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.

Sperm whales (Physeteroidea) from the Pisco Formation, Peru, and their trophic role as fat sources for late Miocene sharks.

Shark-cetacean trophic interactions, preserved as bite marks in the fossil record, mostly correspond to isolated or fragmentary findings that bear limited information about major trophic patterns or roles. Here, we provide evidence of focalized foraging by sharks in the form of tooth bite marks over physeteroids fossil bones from the late Miocene of Peru. These findings indicate that sharks were targeting the forehead of coeval physeteroids to actively feed on their lipid-rich nasal complexes. Miocene physeteroids displayed a broad diversity, including giant predatorial forms, small benthic foragers and suction feeders. Like their extant relatives, these animals exhibited enlarged fatty forehead organs responsible for their sound production capabilities, thus evolving taxon-specific cranial architecture. Bite marks are found on the cranial bones where these structures were attached, indicating that sharks actively targeted this region; but also, in areas that would only be accessible following the consumption of the surrounding soft tissues. The shape of the bite marks and their distribution suggests a series of consecutive scavenging events by individuals of different shark species. Similar bite patterns can be recognized on other Miocene physeteroids fossils from across the globe, suggesting that sharks actively exploited physeteroid carcasses as fat sources.

Miocene fossils from the southeastern Pacific shed light on the last radiation of marine crocodylians.

The evolution of crocodylians as sea dwellers remains obscure because living representatives are basically freshwater inhabitants and fossil evidence lacks crucial aspects about crocodylian occupation of marine ecosystems. New fossils from marine deposits of Peru reveal that crocodylians were habitual coastal residents of the southeastern Pacific (SEP) for approximately 14 million years within the Miocene (ca 19 to 5 Ma), an epoch including the highest global peak of marine crocodylian diversity. The assemblage of the SEP comprised two long and slender-snouted (longirostrine) taxa of the Gavialidae: the giant Piscogavialis and a new early diverging species, Sacacosuchus cordovai. Although living gavialids (Gavialis and Tomistoma) are freshwater forms, this remarkable fossil record and a suite of evolutionary morphological analyses reveal that the whole evolution of marine crocodylians pertained to the gavialids and their stem relatives (Gavialoidea). This adaptive radiation produced two longirostrine ecomorphs with dissimilar trophic roles in seawaters and involved multiple transmarine dispersals to South America and most landmasses. Marine gavialoids were shallow sea dwellers, and their Cenozoic diversification was influenced by the availability of coastal habitats. Soon after the richness peak of the Miocene, gavialoid crocodylians disappeared from the sea, probably as part of the marine megafauna extinction of the Pliocene.

Correction to: Late Neogene evolution of the Peruvian margin and its ecosystems: a synthesis from the Sacaco record.

[This corrects the article DOI: 10.1007/s00531-021-02003-1.].

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.

La hiperdiversificación de Physeteroidea (Cetacea: Mammalia) del Mioceno tardío en el Pacifico Sur, nuevos fósiles del área de Sacaco, Arequipa, Perú / Physeteroidea (Cetacea: Mammalia) hyper-diversification from late Miocene of South Pacific, new fossils from the Sacaco area, Arequipa, Peru

Resumen Los cachalotes (Physeteroidea) son un grupo de cetáceos con una ecología restringida en la actualidad. Las tres especies actuales son el remanente de un grupo que alcanzó su máxima diversificacion durante el Mioceno, incluyendo cinco especies simpátricas descritas en la Formación Pisco, Ica-Arequipa, Perú. Entre estas se incluyen formas piscívoras, bénticas, así como carnívoros de tamaño medio e hipercarnívoros. Se reportan dos dientes aislados provenientes del Mioceno superior de las localidades de Sacaco y Sacaco sur, que corresponden con la morfología típica de Physeteridae. Esta morfología dentaria es similar a la de taxones presentes en el hemisferio norte como Orycterocetus o Aulophyseter del Mioceno medio. Debido a esto se presume que los restos corresponderían a physetéridos con hábitos piscívoros, que quizás poseyeron un desarrollo incipiente de la capacidad de succión. La presencia de este linaje indicaría que a fines del Mioceno los parientes de los modernos Physeter y Kogia compartieron el mar peruano con formas extintas, sin solapamiento de nichos debido a la morfología altamente derivada de cada grupo. Este nuevo registro expande la comunidad de cetáceos fósiles de Sacaco, que se vio favorecida por la gran diversidad de ambientes poco profundos que dominaron el litoral peruano hasta el establecimiento final del Sistema Humboldt.

