The modern giant sperm whale Physeter macrocephalus, one of the largest known predators, preys upon cephalopods at great depths. Lacking a functional upper dentition, it relies on suction for catching its prey; in contrast, several smaller Miocene sperm whales (Physeteroidea) have been interpreted as raptorial (versus suction) feeders, analogous to the modern killer whale Orcinus orca. Whereas very large physeteroid teeth have been discovered in various Miocene localities, associated diagnostic cranial remains have not been found so far. Here we report the discovery of a new giant sperm whale from the Middle Miocene of Peru (approximately 12-13 million years ago), Leviathan melvillei, described on the basis of a skull with teeth and mandible. With a 3-m-long head, very large upper and lower teeth (maximum diameter and length of 12 cm and greater than 36 cm, respectively), robust jaws and a temporal fossa considerably larger than in Physeter, this stem physeteroid represents one of the largest raptorial predators and, to our knowledge, the biggest tetrapod bite ever found. The appearance of gigantic raptorial sperm whales in the fossil record coincides with a phase of diversification and size-range increase of the baleen-bearing mysticetes in the Miocene. We propose that Leviathan fed mostly on high-energy content medium-size baleen whales. As a top predator, together with the contemporaneous giant shark Carcharocles megalodon, it probably had a profound impact on the structuring of Miocene marine communities. The development of a vast supracranial basin in Leviathan, extending on the rostrum as in Physeter, might indicate the presence of an enlarged spermaceti organ in the former that is not associated with deep diving or obligatory suction feeding.

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Eight skulls of beaked whales (Cetacea, Odontoceti, Ziphiidae), in six cases associated with elements of the mandible, were collected from a limited area (about 1.5 km 2 )and roughly from the same stratigraphic horizon at Cerro Colorado, 35 km south-south-west of the city of Ica (Peru), where the late Middle Miocene basal strata of the Pisco Formation crop out. They represent the highest concentration reported of fossil Ziphiidae. These finely preserved Cerro Colorado fossils are described and assigned to a new species Messapicetus gregarius, together with another specimen collected from sediments of the same age at Cerro la Bruja (33 km south-east to Cerro Colorado). Messapicetus gregarius shares with M. longirostris Bianucci, Landini and Varola, 1992 (Tortonian of Italy), an extremely elongated rostrum, but is clearly different from the Italian species in the more distinct maxillary tubercle and prominential notch, the more robust premaxillary crest, and the abrupt ventrolateral descent of the medial margin of the maxilla from the vertex. A parsimony analysis reveals that Messapicetus belongs to a basal clade, which includes other ziphiids with a dorsally closed mesorostral groove and prenarial basin. The high concentration of specimens belonging to the same species (some of them tentatively identified as adult males and females), combined with the presence of a calf, supports the hypothesis of site fidelity; these cetaceans might have lived in a limited region for a long period for both breeding and feeding. Besides the eight specimens of M. gregarius, strata at Cerro Colorado include many other cetacean remains (with several specimens of the pontoporiid Brachydelphis including a foetus), pinnipeds, turtles, fishes, and birds.

The three extant sperm whale species, the giant Physeter macrocephalus and the diminutive Kogia species, are relict members of the superfamily Physeteroidea, whose evolutionary history is traced back to the late Oligocene. Although well-preserved and diagnostic cranial remains are relatively scarce, the physeteroid fossil record reveals a considerable degree of morphological disparity (especially during the Miocene), suggesting that sperm whales occupied a broader range of ecological niches in the past. Here, we provide detailed descriptions and a (re)analysis of several new and established middle-late Miocene stem physeteroids from the Pisco Formation of southern Peru. In particular, we (1) further describe the holotype of Acrophyseter deinodon from the latest Tortonian-Messinian of Sud-Sacaco, with new information on previously unpublished portions of the skull (including the ear bones) and a discussion of its masticatory musculature; (2) diagnose a new species of Acrophyseter, A. robustus sp. nov., based on a finely preserved skull with some associated cranial remains from the late Serravallian-Tortonian (> 9.2 Ma) of Cerro la Bruja; (3) provisionally refer a skull from Cerro los Quesos, which has been radiometrically dated to the Messinian (6.9-6.7 Ma), to Acrophyseter sp.; and (4) further describe the skull of the giant raptorial sperm whale Livyatan melvillei from Cerro Colorado, recently re-dated to the Tortonian (9.9-8.9 Ma) based on the associated diatom fauna and Ar/Ar dating. A phylogenetic analysis based on 53 characters and 21 physeteroid species confirms the monophyly of Acrophyseter and groups this genus with the larger, middle to late Miocene macroraptorial stem physeteroids Brygmophyseter and Zygophyseter. With its unique supracranial basin morphology, Livyatan forms a separate, more crownward stem physeteroid lineage. Combined with biostratigraphic information, our cladistic hypothesis allows us to discuss the time of origin of the main physeteroid clades, as well as trends in the evolution of their body size, dentition, temporal fossa and supracranial basin

Although combined molecular and morphological analyses point to a late middle Eocene (38-39 million years ago) origin for the clade Neoceti (Odontoceti, echolocating toothed whales plus Mysticeti, baleen whales, and relatives), the oldest known mysticete fossil dates from the latest Eocene (about 34 million years ago) of Antarctica [1, 2]. Considering that the latter is not the most stemward mysticete in recent phylogenies and that Oligocene toothed mysticetes display a broad morphological disparity most likely corresponding to contrasted ecological niches, the origin of mysticetes from a basilosaurid ancestor and its drivers are currently poorly understood [1, 3-8]. Based on an articulated cetacean skeleton from the early late Eocene (Priabonian, around 36.4 million years ago) of the Pisco Basin, Peru, we describe a new archaic tooth-bearing mysticete, Mystacodon selenensis gen. et sp. nov. Being the geologically oldest neocete (crown group cetacean) and the earliest mysticete to branch off described so far, the new taxon is interpreted as morphologically intermediate between basilosaurids and later toothed mysticetes, providing thus crucial information about the anatomy of the skull, forelimb, and innominate at these critical initial stages of mysticete evolution. Major changes in the morphology of the oral apparatus (including tooth wear) and flipper compared to basilosaurids suggest that suction and possibly benthic feeding represented key, early ecological traits accompanying the emergence of modern filter-feeding baleen whales' ancestors.