The late Miocene elasmobranch assemblage from Cerro Colorado (Pisco Formation, Peru)
Introduction
The upper Neogene Pisco Formation is a shallow-marine sedimentary succession exposed along the southern coast of Peru from Pisco to Yauca (Dunbar et al., 1990, DeVries, 1998). Due to the abundance of fossil marine vertebrates, to their high diversity, and to the extraordinary quality of their preservation, the Pisco Formation is currently regarded as one of the most significant Cenozoic Fossil-Lagerstätten worldwide (Bianucci et al., 2016a, Bianucci et al., 2016b, Esperante et al., 2015, Gariboldi et al., 2015, Gioncada et al., 2016, Collareta et al., press). More than three decades of paleontological studies on the fossil content of the Pisco Formation led to a plethora of significant works dealing with the systematics, taphonomy, paleoecology, and phylogeny of various lineages of vertebrates, in particular birds and marine mammals (see e.g. Bianucci et al., 2016b and previous references theirein; Collareta et al., 2015, Bianucci et al., 2016b, Lambert et al., 2015, Lambert et al., in press; Stucchi et al., 2016, Gioncada et al., 2016, Collareta et al., press). On the contrary, the literature about the fossil sharks of the Pisco Formation is rather scarce, and most paleontological works only report the presence of teeth belonging to a few large-sized extinct taxa (e.g., Carcharocles megalodon or Cosmopolitodus hastalis) in their respective study areas. In his review of the geology of the coastline comprised between Marcona and Yauca, Ruegg (1968) recognized close similarities between the faunas of El Jahuay and Sacaco by identifying several species of diatoms, foraminifera and chondrichthyan fishes (including C. megalodon, Isurus sp., and Carcharhinus sp.). Later, Muizon and DeVries (1985) recognized six main vertebrate-bearing levels in the Pisco Formation (Cerro la Bruja (CLB), El Jauhay (ELJ), Aguada de Lomas (AGL), Montemar (MTM), Sacaco (SAO), and Sud Sacaco (SAS), listed in order of decreasing geological age); for each of these levels, Muizon and DeVries (1985) provided a list of the recognized taxa and briefly discussed the elasmobranch assemblages, reporting also on an undescribed lamniform species with weakly serrated teeth (somewhat intermediate between those of Cosmopolitodus hastalis and those of Carcharodon carcharias). Alván de la Cruz (2008) provided a preliminary characterization of the fossil selachians of a few sites near Ullujaya (Ocucaje area, Ica desert) where both the Pisco Formation and the underlying Chilcatay Formation are exposed. Ehret et al. (2009a) described an exceptionally well-preserved specimen (including 222 teeth and 45 vertebrae, as well as portions of the jaws) of Carcharodon sp., whereas a partial tooth belonging to the same species (embedded within a fragmentary mysticete dentary) was described by Ehret et al. (2009b); both these records come from the upper Miocene SAS horizon of the Sacaco basin. Based on the almost complete specimen studied by Ehret et al., 2009a, Ehret et al., 2012 instituted the species Carcharodon hubbelli, interpreted as representing the transition between the archaic Cosmopolitodus hastalis and the modern Carcharodon carcharias. Takakuwa (2014) described sixteen fossil teeth, referred to at least two individuals of “Isurus” hastalis, found in close contact with a balaenopterid whale skeleton from the upper Miocene site of Aguada de Lomas. Bianucci et al. (2016b) provided a detailed census of the fossil vertebrate assemblage of Cerro Colorado (a site located in the Ica desert), thus recognizing eight selachian families (based on the determination of more than 3500 teeth) and detailing the geographic and stratigraphic position of the most relevant shark remains. A similar approach was followed by Bianucci et al. (2016a) for characterizing the Messinian vertebrate assemblage of Cerro Los Quesos (Ica desert). Finally, Collareta et al. (in press) described a well-preserved juvenile specimen of Cosmopolitodus hastalis, featuring fish remains as stomach contents, from the late Miocene site of Cerro Yesera (Ica desert).
The present paper focuses on the Cerro Colorado tooth assemblage, preliminary recorded by Bianucci et al. (2016b), in order to: i) provide a more complete view of the Pisco Formation paleobiodiversity; and ii) define the taxonomic composition, the paleoecological relationships, and the paleobiological significance of this fossil elasmobranch assemblage.
Section snippets
Geological and stratigraphical setting
Located in southern Peru (Fig. 1), the (East) Pisco Basin is a 180 × 30 km elongated basin bounded to the east by Mesozoic igneous rocks of the Coastal Batholith (e.g. Cobbing, 1999) and to the west by Precambrian to Jurassic metamorphic, igneous, and sedimentary rocks of the Coastal Cordillera. The basin fill comprises, from bottom to top, the Eocene Caballas Formation and Paracas Group, the late Oligocene-middle Miocene Chilcatay Formation, and the late Miocene-Pliocene Pisco Formation (
Sampling methods
The fossil teeth here studied have been collected during several successive field trips in the last few years (2014–2015). These detailed investigations allowed us to recognize an extraordinarily rich shark tooth-bearing interval in the lower allomember of the Pisco Formation exposed at Cerro Colorado.
From a methodological point of view, two different sampling strategies have been followed: one consisting of a detailed (both formal and informal) sampling procedure, and one consisting of a
Systematic paleontology
Squatiniformes Buen, 1926
Squatinidae Bonaparte, 1838
Squatina Duméril, 1806
Squatina sp.
(Fig. 2G and H)
Referred specimens - Two partially eroded specimens (MUSM 3251).
Stratigraphic level - ST-low1.
Description - Specimens of Squatina sp. range from 6.7 to 6.9 mm in length and from 6.0 to 6.8 mm in width. Lingual face of the crown regularly smooth and convex. Main cusp lingually bent, triangular in shape, with a basal protuberance protruded towards the inside of the mouth. Cutting edges of the cusp
The preservation state
The preservation state of the elasmobranch teeth provides useful information about the diagenetic and/or postdiagenetic events (e.g. decay, fossilization, weathering exposure, wet-dry cracking) they were affected by. Litvinov (2007) reported on rootless and/or partially destroyed teeth which suffered long-lasting exposure on the ocean sea floor. Similar preservation patterns have been also recorded in time-averaged deposits by several authors (e.g. Boessenecker et al., 2014, Behrensmeyer, 1978
Conclusions
Changes in diversity in the three studied shark tooth-bearing levels allow to define the trophic and ecological dynamics of the elasmobranch communities of Cerro Colorado and to recognize their biological relationships with other marine vertebrate communities spread in the same basin.
The rich and diversified association coming from the ST-low1 deposits is largely dominated by C. brachyurus and characterized by a strong juvenile imprint.
The bathimetric distribution and ecological preferences of
Acknowledgments and funding
The authors wish to thank the following institutions: Dipartimento di Scienze della Terra, Università di Pisa (Italy); Museo di Storia Naturale, Università di Pisa (Italy); Museo Zoologico “La Specola”, Università di Firenze (Italy); Museo Civico “A. Doria”, Genua (Italy); Museo Zoologico “P. Doderlein”, Università di Palermo (Italy). We particularly thank Rafael M. Varas-Malca and W. Aguirre, who provided very helpful assistance both in the field and during our stays at the MUSM. Comments by
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