Abstract Sperm whales (Physeteroidea) are a group of cetaceans with a restricted modern ecology. The three extant species are a remnant of a group that reached its diversity peak during the Miocene, including five already-described species from the Pisco Formation, Ica-Arequipa, Peru. We report two isolated teeth from the upper Miocene localities of Sacaco and Sud-Sacaco, which correspond with the typical morphology of Physeteridae. This morphology is similar to the one present in northern hemisphere taxa as Orycterocetus or Aulophyseter. Because of this the remains could be related to physeterids with piscivorous habits, so there would not be a niche overlap with other coeval sperm whales. This new registry expands the fossil cetacean community of Sacaco, which was favored by the great diversity of shallow environments that dominated the Peruvian coast until the final establishment of the Humboldt System.

Extensive Diversity and Disparity of the Early Miocene Platanistoids (Cetacea, Odontoceti) in the Southeastern Pacific (Chilcatay Formation, Peru).

Several aspects of the fascinating evolutionary history of toothed and baleen whales (Cetacea) are still to be clarified due to the fragmentation and discontinuity (in space and time) of the fossil record. Here we open a window on the past, describing a part of the extraordinary cetacean fossil assemblage deposited in a restricted interval of time (19-18 Ma) in the Chilcatay Formation (Peru). All the fossils here examined belong to the Platanistoidea clade as here redefined, a toothed whale group nowadays represented only by the Asian river dolphin Platanista gangetica. Two new genera and species, the hyper-longirostrine Ensidelphis riveroi and the squalodelphinid Furcacetus flexirostrum, are described together with new material referred to the squalodelphinid Notocetus vanbenedeni and fragmentary remains showing affinities with the platanistid Araeodelphis. Our cladistic analysis defines the new clade Platanidelphidi, sister-group to Allodelphinidae and including E. riveroi and the clade Squalodelphinidae + Platanistidae. The fossils here examined further confirm the high diversity and disparity of platanistoids during the early Miocene. Finally, morphofunctional considerations on the entire platanistoid assemblage of the Chilcatay Formation suggest a high trophic partitioning of this peculiar cetacean paleocommunity.

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.

A new large squalodelphinid (Cetacea, Odontoceti) from Peru sheds light on the Early Miocene platanistoid disparity and ecology.

The South Asian river dolphin (Platanista gangetica) is the only extant survivor of the large clade Platanistoidea, having a well-diversified fossil record from the Late Oligocene to the Middle Miocene. Based on a partial skeleton collected from the Chilcatay Formation (Chilcatay Fm; southern coast of Peru), we report here a new squalodelphinid genus and species, Macrosqualodelphis ukupachai. A volcanic ash layer, sampled near the fossil, yielded the 40Ar/39Ar age of 18.78 ± 0.08 Ma (Burdigalian, Early Miocene). The phylogenetic analysis places Macrosqualodelphis as the earliest branching squalodelphinid. Combined with several cranial and dental features, the large body size (estimated body length of 3.5 m) of this odontocete suggests that it consumed larger prey than the other members of its family. Together with Huaridelphis raimondii and Notocetus vanbenedeni, both also found in the Chilcatay Fm, this new squalodelphinid further demonstrates the peculiar local diversity of the family along the southeastern Pacific coast, possibly related to their partition into different dietary niches. At a wider geographical scale, the morphological and ecological diversity of squalodelphinids confirms the major role played by platanistoids during the Early Miocene radiation of crown odontocetes.

