Did the giant extinct shark Carcharocles megalodon target small prey? Bite marks on marine mammal remains from the late Miocene of Peru
Graphical abstract
Introduction
The extinct megatooth shark species Carcharocles megalodon (Agassiz, 1843) (Elasmobranchii, Lamniformes, Otodontidae) is known by large serrated teeth and vertebrae from Neogene marine and brackish-transitional deposits worldwide, ranging in age from the late early Miocene (Burdigalian) to the late Pliocene (e.g., Cappetta, 2012, Pimiento and Clements, 2014, Carrillo-Briceño et al., 2015, Carrillo-Briceño et al., 2016). Reaching an estimated total body length of more than 16 m (Gottfried et al., 1996, Pimiento et al., 2010), C. megalodon is widely regarded as an apex predator that likely filled the top trophic levels of the global ocean (e.g., Aguilera et al., 2008, Ehret, 2010). Despite C. megalodon being interpreted as a whale-eating predator (e.g., Compagno, 1990, Purdy, 1996, Wroe et al., 2008, Ehret, 2010), and its remains being common in Neogene deposits, little direct evidence for the trophic ecology of this giant shark arose from the fossil record to date. This scarce fossil record includes: (1) several large whale bones (mainly vertebrae and forelimb bones) from the Pliocene Yorktown Formation (USA) bearing bite marks made by very large serrated teeth (Purdy, 1996); (2) one cetacean vertebra probably from the Burdigalian to ?early Langhian Cantaure Formation (Venezuela) pierced by a tooth of C. megalodon (Aguilera et al., 2008, Carrillo-Briceño et al., 2016); and (3) one vertebral centrum of a small-sized (ca. 6 m long) whale from the Miocene Chesapeake group of Maryland (USA) presenting a partially healed compression fracture, tentatively attributed to failed predation by C. megalodon (Godfrey and Altman, 2005). In this paper, we describe new shark bite marks attributed to C. megalodon and affecting cetacean and pinniped bones from the late Miocene deposits of the Pisco Formation. The latter is a shallow-marine sedimentary unit exposed along the southern coast of Peru which has recently yielded multiple clues of trophic interactions between marine vertebrates (Ehret et al., 2009b, Collareta et al., 2015, Collareta et al., 2016, Lambert et al., 2015) besides a remarkable fossil record of sharks (Alván de la Cruz et al., 2006, Alván de la Cruz, 2008, Ehret et al., 2009a, Ehret et al., 2009b, Ehret et al., 2012, Altamirano-Sierra, 2012, Takakuwa, 2014, Landini et al., 2017, Collareta et al., 2016). To our knowledge, the trace fossils described herein represent the first record of C. megalodon bite marks from the Southern Hemisphere, and the first instance when a prey (or scavenging item) of C. megalodon is identified at the species level (as Piscobalaena nana, a small-sized cetotheriid baleen whale). This fossil occurrence stimulates various inferences about the trophic habits of C. megalodon and its extinction.
Section snippets
Geographical, geological, and palaeontological context
The Hueso Blanco study area is located in the valley of Aguada de Lomas (indicative geographic coordinates: S 15°28′50″; W 74°48′17″), Lomas area of the Sacaco Basin, where a 287-m-thick succession of upper Miocene beds of the Pisco Formation is exposed (Brand et al., 2011) (Fig. 1). The sediment package exposed at Hueso Blanco (bed LM 10 in Brand et al., 2011) is about 15 m thick and consists mainly of sparsely to moderately bioturbated, well-sorted, fine- to medium-grained sandstones (Electronic
Identification of the bitten mammalian remains
MUSM 3239 (Figs. 2a–b and S2) is a fragment of a mysticete left mandible belonging to a Cetotheriidae s.s., owing to the presence of an angular process protruding posteriorly beyond the edge of the mandibular condyle, the latter being oriented obliquely with respect to the long axis of the bone (El Adli et al., 2014, Gol'din et al., 2014). The shape and size of MUSM 3239 are strikingly reminiscent of Piscobalaena nana, known from upper Miocene deposits of Peru (Pilleri and Siber, 1989, Bouetel
Active predation or scavenging
Except for a few cases in which a shark attack interpretation is favoured due to bite marks preserved on almost complete prey skeletons (Cigala Fulgosi, 1990, Bianucci et al., 2010b, Bianucci and Gingerich, 2011) or to bone healing around a tooth-related wound (Kallal et al., 2012), it is virtually impossible to discriminate between active predation and scavenging when dealing with fossil specimens. Considered a modern analogous of C. megalodon (Purdy, 1996, Ehret, 2010, Pimiento et al., 2010),
Conclusions
Bite marks attributed to the megatooth shark Carcharocles megalodon have been described on fossil cetacean and pinniped bones collected from upper Miocene (about 7.5–7 Ma) deposits of the Pisco Formation exposed at Hueso Blanco (Aguada de Lomas valley, Sacaco basin, southern Peru). One of these bitten bones has been determined as a partial mandible of a small-sized cetotheriid (Piscobalaena nana) which was consumed by a mature individual of C. megalodon; the other bitten mammal bones here
Acknowledgments and funding
The authors wish to thank R. Salas-Gismondi and A. Altamirano-Sierra for fruitful discussions on the palaeoecology of the fossil sharks of the Pisco Formation, W. Aguirre for valuable field and laboratory assistance, and A. Gennari for the life reconstruction of Carcharocles megalodon and Piscobalaena nana in Fig. 3. Comments by the reviewers (O. Aguilera and J. Carrillo-Briceño) greatly improved the quality of this paper. We are also grateful to T. Corrège and T. J. Algeo for their valuable
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