Mechanical Analysis of Feeding Behavior in the Extinct “Terror Bird” Andalgalornis steulleti (Gruiformes: Phorusrhacidae)

The South American phorusrhacid bird radiation comprised at least 18 species of small to gigantic terrestrial predators for which there are no close modern analogs. Here we perform functional analyses of the skull of the medium-sized (∼40 kg) patagornithine phorusrhacid Andalgalornis steulleti (upper Miocene–lower Pliocene, Andalgalá Formation, Catamarca, Argentina) to assess its mechanical performance in a comparative context. Based on computed tomographic (CT) scanning and morphological analysis, the skull of Andalgalornis steulleti is interpreted as showing features reflecting loss of intracranial immobility. Discrete anatomical attributes permitting such cranial kinesis are widespread phorusrhacids outgroups, but this is the first clear evidence of loss of cranial kinesis in a gruiform bird and may be among the best documented cases among all birds. This apomorphic loss is interpreted as an adaptation for enhanced craniofacial rigidity, particularly with regard to sagittal loading. We apply a Finite Element approach to a three-dimensional (3D) model of the skull. Based on regression analysis we estimate the bite force of Andalgalornis at the bill tip to be 133 N. Relative to results obtained from Finite Element Analysis of one of its closest living relatives (seriema) and a large predatory bird (eagle), the phorusrhacid''s skull shows relatively high stress under lateral loadings, but low stress where force is applied dorsoventrally (sagittally) and in “pullback” simulations. Given the relative weakness of the skull mediolaterally, it seems unlikely that Andalgalornis engaged in potentially risky behaviors that involved subduing large, struggling prey with its beak. We suggest that it either consumed smaller prey that could be killed and consumed more safely (e.g., swallowed whole) or that it used multiple well-targeted sagittal strikes with the beak in a repetitive attack-and-retreat strategy.     

Implications of Diet for the Extinction of Saber-Toothed Cats and American Lions

The saber-toothed cat, Smilodon fatalis, and American lion, Panthera atrox, were among the largest terrestrial carnivores that lived during the Pleistocene, going extinct along with other megafauna ∼12,000 years ago. Previous work suggests that times were difficult at La Brea (California) during the late Pleistocene, as nearly all carnivores have greater incidences of tooth breakage (used to infer greater carcass utilization) compared to today. As Dental Microwear Texture Analysis (DMTA) can differentiate between levels of bone consumption in extant carnivores, we use DMTA to clarify the dietary niches of extinct carnivorans from La Brea. Specifically, we test the hypothesis that times were tough at La Brea with carnivorous taxa utilizing more of the carcasses. Our results show no evidence of bone crushing by P. atrox, with DMTA attributes most similar to the extant cheetah, Acinonyx jubatus, which actively avoids bone. In contrast, S. fatalis has DMTA attributes most similar to the African lion Panthera leo, implying that S. fatalis did not avoid bone to the extent previously suggested by SEM microwear data. DMTA characters most indicative of bone consumption (i.e., complexity and textural fill volume) suggest that carcass utilization by the extinct carnivorans was not necessarily more complete during the Pleistocene at La Brea; thus, times may not have been “tougher” than the present. Additionally, minor to no significant differences in DMTA attributes from older (∼30–35 Ka) to younger (∼11.5 Ka) deposits offer little evidence that declining prey resources were a primary cause of extinction for these large cats.     

Diets of sympatric native and introduced carnivores in the Barrington Tops,eastern Australia

Invasive predators have severe impacts on global biodiversity, and their effects in Australia have been more extreme than on any other continent. The spotted‐tailed quoll (Dasyurus maculatus), an endangered marsupial carnivore, coexists with three eutherian carnivores, the red fox (Vulpes vulpes), feral cat (Felis catus) and wild dog (Canis lupus ssp.) with which it did not coevolve. No previous study has investigated dietary overlap between quolls and the suite of three eutherian carnivores. By analysing scats, we aimed to quantify dietary overlap within this carnivore assemblage in eastern Australia, and to detect any differences that may facilitate coexistence. We also sought evidence of intraguild predation. Dietary overlap between predators was extensive, with the greatest similarity occurring between foxes and cats. However, some differences were apparent. For example, cats mainly consumed smaller prey, and wild dogs larger prey. Quolls showed greater dietary overlap with foxes and cats than with dogs. Intraguild predation was evident, with fox remains occurring in 3% of wild dog scats. Our results suggest wild dogs competitively dominate invasive foxes, which in turn are likely to compete with the endangered quoll.     

