A heavyweight early whale pushes the boundaries of vertebrate morphology.

The fossil record of cetaceans documents how terrestrial animals acquired extreme adaptations and transitioned to a fully aquatic lifestyle1,2. In whales, this is associated with a substantial increase in maximum body size. Although an elongate body was acquired early in cetacean evolution3, the maximum body mass of baleen whales reflects a recent diversification that culminated in the blue whale4. More generally, hitherto known gigantism among aquatic tetrapods evolved within pelagic, active swimmers. Here we describe Perucetus colossus-a basilosaurid whale from the middle Eocene epoch of Peru. It displays, to our knowledge, the highest degree of bone mass increase known to date, an adaptation associated with shallow diving5. The estimated skeletal mass of P. colossus exceeds that of any known mammal or aquatic vertebrate. We show that the bone structure specializations of aquatic mammals are reflected in the scaling of skeletal fraction (skeletal mass versus whole-body mass) across the entire disparity of amniotes. We use the skeletal fraction to estimate the body mass of P. colossus, which proves to be a contender for the title of heaviest animal on record. Cetacean peak body mass had already been reached around 30 million years before previously assumed, in a coastal context in which primary productivity was particularly high.

No deep diving: evidence of predation on epipelagic fish for a stem beaked whale from the Late Miocene of Peru.

Although modern beaked whales (Ziphiidae) are known to be highly specialized toothed whales that predominantly feed at great depths upon benthic and benthopelagic prey, only limited palaeontological data document this major ecological shift. We report on a ziphiid-fish assemblage from the Late Miocene of Peru that we interpret as the first direct evidence of a predator-prey relationship between a ziphiid and epipelagic fish. Preserved in a dolomite concretion, a skeleton of the stem ziphiid Messapicetus gregarius was discovered together with numerous skeletons of a clupeiform fish closely related to the epipelagic extant Pacific sardine (Sardinops sagax). Based on the position of fish individuals along the head and chest regions of the ziphiid, the lack of digestion marks on fish remains and the homogeneous size of individuals, we propose that this assemblage results from the death of the whale (possibly via toxin poisoning) shortly after the capture of prey from a single school. Together with morphological data and the frequent discovery of fossil crown ziphiids in deep-sea deposits, this exceptional record supports the hypothesis that only more derived ziphiids were regular deep divers and that the extinction of epipelagic forms may coincide with the radiation of true dolphins.

Piscivory in a Miocene Cetotheriidae of Peru: first record of fossilized stomach content for an extinct baleen-bearing whale.

Instead of teeth, modern mysticetes bear hair-fringed keratinous baleen plates that permit various bulk-filtering predation techniques (from subsurface skimming to lateral benthic suction and engulfment) devoted to various target prey (from small invertebrates to schooling fish). Current knowledge about the feeding ecology of extant cetaceans is revealed by stomach content analyses and observations of behavior. Unfortunately, no fossil stomach contents of ancient mysticetes have been described so far; the investigation of the diet of fossil baleen whales, including the Neogene family Cetotheriidae, remains thus largely speculative. We report on an aggregate of fossil fish remains found within a mysticete skeleton belonging to an undescribed late Miocene (Tortonian) cetotheriid from the Pisco Formation (Peru). Micro-computed tomography allowed us to interpret it as the fossilized content of the forestomach of the host whale and to identify the prey as belonging to the extant clupeiform genus Sardinops. Our discovery represents the first direct evidence of piscivory in an ancient edentulous mysticete. Since among modern mysticetes only Balaenopteridae are known to ordinarily consume fish, this fossil record may indicate that part of the cetotheriids experimented some degree of balaenopterid-like engulfment feeding. Moreover, this report corresponds to one of the geologically oldest records of Sardinops worldwide, occurring near the Tortonian peak of oceanic primary productivity and cooling phase. Therefore, our discovery evokes a link between the rise of Cetotheriidae; the setup of modern coastal upwelling systems; and the radiation of epipelagic, small-sized, schooling clupeiform fish in such highly productive environments.

Taphonomy of marine vertebrates of the Pisco Formation (Miocene, Peru): Insights into the origin of an outstanding Fossil-Lagerstätte.

