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From the Late Miocene Gatun Formation of Panama Catalina Pimiento,1,2,3 Gerardo Gonza´ Lez-Barba,4 Dana J

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From the Late Miocene Gatun Formation of Panama Catalina Pimiento,1,2,3 Gerardo Gonza´ Lez-Barba,4 Dana J Journal of Paleontology, 87(5), 2013, p. 755–774 Copyright Ó 2013, The Paleontological Society 0022-3360/13/0087-755$03.00 DOI: 10.1666/12-117 SHARKS AND RAYS (CHONDRICHTHYES, ELASMOBRANCHII) FROM THE LATE MIOCENE GATUN FORMATION OF PANAMA CATALINA PIMIENTO,1,2,3 GERARDO GONZA´ LEZ-BARBA,4 DANA J. EHRET,5 AUSTIN J. W. HENDY,1,2 1 2 BRUCE J. MACFADDEN, AND CARLOS JARAMILLO 1Florida Museum of Natural History, University of Florida, Gainesville, FL 32611, USA, ,[email protected].; ,[email protected].; ,[email protected].; 2Smithsonian Tropical Research Institute, Center for Tropical Paleoecology and Archaeology, Box 2072, Panama, Republic of Panama, ,[email protected].; 3Department of Biology, University of Florida, Gainesville, FL 32611, USA; 4Museo de Historia Natural, Area de Ciencias del Mar, La Paz, Universidad Autonoma de Baja California Sur, AP 23080, Mexico, ,[email protected].; and 5Alabama Museum of Natural History, University of Alabama, Tuscaloosa, Alabama, 35487, USA, ,[email protected]. ABSTRACT—The late Miocene Gatun Formation of northern Panama contains a highly diverse and well sampled fossil marine assemblage that occupied a shallow-water embayment close to a purported connection between the Pacific and Atlantic (Caribbean) oceans. However, the diverse chondrichthyan fauna has been poorly documented. Based on recent field discoveries and further analysis of existing collections, the chondrichthyan fauna from this unit comprises at least 26 taxa, of which four species are extinct today. The remaining portion of the total chondrichthyan biodiversity has affinities with modern taxa and is therefore comprised of long-lived species. Based on known records of the modern geographic distribution range of the Gatun chondrichthyans, the fauna has mixed biogeographic affinities suggesting that around 10 million yr ago, a connection likely occurred between the Pacific Ocean and the Caribbean Sea. Given the known habitat preferences for modern chondrichthyans, the Gatun fauna was primarily adapted to shallow waters within the neritic zone. Finally, comparisons of Gatun dental measurements with other faunas suggest that many of the taxa have an abundance of small individuals, in agreement with previous studies that proposed this area as a paleonursery habitat for the species Carcharocles megalodon. INTRODUCTION noted three species from the Miocene deposits of Monkey Hill HE LATE Miocene Gatun Formation comprises fossiliferous (present-day Colon). Of most significance to the current study, T sediments that crop out on the Caribbean coast of the Gillette (1984) described the fossil chondrichthyans from the Isthmus of Panama, and contains a highly diverse macro- and Gatun Formation. Based on screenwashing of sediments at a microfossil fauna (e.g., Woodring, 1957–1982; Collins et al., now inaccessible (i.e., covered by urban development) locality, 1996, 1999; Jackson and Budd, 1996; Jackson et al., 1999; Uhen he described the diversity, ecology, and biogeography of 14 et al., 2010; Hendy, 2013). Previous studies of the invertebrate chondrichthyan taxa. Most recently, Pimiento et al. (2010) fauna from the Gatun Formation indicate this area was a studied the natural history of the species Carcharocles shallow-water embayment that supported a neritic environment megalodon fromtheGatunFormationandproposedthatthey (Teranes et al., 1996). It has been postulated that this area was a used this area as a nursery habitat. Lastly, Pimiento et al. (2013) strait connecting the Pacific Ocean and the Caribbean Sea described the chondrichthyan fauna from the early Miocene (Coates and Obando, 1996), although subsequent workers have Culebra Formation and studied their paleoenvironmental setting suggested that any connection likely existed in a more distal and paleobiogeography. location (Collins et al., 1996, 1999; Teranes et al., 1996; Kirby Over the past 20 yr, the Gatun Formation localities have been et al., 2008). Regardless, all these studies show that the Gatun extensively quarried to extract sediment for construction marine faunas existed during a time of active transoceanic projects in the Colon region and contour surfaces for residential interchange and dispersal before the time of the final closure of developments. These quarries have both uncovered fresh the Central American Seaway (CAS). outcrops, but these are typically ephemeral (e.g., Gillette’s It is generally accepted that the final closure of the CAS took locality). Extraction activities have increased substantially in place about 3 million yr ago (Duque-Caro, 1990; Coates et al., recent years and numerous well-established outcrops