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Agassizia excentrica Valenciennes, in Agassiz & Desor, 1847, San Gregorio Formation, Falcón state, Venezuela, UNEFM-IF-030. 1, detail of petals, showing anterior petal with only one row of pore pairs, and short posterior petal with less developed upper part of the anterior series of the pore pairs. 2, test, lateral view. Scale bar equals 1 mm.
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The extensive Venezuelan coastline is very important for understanding the evolu-tion of the Caribbean marine fauna. We report new fossil material collected from three Neogene fossil sites in the Falcón Basin and present the first diversity analysis of the known fossil echinoids from Venezuela and other Caribbean regions. Five species are reported...
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Citations
... In contrast, the fossil record of the genus Arbacia is poor in South America. It is restricted to representatives of the extant species A. stellata from the upper Pliocene-Pleistocene of Ecuador (Canoan Formation, Di Celma et al., 2002) and A. punctulata from the lower Pleistocene of Venezuela (San Gregorio Formation;Mihaljević et al., 2010). This poor fossil record departs from the present diversity of the genus that comprises the four extant species A. stellata (blainville, 1825), A. punctulata (Lamarck, 1816), A. dufresnii blainville, 1825, and A. spatuligera (Valenciennes, 1846), without mentioning the closely related species Tetrapygus niger (Molina, 1782). ...
... The first fossils of the genus Arbacia ever reported from 2002) and a specimen of Arbacia punctulata from Venezuela (Mihaljević et al., 2010), the genus Arbacia has been mainly reported as a fossil from North America so far, in addition to few specimens from the West Indies (Curaçao, Cuba;de buisonjé, 1974). Therefore, the new material described here fills in an important gap in the fossil record of the genus and improves our knowledge of its paleobiogeography. ...
Abstract. Recent fieldwork on Neogene deposits of northern Chile led to the discovery of new material, including three new species of the echinoid genus Arbacia: Arbacia quyllur sp. nov. from the Miocene, Arbacia terraeignotae sp. nov., and Arbacia larraini sp. nov. from the Pliocene. In the Pleistocene, the new material includes the first fossil occurrence of the extant species Arbacia spatuligera and new specimens of the extant species Tetrapygyus niger. The specimens and new species described here reveal the significant past diversity of the genus Arbacia and increase our knowledge of the overall diversity of the genus in South America, showing an older origin (upper Miocene) and higher species richness of the genus Arbacia along the Chilean coasts than previously thought.
Key words. Paleobiogeography. Fossils. Echinoid. Arbacia. Chile.
Resumen. Recientes trabajos de campo en el Neógeno del norte de Chile han permitido descubrir nuevos materiales, entre ellos tres nuevas especies del género de equinoideo Arbacia: Arbacia quyllur sp. nov. (Mioceno), Arbacia terraeignotae sp. nov. y Arbacia larraini sp. nov. (Plioceno), pero también la primera aparición de Arbacia spatuligera y nuevo material de Tetrapygyus niger (Pleistoceno). Los especímenes y las nuevas especies aquí descritas revelan la diversidad pasada del género Arbacia. Estos descubrimientos muestran la existencia de una mayor riqueza de Arbacia a lo largo de las costas chilenas y un origen más antiguo del género en la costa del Pacífico Sureste (Mioceno superior).
Palabras clave. Paleobiogeografía. Fósiles. Equinoideo. Arbacia. Chile.
... Echinoderms from the tropical Neogene marine deposits of America have been recorded in different countries and specific sedimentary basins (Antigua: Blake et al., 2015;Belize: Donovan et al., 2005; Brazil: discussed here; Costa Rica: Alvarado et al., 2006;Cuba: Roig andBrodermann, 1949, Donovan, 1994;Guadeloupe: Donovan, 1994;Grenadines: Jagt et al., 2014;Jamaica: Donovan, 1994, Donovan et al., 2005, Blake et al., 2015Puerto Rico: Vélez-Juarbe and Santos, 2008;Trinidad: Cooke, 1961;and Venezuela: Jeannet, 1928, Cooke, 1941, 1961, Mihaljević et al., 2010, Lodeiros et al., 2013. However, the potential use of echinoderms for detailed paleostratigraphy interpretation continued to be scarce. ...
... These chaotic arrangement of fossil in both Ponta do Castelo and Fazenda outcrops were discussed by Ferreira and Cunha (1957). Similar high-energy event in tropical America is exhibited in the northwestern Caribbean coast of Venezuela (Mihaljević et al., 2010), in which Lytechinus A. Agassiz (1863), Prionocidaris, and Clypeaster were combined in the coarse-grained sandy carbonate rocky matrix. ...
