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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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The present paper describes a disarticulated skull of Allqokirus australis Marshall & Muizon, 1988, a basal sparassodont (Metatheria, Mammalia) from the early Palaeocene (c. 65 Ma.) of Tiupampa (Bolivia). The specimen includes the rostrum and palate with right premaxilla, both maxillae, left lacrimal, palatines and most upper teeth. The second largest element includes the frontals, the left squamosal, the parietals, the supraoccipital, the basisphenoid, the presphenoid, the alisphenoid, and part of the pterygoids. The nasals, basioccipital and exoccipitals are missing. Other elements are the left petrosal, the right jugal and squamosal, and both dentaries. The elements of the specimen allow for a good reconstruction of the skull, which is thoroughly described and compared to that of other sparassodonts and to the Tiupampa pucadelphyids, Pucadelphys and Andinodelphys. The dental morphology of Allqokirus australis is extremely similar to that of Patene simpsoni from the early Eocene of Itaboraí (Brazil) and presents distinct (although incipient) carnivorous adaptations. Furthermore, some characters of the ear region (e.g. medial process of the squamosal, deep groove for the internal carotid artery at the ventral apex of the petrosal) are also present in most other sparassodonts and in the pucadelphyids from the same locality. A parsimony analysis performed on the basis of a data matrix of 364 characters and 38 taxa placed Allqokirus in a sparassodont clade (the Mayulestidae) that also included Mayulestes and Patene. This family constitutes the sister group of all other sparassodonts. Our analysis also retrieved a large clade composed of the sparassodonts and the pucadelphyids, formally named Pucadelphyda n. superord. This superorder represents the large metatherian carnivorous radiation of the Tertiary of South America, which is first known at Tiupampa, and which started to diversify probably slightly earlier, during the late Cretacous in South America. So far, no representative of Pucadelphyda has been discovered in North America. At Tiupampa, Allqokirus and Mayulestes are the largest metatherians of the fauna and they fill the predaceous mammalian ecological niche. They are the earliest representatives of Sparassodonta, a successful metatherian carnivorous radiation which persisted in South America until the late Pliocene, i.e., during more than 63 Ma.

Alcidedorbignya inopinata Muizon & Marshall, 1987 is a basal pantodont (Placentalia, Mammalia) of small body size, from the early Palaeocene of the Santa Lucia Formation at Tiupampa, Bolivia. Tiupampa is the type locality for the Tiupampan, a South American Land Mammal age (SALMA), which is assigned an age equivalent to the basal Torrejonian 1 of North America (c. 65 Ma). Alcidedorbignya is known by exceptionally preserved specimens, which are described here. The two major specimens are an almost complete skeleton (MHNC 8372) and a partial skull (MHNC 8399), the former representing one of the best-preserved fossil placentals from the early Palaeocene and probably the oldest placental skeleton that is so completely known. These specimens are also the first eutherian skulls and skeleton ever discovered at Tiupampa, a locality which has yielded numerous metatherian skulls and partial skeletons. The remarkable preservation of the two skulls allows a detailed description of the cranial anatomy with well-identified sutures and foramina, including those of the auditory region. Through CT scanning of the skulls, the arterial and venous circulation pattern in the basicranium as well as the bony labyrinth of the inner ear were tentatively reconstructed. A thorough description of the postcranial skeleton of MHNC 8372 is also provided. Among pantodonts, Alcidedorbignya presents the closest morphological similarities with Pantolambda, the oldest and earliest diverging North American pantodont (known by skulls and skeletons), from the late early Palaeocene (Torrejonian 2 and 3) of New Mexico. Alcidedorbignya is one-third the size, much more gracile, and clearly exhibits more plesiomorphic features than Pantolambda. It is also at least 3 Ma older. Interesting similarities are also observed between the skull of Alcidedorbignya and several "condylarths", such as Maiorana, Baioconodon, Arctocyon, and Arctocyonides. The basicranium of Alcidedorbignya is also similar to that of some extant afrotheres (e.g., Tenrec) or Lipotyphla (e.g., 398GEODIVERSITAS • 2015 • 37 (4) Muizon C. de et al. Solenodon), but most of these similarities may represent placental symplesiomorphies. In fact, the cranial anatomy of Alcidedorbignya, beyond the simple thorough description of a basal pantodont, sheds light on the cranial anatomy of placentals from the earliest Paleocene, previously unknown in this detail. The postcranial skeleton of MHNC 8372 together with some isolated specimens referred to A. inopinata, is compared to adequate morphofunctional models (e.g., Solenodon, Dendrohyrax, Sciurus), which indicates that it was a moderately agile, plantigrade, generalized terrestrial mammal with good climbing ability (scansorial) and occasionally capable of standing in a bipedal position. The scutiform ungual phalanges were probably bearing nail-like hooves (or primate-like nails) and because of the absence of claws, fossorial habits are unlikely.A parsimony analysis of a data matrix including 72 taxa and 426 characters (cranial and postcranial...

