The Giant Bite of a New Raptorial Sperm Whale from the Miocene Epoch of Peru

Home , Aulophyseter, Livyatan, Orycterocetus, Physeter, Physeteroidea, Praekogia

Vol 466 | 1 July 2010 | doi:10.1038/nature09067 LETTERS

The giant bite of a new raptorial sperm whale from the Miocene epoch of Peru

Olivier Lambert1*{, Giovanni Bianucci2*, Klaas Post3, Christian de Muizon4, Rodolfo Salas-Gismondi5, Mario Urbina5 & Jelle Reumer3,6

The modern giant sperm whale Physeter macrocephalus, one of the the Miocene of the Pisco basin, Peru, with an associated set of very largest known predators, preys upon cephalopods at great large teeth and mandible. depths1,2. Lacking a functional upper dentition, it relies on suction 3 Cetacea for catching its prey ; in contrast, several smaller Miocene sperm Neoceti whales (Physeteroidea) have been interpreted as raptorial (versus 4,5 Odontoceti suction) feeders , analogous to the modern killer whale Orcinus Physeteroidea orca. Whereas very large physeteroid teeth have been discovered in Leviathan melvillei gen. et sp. nov. various Miocene localities, associated diagnostic cranial remains have not been found so far6–8. Here we report the discovery of a Etymology. From Hebrew ‘Livyatan’ (‘Leviathan’ in Latin); name new giant sperm whale from the Middle Miocene of Peru (approxi- applied to large marine monsters in popular and mythological stories. mately 12–13 million years ago), Leviathan melvillei, described on Species is dedicated to novelist Herman Melville (1819–1891). the basis of a skull with teeth and mandible. With a 3-m-long head, Holotype. Museo de Historia Natural, Universidad Nacional Mayor very large upper and lower teeth (maximum diameter and length de San Marcos (MUSM) 1676, 75% complete skull, including rostrum of 12 cm and greater than 36 cm, respectively), robust jaws and a with damaged maxillary teeth embedded and the right side of the temporal fossa considerably larger than in Physeter, this stem cranium with dentaries and ten isolated teeth associated (Fig. 1, Sup- physeteroid represents one of the largest raptorial predators plementary Figs 1–5 and Supplementary Tables 1–6). and, to our knowledge, the biggest tetrapod bite ever found. The Locality. Cerro Colorado, Pisco-Ica desert, 35 km south-southwest appearance of gigantic raptorial sperm whales in the fossil record of Ica, southern coast of Peru, 14u 209 13.499 S, 75u 549 25.299 W. coincides with a phase of diversification and size-range increase of Horizon. Lowest beds of Pisco Formation, late Middle Miocene, the baleen-bearing mysticetes in the Miocene. We propose that Serravallian stage, about 12–13 million years ago (see Supplemen- Leviathan fed mostly on high-energy content medium-size baleen tary Information). whales. As a top predator, together with the contemporaneous Diagnosis. Large stem physeteroid (Fig. 2) defined by two unequi- giant shark Carcharocles megalodon, it probably had a profound vocal autapomorphies: great anterior expansion of the premaxilla on impact on the structuring of Miocene marine communities. The the wide rostrum, reaching the lateral margin of the latter; and maxi- development of a vast supracranial basin in Leviathan, extending mum tooth diameter greater than 10 cm for most of the dentition. It on the rostrum as in Physeter, might indicate the presence of an further differs from all other physeteroids except Physeter in the enlarged spermaceti organ in the former that is not associated with elongated supracranial basin extending along the whole length of deep diving or obligatory suction feeding. the rostrum, and from Kogia, Physeter and some related fossil forms With adult males reaching a body length of 18.3 m, the modern in the retention of a functional upper dentition and enamel on the sperm whale Physeter macrocephalus is one of the largest macro- teeth. predators ever found. It preys primarily on squid, which it usually With a skull length of about 3 m and a skull width of 190 cm, hunts at great depths1,2. Lacking functional upper dentition, Physeter Leviathan melvillei is the largest known fossil physeteroid. Body length ingests and swallows its prey by suction3, whereas its lower teeth are is estimated at 13.5–17.5 m (see Supplementary Information), in the mostly used in social interactions9,10. However, a raptorial feeding range of adult male Physeter1. The rostrum is short, slightly longer than behaviour analogous to that of the modern killer whale Orcinus orca the cranium, with a wide base and thick outer margins laterally convex has been proposed for several Miocene fossil sperm whales bearing in the anterior portion (Fig. 1). The laterally expanded premaxillae functional upper teeth4,5,11. All these taxa are considerably smaller than form the floor of the distal part of the elongated supracranial basin. Physeter. For more than a century, isolated large physeteroid teeth This anterior extension of the basin, only observed as such in Physeter, with a diameter of 7–9 cm and a maximum length of 27 cm, found is an indication of the presence of a large spermaceti organ, which, as in in various Neogene localities worldwide, indicated that unusually Physeter, would have reached the anterior tip of the rostrum. The deep large sperm whales roamed the oceans in the past6,7. But except for neurocranial portion of the supracranial basin is margined laterally by fragments of rostrum and mandible from the Miocene of California8, the maxilla, which leaves the posterior wall of the basin uncovered. no known fossil sperm whale could match the size of these teeth, Eleven deep alveoli are found in the dentary, and nine in the precluding any hypothesis on the diet and feeding strategy of these maxilla. Except for some specimens of the extant dwarf and pygmy animals. Here we report the discovery of a giant physeteroid skull from sperm whales Kogia spp.12, this is the lowest dentary tooth count in a