... However, some authors report closer faunal relationships between similarly aged units from the northern coast of Brazil and the Caribbean (foraminifera: Petri, 1957;bryozoans: Ramalho et al., 2017;ostracods: Nogueira and Nogueira, 2017;Nogueira et al., 2019;mollusks: Maury, 1925). The echinoid assemblages from the Pirabas Formation studied here show a lower generic diversity (early/middle Miocene: 11 genera) compared with those of the Caribbean (early Miocene: 42 and middle Miocene: 40), Greater Antilles (26 and 26), Venezuela (12 and 21), and Cuba (21 and 19) (Mihaljević et al., 2010). The low diversity and the specific endemism from the Pirabas Formation could be an artifact caused by insufficient sampling. ...
Echinoderms are one of the most important invertebrates in the early to middle Miocene heterozoan carbonate deposits of the Pirabas Formation of the northern Brazil. The well-preserved echinoid tests were useful for accurate identification, whereas the sediment filling the tests provide significant information regarding the rock matrix. The dominance of disarticulated ophiuroid, crinoid, and asteroid plates, and damage on the echinoderm test provides a basis for taphonomic processes to elucidate a paleoenvironmental interpretation based on the echinoderm assemblages. Nineteen echinoderm taxa (ten valid species and nine species in open nomenclature) were recorded based on well-preserved and/or fragmented tests and/or spines, including Abertellidae †Abertella pirabensis (Santos, 1958), Placatenellidae †Placatenella complanata (Brito, 1981), Clypeasteridae †Clypeaster lamegoi Santos1958, †Clypeaster paraensis Brito1979, and †Clypeaster paulinoi Santos1958, Cassidulidae †Anisopetalus oliveirai (Santos, 1958) and †Rhyncholampas candidoi Fernandes and Morais1994, Echinolampadidae †Echinolampas paraense Santos1958, Schizasteridae †Agassizia eugeniae Brito and Ramires1974 and Schizaster sp., Cidaridae Cidaris sp., †Phyllacanthus priscus Brito and Ramires1974, and Prionocidaris sp. The smaller echinoderm fragments were dominated by disarticulated plates of Gorgonocephalidae ophiuroids, Comatulidae crinoid Sievertsella sp., Goniasteridae asteroids, as well as spines and tests fragments of Cidaridae echinoids. The echinoderm assemblages suggest an inshore and shallow water paleoenvironment over a soft sandy carbonate bottom. MicroCT analyses of sediments filling echinoid tests exhibited a microfossil assemblage, including foraminifera, sponges, ahermatypic corals, bryozoans, and mollusks. Broken and disarticulated tests and plates in chaotic orientations suggest high-energy paleoenvironment. Evidence of test damage indicates complex predator-prey interactions. Overall, the microCT analyses of echinoids tests proved to be a high accuracy non-destructive, analytical approach for investigating fossil echinoderms and their paleoenvironment.
... Isolated ossicles and fragments of test and broken spines are insufficient for allowing an accurate identification (Santos, 1958(Santos, , 1967Brito and Ramires, 1974;Fernandes and Morais, 1994;Mooi et al., 2018) are not observed in the Aricuru outcrop. The paleodiversity of Echinodermata from the Aricuru is low compared to other nearshore paleoenvironments from the Caribbean (e.g., the Oligocene asteroids (Goniasteridae) from Jamaica and Antigua: Blake et al., 2015; the middle Miocene asteroids (Goniasteridae) from the Grenadines: Jagt et al., 2014; late Miocene to Pliocene echinoids (Cidaridae and Clypeasteridae) from Venezuela: Mihaljević et al., 2010, Lodeiros et al., 2013; late Oligocene to early Miocene echinoids (Prenasteridae) from Trinidad: Jeannet, 1928). ...