are being 1992, 2003, 2004; Coates and Obando, 1996; Bartoli et al., completely removed or covered by urban development. We 2005). However, a number of recent studies suggest that the predict that many of the critical outcrops of the Gatun Formation shoaling of the Isthmus began during the middle–late Eocene will soon disappear. Given that the only systematic paleontology and likely resulted in the final closure during the middle paper on the chondrichthyans of the Gatun Formation was Miocene (Farris et al., 2011; Montes et al., 2012a, 2012b). In published 28 yr ago based on a single locality and collection any case, this closure was a key vicariant event for tropical method, we consider it important to recover and document new biotic evolution (Cronin and Dowsett, 1996) that resulted in the specimens from different sections before they are no longer separation of the once continuous marine distribution, and an available. Accordingly, we have collected and/or studied a total increase of habitat and biogeographic complexity (e.g., Wood- of 800 specimens from 15 localities from the Gatun Formation. ring, 1974; Rosen, 1975; Jackson and Budd, 1996; Aguilera et This material includes the original specimens described in al., 2011). Gillette (1984), new specimens collected by our team, and Fossil chondrichthyans from Panama are poorly known in the additional specimens in the Smithsonian National Museum of literature. The earliest record was reported by Blake (1862) who Natural History, Washington, D.C. 755 756 JOURNAL OF PALEONTOLOGY, V. 87, NO. 5, 2013 The purpose of this report is to revise and update the fossil Formation, Maryland (Tedford et al., 2004); Can´ımar Forma- record of the chondrichthyan fauna from the Gatun Formation. tion, Cuba (Iturralde-Vinent, 1969); Grand Bay Formation of Here, we review and describe all relevant known material. Carriacou; Grenadines, Lesser Antilles (Donovan and Gunter, Furthermore, based on our study of the fossil specimens and the 2001); and Rio Banano Formation, Costa Rica (Taylor, 1975; information available on extant related species, we evaluate the Laurito, 1999); 3) younger, early Pliocene assemblages include: diversity, taxonomic composition, paleobiogeography, paleo- Yorktown Formation, North Carolina (Ward and Bohaska, environment, and paleoecology of the chondrichthyan fauna of 2008); Bone Valley, Florida (Morgan, 1994; Tedford et al., the Gatun Formation. Results of this study enhance our 2004); Pisco Formation (upper part), Peru (De Muizon and understanding of the chondrichthyans of the ancient Neotropics DeVries, 1985); and Cubagua Formation, Venezuela (Aguilera, at a time when the Pacific Ocean and the Caribbean Sea were 2010). connected through Central America. With respect to the paleogeography, it has been postulated that the Gatun Formation was deposited within an archipelagic GEOLOGICAL SETTING strait connecting the Pacific Ocean and the Caribbean Sea The fossil chondrichthyan teeth described in this paper were (Coates and Obando, 1996). Subsequent workers indicated that collected from late Miocene marine sediments of the Gatun waters had shoaled to the point the strait was restricted (Collins Formation. This formation crops out in a broad area of northern et al., 1996, 1999). Most recently, it has been suggested that Panama (Fig. 1) extending along the northern shore of Lake contrary to an archipelago, a subaerial peninsula connected to Gatun, approximately 15 km northward to the Caribbean Sea, North America existed during this time (Kirby and MacFadden, and approximately 50 km east and west of Colon City, within 2005; Kirby et al., 2008). Regardless, all these studies show that the Panama Canal structural basin. In the study area, the gently a passageway existed between Central and South America, and dipping (58) Gatun Formation is thought to overlie unnamed it was located near to the deposition of sediments of the Gatun Cretaceous–Paleogene volcanic and volcaniclastic sediments or Formation. Therefore, the Gatun faunas existed during a time of the upper Oligocene sediments of the Caimito Formation active transoceanic interchange and dispersal (Collins et al., (Woodring, 1957–1982; Coates et al., 1992; Coates, 1999) 1996, 1999; Newkirk and Martin, 2009). (Fig. 2). With regard to the paleoecology of the Gatun Formation, The Gatun Formation, with a stratigraphic thickness of 500 m, foraminifera (Collins, 1996), invertebrates (e.g., Coates and consists of three informal members (Fig. 2) referred to as the lower, middle, and upper Gatun in most publications (e.g., Obando, 1996; Collins et al., 1996; Jackson et al., 1999; Hendy, Woodring, 1957–1982; Coates, 1999). Of relevance to this 2013) and fish otoliths (Aguilera and Rodrigues de Aguilera, report, most of the fossils collected during our study, and likely 1999) indicate a shallow-water marine shelf neritic environ- those of Gillette (1984),
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