Differences in diet and trophic interactions of Patagonian carnivores between areas with mostly native or exotic prey

The prey base for Patagonian carnivores has been altered greatly over the past 150 years due to widespread overgrazing by livestock, invasions by exotic wildlife, and hunting. On ranches in northern Patagonia carnivores consume mostly exotic species, and native herbivores are ecologically extinct in their role as prey. In this study we compare diets of the culpeo (Lycalopex culpaeus), chilla (L. griseus), puma (Puma concolor), Geoffroy's cat (Leopardus geoffroyi), colocolo (L. colocolo), and hog-nosed skunks (Conepatus chinga and C. humboldtii) in a reserve with a mostly native prey base to their diets on the ranches, and evaluate how differences in prey bases affect trophic interactions among carnivores. Carnivores in the reserve consumed mostly native prey. Dietary overlap among carnivores was not significant on the reserve, but was highly significant on the ranches. This homogenization of diets where densities of native species are reduced could lead to stronger negative interactions among carnivores, altering the composition of the carnivore assemblage to the detriment of the more specialized species. Study of carnivore diets may be a relatively quick way to evaluate the conservation status and ecological functionality of prey assemblages in Patagonia and other areas where these have been altered.     

Immunological Insights into the Life and Times of the Extinct Tasmanian Tiger (Thylacinus cynocephalus)

The thylacine (Thylacinus cynocephalus) was Australia’s largest marsupial carnivore until its extinction within the last century. There remains considerable interest and debate regarding the biology of this species. Studies of thylacine biology are now limited to preserved specimens, and parts thereof, as well as written historical accounts of its biology. This study describes the development of the immune tissues of a pouch young thylacine, one of only eleven in existence, and the only specimen to be histologically sectioned. The appearance of the immune tissue of the developing pouch young thylacine is compared to the immune tissues of extant marsupials, providing insights into the immunity, biology and ecology of the extinct thylacine.     

Forelimb anatomy and the discrimination of the predatory behavior of carnivorous mammals: The thylacine as a case study

Carnivorous mammals use their forelimbs in different ways to capture their prey. Most terrestrial carnivores have some cursorial (running) adaptations, but ambush predators retain considerable flexibility in their forelimb movement, important for grappling with their prey. In contrast, predators that rely on pursuit to run down their prey have sacrificed some of this flexibility for locomotor efficiency, in the greater restriction of the forelimb motion to the parasagittal plane. In this article, we measured aspects of the forelimb anatomy (44 linear measurements) in 36 species of carnivorous mammals of known predatory behavior, and used multivariate analyses to investigate how well the forelimb anatomy reflects the predatory mode (ambush, pursuit, or pounce‐pursuit). A prime intention of this study was to establish morphological correlates of behavior that could then be applied to fossil mammals: for this purpose, five individuals of the recently extinct thylacine (Thylacinus cynocephalus) were also included as unknowns. We show that the three different types of predators can be distinguished by their morphology, both in analyses where all the forelimb bones are included together, and in the separate analyses of each bone individually. Of particular interest is the ability to distinguish between the two types of more cursorial predators, pursuit and pounce‐pursuit, which have previously been considered as primarily size‐based categories. Despite a prior consideration of the thylacine as a “pounce‐pursuit” or an “ambush” type of predator, the thylacines did not consistently cluster with any type of predatory carnivores in our analyses. Rather, the thylacines appeared to be more generalized in their morphology than any of the extant carnivores. The absence of a large diversity of large carnivorous mammals in Australia, past and present, may explain the thylacine's generalized morphology. J. Morphol. 275:1321–1338, 2014. © 2014 Wiley Periodicals, Inc.     

Diel patterns in space use, food and metabolic activity of Galaxias maculatus (Pisces: Galaxiidae) in the littoral zone of a shallow Patagonian lake

We detected that Galaxias maculatus exhibits a pattern where metabolic activity increases after sunrise and peaks between noon and sunset, but this species feeds in the afternoon, until several hours after sunset. Moreover, we showed that G. maculatus is observed in the littoral zone during the day, disappears completely from this zone after sunset and returns at sunrise. Littoral prey species are common in the diet of G. maculatus, but this study showed that pelagic prey is also present during twilight and night hours in smaller individuals (<50 mm), which is related to habitat use. These behavioural rhythms are especially important for G. maculatus, which runs a high predation risk when consuming prey that is widely available outside the littoral zone. This risk is ameliorated under the protection of low light intensity. Thus, G. maculatus is a key species linking lower trophic levels, such as the plankton community, to higher levels of native and exotic piscivores. These displacements of G. maculatus generate an active flow of energy and matter between habitats, with a potentially profound effect on the entire food network and energy dynamics of the lake.     