The Miocene Pisco Formation, broadly exposed in the Ica Desert of southern Peru, is among the most outstanding Cenozoic marine Fossil-Lagerstätten worldwide. It is renowned for its exceptional preservation and abundance of vertebrate fossils, including a rich assemblage of whales and dolphins (Cetacea). Here, we integrate taphonomic data on 890 marine vertebrate fossils, gathered through 16 different localities. Our observations range from the taxonomic distribution, articulation, completeness, disposition and orientation of skeletons, to the presence of bite marks, associations with shark teeth and macro-invertebrates, bone and soft tissue preservation, and the formation of attendant carbonate concretions and sedimentary structures. We propose that the exceptional preservation characterising many Pisco vertebrates, as well as their exceptionally high abundance, cannot be ascribed to a single cause like high sedimentation rates (as proposed in the past), but rather to the interplay of several favourable factors including: (i) low levels of dissolved oxygen at the seafloor (with the intervention of seasonal anoxic events); (ii) the early onset of mineralisation processes like apatite dissolution/recrystallisation and carbonate mineral precipitation; (iii) rapid burial of carcasses in a soupy substrate and/or a novel mechanism involving scour-induced self-burial; and (iv) original biological richness. Collectively, our observations provide a comprehensive overview of the taphonomic processes that shaped one of South America's most important fossil deposits, and suggest a model for the formation of other marine vertebrate Fossil-Lagerstätten.

An Amphibious Whale from the Middle Eocene of Peru Reveals Early South Pacific Dispersal of Quadrupedal Cetaceans.

Cetaceans originated in south Asia more than 50 million years ago (mya), from a small quadrupedal artiodactyl ancestor [1-3]. Amphibious whales gradually dispersed westward along North Africa and arrived in North America before 41.2 mya [4]. However, fossil evidence on when, through which pathway, and under which locomotion abilities these early whales reached the New World is fragmentary and contentious [5-7]. Peregocetus pacificus gen. et sp. nov. is a new protocetid cetacean discovered in middle Eocene (42.6 mya) marine deposits of coastal Peru, which constitutes the first indisputable quadrupedal whale record from the Pacific Ocean and the Southern Hemisphere. Preserving the mandibles and most of the postcranial skeleton, this unique four-limbed whale bore caudal vertebrae with bifurcated and anteroposteriorly expanded transverse processes, like those of beavers and otters, suggesting a significant contribution of the tail during swimming. The fore- and hind-limb proportions roughly similar to geologically older quadrupedal whales from India and Pakistan, the pelvis being firmly attached to the sacrum, an insertion fossa for the round ligament on the femur, and the retention of small hooves with a flat anteroventral tip at fingers and toes indicate that Peregocetus was still capable of standing and even walking on land. This new record from the southeastern Pacific demonstrates that early quadrupedal whales crossed the South Atlantic and nearly attained a circum-equatorial distribution with a combination of terrestrial and aquatic locomotion abilities less than 10 million years after their origin and probably before a northward dispersal toward higher North American latitudes. VIDEO ABSTRACT.

Earliest Mysticete from the Late Eocene of Peru Sheds New Light on the Origin of Baleen Whales.

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.

New beaked whales from the late Miocene of Peru and evidence for convergent evolution in stem and crown Ziphiidae (Cetacea, Odontoceti).

The Ziphiidae (beaked whales) represent a large group of open-ocean odontocetes (toothed cetaceans), whose elusive and deep diving behavior prevents direct observation in their natural habitat. Despite their generally large body size, broad geographical distribution, and high species number, ziphiids thus remain poorly known. Furthermore, the evolutionary processes that have led to their extreme adaptations and impressive extant diversity are still poorly understood. Here we report new fossil beaked whales from the late Miocene of the Pisco Formation (southern Peru). The best preserved remains here described are referred to two new genera and species, the Messinian Chavinziphius maxillocristatus and the Tortonian Chimuziphius coloradensis, based on skull remains from two marine vertebrate-rich localities: Cerro Los Quesos and Cerro Colorado, respectively. C. maxillocristatus is medium sized retains a complete set of functional lower teeth, and bears robust rostral maxillary crests similar to those of the extant Berardius. By contrast, C. coloradensis is small and characterized by large triangular nasals and moderately thickened premaxillae that dorsally close the mesorostral groove. Both species confirm the high past diversity of Ziphiidae, the richest cetacean family in terms of the number of genera and species. Our new phylogenetic and biogeographical analyses depart markedly from earlier studies in dividing beaked whales into two major clades: the Messapicetus clade, which, along with other stem ziphiids, once dominated the southeastern Pacific and North Atlantic; and crown Ziphiidae, the majority of which are found in deep-water regions of the Southern Ocean, with possible subsequent dispersal both globally (Mesoplodon and Ziphius) and to the cooler waters of the northern oceans (Berardius and Hyperoodon). Despite this relatively clear separation, both lineages seem to follow similar evolutionary trends, including (1) a progressive reduction of dentition; (2) an increase in the compactness and thickness of the rostral bones; (3) similar changes in facial morphology (e.g., elevation of the vertex); and (4) an increase of body size. We suggest that these trends may be linked to a convergent ecological shift to deep diving and suction feeding.