 1028
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
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1051
1052
1053
Plate 2 1054
1055
1056
1057
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1058
1059
1060
1061
1062
1063
Plate 3 1064
1065
General view of the colonies. 1066
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
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1078
1079
Plate 3 1080
1081
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1083
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Amaral, F.M.D., Steiner, A.Q., Broadhurst, M.K., and Cairns, S.D., 2008, an overview of the 1087
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Budd, A.F., 1980, Ecology and morphology of the Caribbean Mio-Pliocene reef-coral 1093
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