Outcrops of Neogene carbonates of the North Brazilian platform occur exclusively along the equatorial coast and represent some of the few existing examples of Neogene carbonate systems of the Atlantic coast of South America. The Pirabas Formation (early-middle Miocene) is the northernmost part of this platform. Although onshore mainly consists of small and scattered outcrops, it considerably extends in the subsurface keeping record of relevant geological and paleontological episodes of tropical South American history. Intending to improve the knowledge of South American carbonate and providing a solid basis for future comparisons between the Pirabas Basin and other, largely subsurface, Cenozoic basins, this research investigates the Aricuru outcrops by combining a standard petrographical and paleontological approach with advanced microCT analyses. The Aricuru area is characterized by mixed siliciclastic-carbonate sand-sized deposits, which probably deposited after the early Miocene according to the palynological assemblage. The bioclastic fraction of the rock is dominated by benthic foraminifera (mostly soritids, amphisteginids, small rotaliids, and small miliolids, typical of marginal marine environment), bryozoans, calcareous algae (Halimeda), echinoderms and mollusks. The abundant siliciclastic fraction together with the taxonomic composition of the foraminiferal, echinodermal, crustacean, ichnofossil and fish assemblages indicates deposition in a tropical coastal environment featuring both protected and more exposed sectors. The carbonate system probably developed under abundant nutrient supply, which fostered heterotroph suspension feeders over hermatypic corals. The demise of this system was most likely caused by a growth in siliciclastic input due to increased rainfall in the coastal area. The sedimentary evolution of the Pirabas Formation is similar to the one of the Foz do Amazonas Basin, and fits well with the general evolutionary trend of Cenozoic carbonate factories of the region, indicating the potential of the Pirabas subsurface record for understanding other Cenozoic basins, their paleoenvironmental significance, and their potential as oil, gas and water reservoirs.
... The type section of the Cocuiza Member includes 80 m and forms 9 to 18 layers with plenty of fossils, especially bivalve mollusks (González de Juana et al., 1980;Léxico Estratigráfico de Venezuela, 1997). The higher foraminifera diversity in the Cocuiza Member suggests a nearshore paleoenvironment with carbonate influence (Smith et al., 2010), which is consistent with the occurrence of tropical, shallow-water fossil echinoids (Mihaljević et al., 2010) and crustacean decapods . ...
The examined Ariidae marine catfish from the Neogene of tropical America consisted of isolated skulls, otoliths and bone fragments, some of which were described independently as otolith-based species or skull-based species. We used three-dimensional digital rendering (microCT) of skull and otolith reconstructions to recognize anatomical patterns including skull-otolith morphology, spatial allocations of otoliths in the endocranium for taxonomic identifications. We recognized isolated Proto-Caribbean otoliths of Cathorops sp. from the late early Miocene to early Pliocene formations and isolated otoliths of †Aspistor verumquadriscutis, †Bagre urumacoensis and Notarius sp. from the late Miocene. We explored the endocrania of four fossil Ariidae skulls from the late Oligocene to late early Miocene Proto-Caribbean to determine their internal otolith-cranial morphology, and we identified and described the skulls of †Bagre protocaribbeanus and †Cantarius nolfi and erected the new species of †Bagre castilloensi n. sp. and †Bagre ornatus n. sp. based on the internal otolith-skull association. The first fossil record of Bagre marinus from the early Pliocene Cubagua Formation to the late Pliocene San Gregorio Formation completed the ariid geochronological sequence. We discuss the differential stages of fossil preservation of bioapatite skulls and aragonite otoliths according to the diagenetic processes as well as the paleoenvironmental conditions in the sedimentary basins. Detailed microCT, 3D reconstructions, X-rays, dry prepared skeletons and digital photos of otolith and skull are shown to elucidate the in-skull otoliths species descriptions.
... Additionally, Cooke (1961) described E. falconensis from the Miocene-Pliocene of Venezuela; however, this species has since been placed into the genus Mellitella (Mihaljevic et al., 2010). The Miocene of Venezuela also contains E. kugleri Jeannet, 1928;E. ...
... The Miocene of Venezuela also contains E. kugleri Jeannet, 1928;E. vonderschmitti Jeannet, 1928[which Cooke (1961 did not discuss in his review of Venezuelan species, but Mihaljevic et al. (2010) retained]; E. wiedenmayeri Jeannet, 1928, and the Pliocene species E. secoensis Cooke, 1961. Cooke (1961 fig. ...
The lower Pleistocene Waccamaw Formation of northeastern South Carolina and southeastern North Carolina is recently understood to contain at least ten species of echinoids making it the most diverse Pleistocene echinoid fauna along the east coast of North America. An exposure along the Intracoastal Waterway in North Myrtle Beach, South Carolina displays the basal bed of the unit with an exceptionally diverse echinoid assemblage. A new species (Encope dubarorum) recently described by some of the authors has attributed to this increase in local diversity. Additionally, the variable morphologies of Mellita caroliniana may either represent other new species or a single, highly-plastic species. These two recognized species are abundant components of the Waccamaw Formation, occurring more frequently within the basal shelly, calcareous sand of this unit. Together, they both account for over half of the relative abundance of echinoid species.