Sparassocynus (Marsupialia, Didelphidae), a peculiar mammal from the late Cenozoic of Argentina

The extinct genus Sparassocynus is known from the Montehermosan, Chapadmalalan, and Uquian stages of Argentina, spanning the Pliocene-Pleistocene transition, and a relative occurs in the older Huayquerian. After a taxonomict review, the morphology of skull, mandible, and dentition is described. The short, triangular face, wide zygomata, broad cranium with large completely bone-enclosed hypo- and epitympanic sinuses, and carnassialized molars are unique among known American marsupials. All these characters are functionally convergent toward living Australian phascogaline dasyurids, small marsupial carnivores, although phylogenetically Sparassocynus clearly represents a moderately specialized offshoot of the American family Didelphidae. It represents a small steppe carnivore now ecologically replaced, at least in part, by placental carnivores that spread from North America in the late Pliocene and Pleistocene.     

Tooth root morphology as an indicator for dietary specialization in carnivores (Mammalia: Carnivora)

The Carnivora occupy a wide range of feeding niches in concordance with the enormous diversity in their skull and dental form. It is well established that differences in crown morphology are linked to variations in the material properties of the foods ingested and masticated. However, how tooth root form is related to dietary specialization is less well known. In the present study, we investigate the relationship between tooth root morphology and dietary specialization in terrestrial carnivores (canids, felids, hyaenids, and ursids). We specifically address the question of how variation in tooth root surface area is related to bite force potentials as one of the crucial masticatory performance parameters in feeding ecology. We applied computed tomography imaging to reconstruct and quantify dental root surface area in 17 extant carnivore species. Moreover, we computed maximal bite force at several tooth positions based on a dry skull model and assessed the relationship of root surface area to skull size, maximal bite force, food properties, and prey size. We found that postcanine tooth root surface areas corrected for skull size serve as a proxy for bite force potentials and, by extension, dietary specialization in carnivores. Irrespective of taxonomic affinity, species that feed on hard food objects have larger tooth roots than those that eat soft or tough foods. Moreover, carnivores that prey on large animals have larger tooth root surface areas. Our results show that tooth root morphology is a useful indicator of bite force production and allows inferences to be made about dietary ecology in both extant and extinct mammals. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105, 456–471.     

Bite club: comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa

We provide the first predictions of bite force (BS) in a wide sample of living and fossil mammalian predators. To compare between taxa, we calculated an estimated bite force quotient (BFQ) as the residual of BS regressed on body mass. Estimated BS adjusted for body mass was higher for marsupials than placentals and the Tasmanian devil (Sarcophilus harrisii) had the highest relative BS among extant taxa. The highest overall BS was in two extinct marsupial lions. BFQ in hyaenas were similar to those of related, non-osteophagous taxa challenging the common assumption that osteophagy necessitates extreme jaw muscle forces. High BFQ in living carnivores was associated with greater maximal prey size and hypercarnivory. For fossil taxa anatomically similar to living relatives, BFQ can be directly compared, and high values in the dire wolf (Canis dirus) and thylacine (Thylacinus cynocephalus) suggest that they took relatively large prey. Direct inference may not be appropriate where morphologies depart widely from biomechanical models evident in living predators and must be considered together with evidence from other morphological indicators. Relatively low BFQ values in two extinct carnivores with morphologies not represented among extant species, the sabrecat, Smilodon fatalis, and marsupial sabretooth, Thylacosmilus atrox, support arguments that their killing techniques also differed from extant species and are consistent with 'canine-shear bite' and 'stabbing' models, respectively. Extremely high BFQ in the marsupial lion, Thylacoleo carnifex, indicates that it filled a large-prey hunting niche.     