1De´partement de Paleóntologie, Institut Royal des Sciences Naturelles de Belgique, Brussels 1000, Belgium. 2Dipartimento di Scienze della Terra, Universita` di Pisa, Pisa 56126, Italy. 3Natuurhistorisch Museum Rotterdam, Rotterdam 3001 KL, The Netherlands. 4De´partement Histoire de la Terre, Museúm National d’Histoire Naturelle, Paris 75005, France. 5Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima 14, Peru. 6Department of Geosciences, Utrecht University, Utrecht 3508TA, The Netherlands. {Present address: De´partement Histoire de la Terre, Museúm National d’Histoire Naturelle, Paris 75005, France. *These authors contributed equally to this work. 105 ©2010 Macmillan Publishers Limited. All rights reserved LETTERS NATURE | Vol 466 | 1 July 2010

abcPmx

Pmx 1 m Mx

Vo Iof An Vo Mx La La Fr Paf Pmf Fg Mx Msb Fr Mt Pmx Fr Tf Rn Tf Zp Mx Sq As Pty

efghi d 10 cm Pmx Wf Fr Sb Tf Mx Or La Mx Ex Sq

Mc Wf

Figure 1 | Skull, mandible and tooth morphology of the holotype of L. Mc, mandibular condyle; Mf, mandibular foramen; Mg, mesorostral groove; melvillei MUSM 1676. a–d, Schematic drawings of skull and mandible. Skull Msb, margin supracranial basin; Mt, mesethmoid; Mx, maxilla; Or, orbit; in dorsal (a) and ventral (b) view, mandible in dorsal view (c), skull and Paf, palatine foramen; Pmf, premaxillary foramen; Pmx, premaxilla; Pty, mandible in right lateral view (d). e–g, Right lower teeth in labial view. posterior process tympanic; Rn, right bony naris; Sb, supracranial basin; Sq, h, i, Teeth of modern sperm whale Physeter (h) and killer whale Orcinus (i) for squamosal; Tf, temporal fossa; Vo, vomer; Wf, wear facet; Zp, zygomatic comparison. An, antorbital notch; As, alisphenoid; Ex, exoccipital; process squamosal. Fg, frontal groove; Fr, frontal; Iof, infraorbital foramen; La, lacrimal; physeteroid. Together with the premaxilla and the vomer, the maxilla tooth-bearing portion of the dentary and a vast temporal fossa. This reaches the apex of the rostrum; no premaxillary alveoli are present. is indicative of feeding by means of raptorial predation for Leviathan,as The maxillary alveolar groove is offset internally from the lateral in several other, much smaller, Miocene physeteroids, but strongly margin of the rostrum, a condition differing from other fossil phy- contrasts with the suction feeding adaptations of Physeter. seteroids. Only partial roots of maxillary teeth were preserved in situ, This sperm whale could firmly hold large prey with its interlocking but several lower teeth are complete. With a maximum diameter teeth, inflict deep wounds and tear large pieces from the body of the ranging from 8.1 to 12.1 cm and a total length of more than victim, as performed on a smaller scale by the delphinid Orcinus, the 36.2 cm, these teeth are much larger than the largest recorded teeth main modern meat-eating cetacean15,16. With a maximum body size of Physeter (25 cm1; Fig. 1h). Combined with the length of the jaws, under 9 m and a skull length of about 1 m, Orcinus has been reported this represents one of the largest bites for vertebrates and certainly the occasionally to cooperatively attack and kill most of the large