In addition to the general morphology of these echinoids, their oral plate sutures and patterns are diagnostic and used to distinguish closely related species. Given that E. dubarorum is an exceptionally robust, thick margined species, with a somewhat inflated oral surface, the oral suture plates and patterns were not easily discerned. The visibility of these sutures were enhanced by gentle polishing with carbide and subsequent staining with a mixture of Alizarin Red S and potassium ferricyanide. In future studies, such stains may be applied to better observe plate margins and suture patterns.
... Additionally, Cooke (1961) described E. falconensis from the Miocene-Pliocene of Venezuela; however, this species has since been placed into the genus Mellitella (Mihaljevic et al., 2010). The Miocene of Venezuela also contains E. kugleri Jeannet, 1928;E. ...
... The Miocene of Venezuela also contains E. kugleri Jeannet, 1928;E. vonderschmitti Jeannet, 1928[which Cooke (1961 did not discuss in his review of Venezuelan species, but Mihaljevic et al. (2010) retained]; E. wiedenmayeri Jeannet, 1928, and the Pliocene species E. secoensis Cooke, 1961. Cooke (1961 fig. ...
The lower Pleistocene Waccamaw Formation of northeastern South Carolina and southeastern North Carolina is now known to contain ten species of echinoids making it the most diverse Pleistocene echinoid fauna along the east coast of North America. Herein, the echinoid fauna of the Waccamaw Formation and underlying upper Pliocene Goose Creek Limestone is discussed. The new species Encope dubarorum is described and documented from both units in Horry County, South Carolina. Remarks on an exposure along the Intracoastal Waterway in North Myrtle Beach, South Carolina, that displays the Waccamaw Formation and Goose Creek Limestone contact is also included. The basal bed of the Waccamaw Formation at this site contains an exceptionally diverse echinoid assemblage and represents the this echinoid fauna is documented and discussed.
... He also presented more information regarding the locality where the specimens were collected, Punta Gavilán; nonetheless, he also assigned C. falconensis to the Middle Miocene. Mihaljević et al. (2010) reported two records for this species: Eurhodia falconensis, as originally described by Jeannet (1928) placed in the original description; and Cassidulus (Cassidulus) falconensis, which was the classification proposed by Cooke (1961). Both records were assigned to the Middle Miocene. ...
Inclusion of fossils can be crucial to address evolutionary questions, because their unique morphology, often drastically modified in recent species, can improve phylogenetic resolution. We performed a cladistic analysis of 45 cassidulids with 98 characters, which resulted in 24 most parsimonious trees. The strict consensus recovers three major cassiduloid clades, and the monophyly of the family Cassidulidae is not supported. Ancillary analyses to determine the sensitivity of the phylogeny to missing data do not result in significantly different topologies. The taxonomic implications of these results, including the description of a new cassiduloid family and the evolution of some morphological features, are discussed. Cassiduloids (as defined here) most probably originated in the Early Cretaceous, and their evolutionary history has been dominated by high levels of homoplasy and a dearth of unique, novel traits. Despite their high diversity during the Palaeogene, there are only seven extant cassiduloid species, and three of these are relicts of lineages dating back to the Eocene. Future studies of the biology of these poorly known species, some of which brood their young, will yield further insights into the evolutionary history of this group.
... The sediments of San Gregorio Fm. are composed mainly of mudstones and limestones, with intercalations of sandstones and pebbly conglomerates (Quiroz and Jaramillo, 2010;Vucetich et al., 2010), with a total of more than 300 m at the type section (Ministerio de Energía y Minas, 1997). Recent paleontological studies of this formation described invertebrates (e.g., molluscs, crustaceans, foraminiferans) from the overlaying marine Cocuiza Member (Hambalek et al., 1994;Aguilera et al., 2010;Mihaljević et al., 2010). From its basal Vergel Member, crocodylians (Crocodylus falconensis, in Scheyer et al., 2013), caviomorph rodents (Cardiatherium sp. and cf. ...
A skull of a ground sloth from the Pliocene San Gregorio Formation documents a northern neotropical occurrence of a megatheriine that addresses issues on intraspecific variation and biogeography. The new specimen is broadly similar in size and morphology to that of Proeremotherium eljebe from the underlying Codore Formation in the Urumaco Sequence, differing in several features such as a longer basicranial area and a more posteriorly projected basioccipital between the condyles. The living sloths species of Bradypus and Choloepus do not have unequivocal anatomical features that
indicate sexual dimorphism. Nevertheless, fossil sloths may have shown such dimorphism, and speculations on this subject are part of the considerations that can be made when allocating fragmentary fossils (e.g., in the new skull the presence of a long sagittal crest could indicate a male individual and the absence of an extended crest in Proeremotherium eljebe a female one). We speculate that as early as the late middle Miocene, two main lines of Megatheriinae had clearly separated in two geographic areas, one in the rising Andean area and one at low latitudes on the lowlands of central and northern South America.