Inferring palaeoecology in extinct tremarctine bears (Carnivora,Ursidae) using geometric morphometrics

Figueirido, B. & Soibelzon, L.H. 2009: Inferring palaeoecology in extinct tremarctine bears (Carnivora, Ursidae) using geometric morphometrics. Lethaia, Vol. 43, pp. 209–222. In this study we explore the ecomorphological patterns of extinct tremarctine bears in South America during the Great American Biotic Interchange (GABI). These patterns are used to derive palaeoautoecological inferences in extinct tremarctines and their palaeosinecological relationships within Plio‐Pleistocene ecosystems. We used geometric morphometrics of landmark data to recover the shape of the craniomandibular skeleton of bears. The results reveal different ecomorphological specializations in extinct tremarctines during the Plio‐Pleistocene of South America. Indeed, these bears could have increased the percentage of plant matter in their diets according with the increased diversity of large carnivores in South America after the GABI. Omnivorous bears retain the ability to behave as carnivores or herbivores depending on resource availability. This fact strongly supports that bears are one of the most ecologically and morphologically adaptable members of the large carnivore guild. Moreover, their skull morphology could reflect ecological adaptations under different selection pressures with the required evolutionary time. □Evolution, GABI, geometric morphometrics, palaeoecology, Tremarctinae.     

An unusual case of cooperative hunting in a solitary carnivore

Cooperative hunting has been documented for several group-living carnivores and had been invoked as either the cause or the consequence of sociality. We report the first detailed observation of cooperative hunting for a solitary species, the Malagasy fossa (Cryptoprocta ferox). We observed a 45 min hunt of a 3 kg arboreal primate by three male fossas. The hunters changed roles during the hunt and subsequently shared the prey. We hypothesize that social hunting in fossas could have either evolved to take down recently extinct larger lemur prey, or that it could be a by-product of male sociality that is beneficial for other reasons.     

Do Australian small mammals respond to native and introduced predator odours?

Abstract Coevolution is thought to have led to many small mammal species avoiding the scent marks of their main mammalian predators, as they provide a reliable cue to predation risk. Most support for this hypothesis comes from northern hemisphere predator/prey systems, however, it is unclear whether this avoidance of predator faecal odour occurs in Australia's mammalian fauna, which has evolved in relative isolation from the rest of the world, and is dominated by marsupials rather than placentals. We tested this theory for an Australian system with marsupial and placental predators and prey, that share a long‐term (>1 million years) or short‐term (<150 years) exposure to each other. The predators were the native marsupial tiger quoll Dasyurus maculatus and the introduced placental red fox Vulpes vulpes. The potential prey were three native rodent species, the bush rat Rattus fuscipes, the swamp rat Rattus lutreolus, the eastern chestnut mouse Pseudomys gracilicaudatus, and the marsupial brown antechinus Antechinus stuartii. Small mammals were captured in Elliott traps with 1/3 of traps treated with fox faeces, 1/3 treated with quoll faeces and the remainder left untreated. The native rodent species all showed avoidance of both tiger quoll and red fox odours whereas the marsupial antechinus showed no responses to either odour. Either predator odour avoidance has not evolved in this marsupial or their reaction to predator odours may be exhibited in ways which are not recognizable through trapping. The avoidance by the rodents of fox odour as well as quoll odour indicates this response may either be due to common components in fox and quoll odour, or it may be a recently evolved response.     

Evidence of intraspecific agonistic interactions in Smilodon populator (Carnivora,Felidae)

The saber-toothed cat Smilodon is a characteristic genus of the Pleistocene faunas of the American continent. Smilodon belongs to an extinct clade of felids that had hypertrophied blade-like upper canines. Because the length of the canines is so extreme, the killing bite of Smilodon is a hotly debated topic in vertebrate paleontology. Some authors have proposed that saber-toothed cats had a weak bite and their canines were fragile, not useful for attacking prey or penetrating bones. The aim of the present contribution is to describe two new specimens of Smilodon populator that have injuries on their skulls. Although it cannot be ruled out that the injuries were caused by a potential prey kicking the skull, the size, shape and general features of the injuries suggest that they were inflicted by the upper canines of another Smilodon individual during agonistic interactions.     

A unique feeding strategy of the extinct marine mammal Kolponomos: convergence on sabretooths and sea otters

Mammalian molluscivores feed mainly by shell-crushing or suction-feeding. The extinct marine arctoid, Kolponomos, has been interpreted as an otter-like shell-crusher based on similar dentitions. However, neither the masticatory biomechanics of the shell-crushing adaptation nor the way Kolponomos may have captured hard-shelled prey have been tested. Based on mandibular symphyseal morphology shared by Kolponomos and sabre-toothed carnivores, we hypothesize a sabretooth-like mechanism for Kolponomos prey-capture, whereby the mandible functioned as an anchor. Torque generated from jaw closure and head flexion was used to dislodge prey by prying, with prey then crushed using cheek teeth. We test this hypothesized feeding sequence using phylogenetically informed biomechanical simulations and shape analyses, and find a strongly supported, shared high mandibular stiffness in simulated prey-capture bites and mandibular shape in Kolponomos and the sabre-toothed cat Smilodon. These two distantly related taxa converged on using mandibles to anchor cranial torqueing forces when prying substrate-bound prey in the former and sabre-driving forces during prey-killing in the latter. Simulated prey-crushing bites indicate that Kolponomos and sea otters exhibit alternative structural stiffness-bite efficiency combinations in mandibular biomechanical adaptation for shell-crushing. This unique feeding system of Kolponomos exemplifies a mosaic of form-function convergence relative to other Carnivora.     