baleen biggest bite for tetrapods. The enamelled crown is low with a small whale species and sperm whales17,18, as well as smaller odontocetes, diameter. The robust root is covered with a thick layer of cement, pinnipeds, fish, seabirds and cephalopods1. typical of continuously growing sperm whale teeth13. Apical lower Besides small-to-medium size odontocetes, the most common teeth are more recurved than the nearly cylindrical and massive other marine mammals in Cerro Colorado are mysticetes (more than 20 teeth. Deep vertical wear facets on either the distolabial or mesiolabial skeletons of a new cetotheriid with a body length approaching 10 m surface of the dentary teeth indicate that the upper and lower teeth have been found). A similarly large number of 5–13-m-long mys- occluded for a long distance, suggesting an important shearing ticetes has been described in other Miocene localities of the Pisco component during the bite. To accommodate the large roots, the Formation, related to high local productivity19. More generally, alveolar-bearing part of the mandible is deep and wide. The anterior Serravallian to Tortonian stages correspond to a peak of diversity part of the rostrum is upturned, and the anterior upper and lower for cetaceans20,21, and particularly for mysticetes22 (Fig. 4a and Sup- teeth project forwards at an angle of approximately 45u. plementary Information). The enormous temporal fossa is much longer anteroposteriorly With their large size and robust jaws, Leviathan adults were surely than the orbit area of the skull, which differs from the small fossa of free from predation. However, besides being an antipredator strategy Physeter (Fig. 3). The wide space between the zygomatic process of (large sharks are abundant in Cerro Colorado) and/or having thermal squamosal and the medial surface of the temporal fossa indicates that advantages, when associated with an efficient raptorial feeding appar- the fossa could accommodate a large volume of musculus temporalis. atus gigantism allows preying upon larger animals23,24. A correlation There is a short and wide rostrum allowing a more powerful bite by between the emergence of the giant shark Carcharocles megalodon and the anterior teeth and better resistance to lateral movements of the the Miocene diversification of the mysticetes has already been pro- struggling prey14, procumbent anterior teeth for grasping voluminous posed22,25. Considering the distinct increase in the range of sizes for prey with moderately convex body surfaces, very robust upper and mysticetes since the Langhian stage, progressively reaching greater lower teeth deeply embedded in the jaw bones, a deep and wide maximum sizes (Fig. 4b and Supplementary Information), the record 106 ©2010 Macmillan Publishers Limited. All rights reserved NATURE | Vol 466 | 1 July 2010 LETTERS

Zygorhiza† a 30

Agorophius† 25 Eudelphis†

Acrophyseter† 20 Zygophyseter† 15 Brygmophyseter†

Genera Leviathan†

10 Stem Physeteroidea Stem Placoziphius† ?

Orycterocetus† 5

Physeterula†

Aulophyseter† hyseteroidea 35 30 25 20 15 10 5 0 P Time (Myr ago) Physeter

Physeteridae b 350

Aprixokogia† 30 300 Thalassocetus† ?