... The preservation of body fossils of Eupatagus ornatus (family Eupatagidae) in life position within B. monastiriensis in Oligocene deposits of Italy (Bernardi et al., 2010) also reinforces the idea that other families of spatangoidscan produce Bichordites. The genus Echinocardium is unknown in the Caribbean seas, including in Cuba (Mihaljevi c et al., 2010). Moreover, Pickerill et al. (1993) did not ascribe Echinocardium to be the producer of Bichordites of the Pleistocene deposits of Jamaica, because there is no record of that genus known in the Cenozoic or recent of the entire Caribbean region. ...
... Moreover, Pickerill et al. (1993) did not ascribe Echinocardium to be the producer of Bichordites of the Pleistocene deposits of Jamaica, because there is no record of that genus known in the Cenozoic or recent of the entire Caribbean region. Nevertheless, the fossils of the Maretiidae and Eupatagidae families are known in Venezuela, Barbados, Jamaica, Cuba, Puerto Rico, Antigua, and other Caribbean regions (Jackson, 1922;Clark, 1927, 1934;S anchez Roig, 1926S anchez Roig, , 1949S anchez Roig, , 1953Cooke, 1961;Kier, 1984;Donovan, 1993;Donovan and Lewis, 1993;Donovan et al., 2005;Mihaljevi c et al., 2010). In particular, the body fossils of Eupatagus and/or Maretia, which are also interpreted to be the Bichordites tracemakers as noted above, occur in Cuba (S anchez Roig, 1926;Kier, 1984;Donovan and Lewis, 1993) and Eupatagus in Jamaica (Jackson, 1922;Clark, 1927, 1934). ...
The trace fossil Bichordites monastiriensis is found in early Eocene turbiditic sandstones of the upper-slope deposits from the Capdevila Formation in Los Palacios Basin, Pinar del Río region, western Cuba. The potential tracemakers of B. monastiriensis include fossil spatangoids from the family Eupatagidae. The record of Bichordites in the deposits from Cuba allows to suppose that Eupatagidae echinoids were the oldest potential tracemakers of Bichordites isp. and reinforce the hypothesis that the ichnological record are relevant in envisaging the evolutionary history of the spatangoids.
... The Diodontidae are durophagous fishes that use the power of the fused jaws and the crushing tooth battery to feed on prey with a hard protected shell (mollusks), carapace (crustaceans) or test (echinoids) [41,42,43,44]. Consequently the broad distribution in tropical marine shallow waters over sandy, rocky and coral reef bottoms is likely to be related to the available food resources in coastal paleoenvironments, particularly the high abundance and diversity of Neogene tropical American mollusks [54,72,89,99,123], crustaceans [12,124,125,126] and echinoids [127,128,129,130] in all of sedimentary basins where the fossils were found. ...
Fossil Diodontidae in Tropical America consist mostly of isolated and fused beak-like jawbones, and tooth plate batteries. These durophagous fishes are powerful shell-crushing predators on shallow water invertebrate faunas from Neogene tropical carbonate bottom, rocky reefs and surrounding flats. We use an ontogenetic series of high-resolution micro CT of fossil and extant species to recognize external and internal morphologic characters of jaws and tooth plate batteries. We compare similar sizes of jaws and/or tooth-plates from both extant and extinct species. Here, we describe three new fossil species including †Chilomycterus exspectatus n. sp. and †Chilomycterus tyleri n. sp. from the late Miocene Gatun Formation in Panama, and †Diodon serratus n. sp. from the middle Miocene Socorro Formation in Venezuela. Fossil Diodontidae review included specimens from the Neogene Basins of the Proto-Caribbean (Brazil: Pirabas Formation; Colombia: Jimol Formation, Panama: Gatun and Tuira formations; Venezuela: Socorro and Cantaure formations). Diodon is present in both the Atlantic and Pacific oceans, whereas the distribution of Chilomycterus is highly asymmetrical with only one species in the Pacific. It seems that Diodon was as abundant in the Caribbean/Western Atlantic during the Miocene as it is there today. We analyze the paleogeographic distribution of the porcupinefishes group in Tropical America, after the complete exhumation of the Panamanian isthmus during the Pliocene.