How whales used to filter: exceptionally preserved baleen in a Miocene cetotheriid.

Baleen is a comb-like structure that enables mysticete whales to bulk feed on vast quantities of small prey, and ultimately allowed them to become the largest animals on Earth. Because baleen rarely fossilises, extremely little is known about its evolution, structure and function outside the living families. Here we describe, for the first time, the exceptionally preserved baleen apparatus of an entirely extinct mysticete morphotype: the Late Miocene cetotheriid, Piscobalaena nana, from the Pisco Formation of Peru. The baleen plates of P. nana are closely spaced and built around relatively dense, fine tubules, as in the enigmatic pygmy right whale, Caperea marginata. Phosphatisation of the intertubular horn, but not the tubules themselves, suggests in vivo intertubular calcification. The size of the rack matches the distribution of nutrient foramina on the palate, and implies the presence of an unusually large subrostral gap. Overall, the baleen morphology of Piscobalaena likely reflects the interacting effects of size, function and phylogeny, and reveals a previously unknown degree of complexity in modern mysticete feeding evolution.

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.

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.

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.

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

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.

Osteology of Icadyptes salasi, a giant penguin from the Eocene of Peru.

We present the first detailed description of the giant Eocene penguin Icadyptes salasi. The species is characterized by a narrow skull with a hyper-elongate spear-like beak, a robust cervical column and a powerful flipper. The bony beak tip of Icadyptes is formed by fusion of several elements and is unique among penguins, differing markedly from previously described giant penguin beaks. Vascular canal patterning similar to that of boobies, frigatebirds and albatrosses suggests I. salasi may have had a thin, sheet-like rhamphotheca unlike the thick rugose rhamphotheca of modern penguins. Together, these features suggest a novel ecology for I. salasi, most likely involving the capture of larger prey items via spearing. As the first described giant penguin specimen to preserve a complete wing skeleton, the I. salasi holotype yields significant insight into the shape, proportions and orientation of the wing in giant penguins. In articulation, the forelimb of I. salasi is straighter, permitting less manus and antibrachium flexion, than previous depictions of giant penguin wings. Cross-sections of the humerus and ulna reveal a level of osteosclerosis equalling or surpassing that of extant penguins. Based on ontogenetic data from extant penguins and the morphology of the carpometacarpus of I. salasi, we infer the retention of a free alular phalanx in basal penguins. Previously, the status of this element in penguins was disputed. Differences in the proportions of the manual phalanges contribute to a more abruptly tapering wingtip in I. salasi compared with crown penguins. Fossils from Peru, including the I. salasi holotype specimen, document that penguins expanded to nearly the whole of their extant latitudinal range early in their evolutionary history and during one of the warmest intervals in the Cenozoic.

Paleogene equatorial penguins challenge the proposed relationship between biogeography, diversity, and Cenozoic climate change.

New penguin fossils from the Eocene of Peru force a reevaluation of previous hypotheses regarding the causal role of climate change in penguin evolution. Repeatedly it has been proposed that penguins originated in high southern latitudes and arrived at equatorial regions relatively recently (e.g., 4-8 million years ago), well after the onset of latest Eocene/Oligocene global cooling and increases in polar ice volume. By contrast, new discoveries from the middle and late Eocene of Peru reveal that penguins invaded low latitudes >30 million years earlier than prior data suggested, during one of the warmest intervals of the Cenozoic. A diverse fauna includes two new species, here reported from two of the best exemplars of Paleogene penguins yet recovered. The most comprehensive phylogenetic analysis of Sphenisciformes to date, combining morphological and molecular data, places the new species outside the extant penguin radiation (crown clade: Spheniscidae) and supports two separate dispersals to equatorial (paleolatitude approximately 14 degrees S) regions during greenhouse earth conditions. One new species, Perudyptes devriesi, is among the deepest divergences within Sphenisciformes. The second, Icadyptes salasi, is the most complete giant (>1.5 m standing height) penguin yet described. Both species provide critical information on early penguin cranial osteology, trends in penguin body size, and the evolution of the penguin flipper.