Scaphokogia† 250 Kogiidae Praekogia† ? 200 Kogia 13.5 150 Figure 2 | Phylogeny of physeteroids illustrating the relationships of 10.5 Leviathan with other stem physeteroids, physeterids and kogiids. A 9.5 9.2 8.1 maximum parsimony cladogram obtained with a PAUP (phylogenetic Bizygomatic width (cm) 100 analysis using parsimony) branch-and-bound search, using a data set 6.6 5.2 6.1 modified from previous works. The consistency (CI) and retention (RI) 4.3 indices of the single shortest tree are 0.61 and 0.65, respectively. Schematic 50 3.3 5.2 4.1 skulls illustrate the presence or absence of functional upper dentition in each 3.2 3.2 3.2 3.6 2.4 taxon. Question marks indicate unknown condition. The daggers indicate 1.6 2.1 extinct species. See Supplementary Information for further details. 35 30 25 20 15 10 5 0 Time (Myr ago) of two giant marine predators, C. megalodon and L. melvillei, in the Figure 4 | Evolution of the diversity and size of mysticetes from Oligocene same mysticete-rich Serravallian locality might be relevant. Fun- to present. a, Plot of the number of mysticete genera for each stage against ctionally able to feed on other groups of marine animals, including time. b, Plot of the bizygomatic width (in centimetres) of the skull of large fish, pinnipeds and odontocetes, Leviathan may have predomi- mysticete specimens for each stage against time. The numbers on the graph nantly preyed on higher-energy content medium-size mysticetes, indicate total body length estimates (in metres) for maximum and minimum which would have provided the large amount of fat required to fulfil bizygomatic width for each stage (see Supplementary Information for data and equation). The thick vertical line corresponds to the age of the sediments in which Leviathan was discovered. Pliocene is considered as a single time interval. Myr, millions of years. a the high caloric demands of this huge endothermic aquatic predator (for comments on energetic value of the prey species of Orcinus, see ref. 26). Furthermore, large physeteroid teeth found in other Miocene localities worldwide6,7 indicate that giant raptorial sperm whales occupied a top predator position in various marine regions during the Miocene, a role now mostly taken by Orcinus. As such, the b c appearance of these sperm whales probably had a profound impact 1 m on the structuring of Middle to Late Miocene marine communities and food chains, similar to the impact of Orcinus15,22,27. Although they bear a notably similar long supracranial basin, the modern physeterid Physeter and the stem physeteroid Leviathan are not closely related and the latter seems to have occupied a com- pletely different ecological niche. This information should be taken into account during future discussion of the potential function(s) (buoyancy regulation, echolocation, acoustic stunning of prey, Figure 3 | Comparison of the outline of the skull and mandible of Leviathan acoustic display, battering ram for intraspecific aggressions) and with modern giant sperm whale Physeter and killer whale Orcinus.a–c, Skull evolutionary history of the spermaceti organ, which fills the supra- 28,29 and mandible of Leviathan (a), Physeter (b) and Orcinus (c), in right lateral cranial basin in Physeter . We propose that the enlarged basin was view. Note the large size and proportionally large temporal fossa and teeth of not associated with deep diving, obligate suction feeding or teutho- Leviathan. All outlines are to the same scale. phagy in Leviathan. 107 ©2010 Macmillan Publishers Limited. All rights reserved LETTERS NATURE | Vol 466 | 1 July 2010

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CORRIGENDUM doi:10.1038/nature09381 The giant bite of a new raptorial sperm whale from the Miocene epoch of Peru Olivier Lambert, Giovanni Bianucci, Klaas Post, Christian de Muizon, Rodolfo Salas-Gismondi, Mario Urbina & Jelle Reumer

Nature 466, 105–108 (2010) The genus name Leviathan, proposed in this Letter for a new fossil physeteroid from the Miocene of Peru, is preoccupied by Leviathan Koch, 1841 (ref. 1), a junior subjective synonym of Mammut Blumenbach, 1799 (ref. 2). We propose here a replacement name Livyatan gen. nov. The type species is placed in this genus to form the binomial Livyatan melvillei. The diagnosis and content of the new genus follow our Letter. ‘Livyatan’ is a Hebrew name applied to large marine monsters in popular and mythological stories. We thank M. P. Taylor and D. Yanega for bringing this to our attention.

1. Koch, A. Description of the Missourium, or Missouri Leviathan 2nd edn (Prentice & Weissinger, 1841). 2. Blumenbach, J. F. Handbuch der Naturgeschichte, 6. Auflage (Go¨ttingen, 1799).