Thalassocnus is a genus of "ground sloths" known from Neogene deposits, for the great majority of specimens, of the Pisco Formation (Peru). Five species are recognized, their description being currently restricted, for the most part, to the skull, mandible, and dentition. The bones of the forelimb are here described, and compared among the species of Thalassocnus and to other pilosans. The main characteristics of the forelimb of Thalassocnus relative to other sloths are the shortness of the humerus and radius, and the specialized digits. Moreover, the late species of the genus are characterized by the development of the pronator ridge of the radius, stoutness of the ulna, widening of the proximal carpal row, and shortening of the metacarpals. Analogies with extant tetrapods are proposed in order to infer plausible aquatic functions of the forelimb of Thalassocnus. In addition to paddling, it is argued that the forelimb of Thalassocnus was involved in bottom-walking, a function similarly found in extant sirenians.However, the function of the forelimb of Thalassocnus differs drastically from that of the latter, since it was likely involved in an activity related to obtaining food such as uprooting seagrass rhizomes.

Odontocetes (toothed whales) rely upon echoes of their own vocalizations to navigate and find prey underwater [1]. This sensory adaptation, known as echolocation, operates most effectively when using high frequencies, and odontocetes are rivaled only by bats in their ability to perceive ultrasonic sound greater than 100 kHz [2]. Although features indicative of ultrasonic hearing are present in the oldest known odontocetes [3], the significance of this finding is limited by the methods employed and taxa sampled. In this report, we describe a new xenorophid whale (Echovenator sandersi, gen. et sp. nov.) from the Oligocene of South Carolina that, as a member of the most basal clade of odontocetes, sheds considerable light on the evolution of ultrasonic hearing. By placing high-resolution CT data from Echovenator sandersi, 2 hippos, and 23 fossil and extant whales in a phylogenetic context, we conclude that ultrasonic hearing, albeit in a less specialized form, evolved at the base of the odontocete radiation. Contrary to the hypothesis that odontocetes evolved from low-frequency specialists [4], we find evidence that stem cetaceans, the archaeocetes, were more sensitive to high-frequency sound than their terrestrial ancestors. This indicates that selection for high-frequency hearing predates the emergence of Odontoceti and the evolution of echolocation.

A major gap in our knowledge of the evolution of marsupial mammals concerns the Paleogene of the northern continents, a critical time and place to link the early history of metatherians in Asia and North America with the more recent diversification in South America and Australia. We studied new exceptionally well-preserved partial skeletons of the Early Oligocene fossil Herpetotherium from the White River Formation in Wyoming, which allowed us to test the relationships of this taxon and examine its adaptations. Herpetotheriidae, with a fossil record extending from the Cretaceous to the Miocene, has traditionally been allied with opossums (Didelphidae) based on fragmentary material, mainly dentitions. Analysis of the new material reveals that several aspects of the cranial and postcranial anatomy, some of which suggests a terrestrial lifestyle, distinguish Herpetotherium from opossums. We found that Herpetotherium is the sister group to the crown group Marsupialia and is not a stem didelphid. Combination of the new palaeontological data with molecular divergence estimates, suggests the presence of a long undocumented gap in the fossil record of opossums extending some 45Myr from the Early Miocene to the Cretaceous.