Contemporaneous Trace and Body Fossils from a Late Pleistocene Lakebed in Victoria,Australia, Allow Assessment of Bias in the Fossil Record

The co-occurrence of vertebrate trace and body fossils within a single geological formation is rare and the probability of these parallel records being contemporaneous (i.e. on or near the same bedding plane) is extremely low. We report here a late Pleistocene locality from the Victorian Volcanic Plains in south-eastern Australia in which demonstrably contemporaneous, but independently accumulated vertebrate trace and body fossils occur. Bite marks from a variety of taxa are also present on the bones. This site provides a unique opportunity to examine the biases of these divergent fossil records (skeletal, footprints and bite marks) that sampled a single fauna. The skeletal record produced the most complete fauna, with the footprint record indicating a markedly different faunal composition with less diversity and the feeding traces suggesting the presence, amongst others, of a predator not represented by either the skeletal or footprint records. We found that the large extinct marsupial predator Thylacoleo was the only taxon apparently represented by all three records, suggesting that the behavioral characteristics of large carnivores may increase the likelihood of their presence being detected within a fossil fauna. In contrast, Diprotodon (the largest-ever marsupial) was represented only by trace fossils at this site and was absent from the site''s skeletal record, despite its being a common and easily detected presence in late Pleistocene skeletal fossil faunas elsewhere in Australia. Small mammals absent from the footprint record for the site were represented by skeletal fossils and bite marks on bones.     

Musculoskeletal anatomy of the Eurasian lynx,Lynx lynx (Carnivora: Felidae) forelimb: Adaptations to capture large prey?

Mammalian carnivores adhere to two different feeding strategies relative to their body masses. Large carnivores prey on animals that are the same size or larger than themselves, whereas small carnivores prey on smaller vertebrates and invertebrates. The Eurasian lynx (Lynx lynx) falls in between these two categories. Lynx descend from larger forms that were probably large prey specialists, but during the Pleistocene became predators of small prey. The modern Eurasian lynx may be an evolutionary reversal toward specializing in large prey again. We hypothesized that the musculoskeletal anatomy of lynx should show traits for catching large prey. To test our hypothesis, we dissected the forelimb muscles of six Eurasian lynx individuals and compared our findings to results published for other felids. We measured the bones and compared their dimensions to the published material. Our material displayed a well‐developed pectoral girdle musculature with some uniquely extensive muscle attachments. The upper arm musculature resembled that of the pantherine felids and probably the extinct sabertooths, and also the muscles responsible for supination and pronation were similar to those in large cats. The muscles controlling the pollex were well‐developed. However, skeletal indices were similar to those of small prey predators. Our findings show that lynx possess the topographic pattern of muscle origin and insertion like in large felids. J. Morphol. 277:753–765, 2016. © 2016 Wiley Periodicals, Inc.     

Tooth Root Morphology in the Early Pliocene African Bear Agriotherium africanum (Mammalia, Carnivora, Ursidae) and its Implications for Feeding Ecology

Tooth root surface areas serve as proxies for bite force potentials, and by extension, dietary specialization in extant carnivorans. Here, we investigate the feeding ecology of the extinct large-bodied ursid Agriotherium africanum, by comparing its root surface areas (reconstructed with the aid of computed tomography and three-dimensional image processing) and bite force estimates, with those of extant carnivorans. Results show that in absolute terms, canine and carnassial bite forces, as well as root surface areas were highest in A. africanum. However, when adjusted for skull size, A. africanum’s canine roots were smaller than those of extant solitary predators. With teeth being the limiting factor in the masticatory system, low canine root surface areas suggest that A. africanum would have struggled to bring down large vertebrate prey. Its adjusted carnassial root sizes were found to be smaller than those of extant hard object feeders and the most carnivorous tough object feeders, but larger than those of extant omnivorous ursids and Ursus maritimus. This and the fact that it displayed its highest postcanine root surface areas in the carnassial region (rather than the most distal tooth in the tooth row) suggest that A. africanum consumed more vertebrate tissue than extant omnivorous ursids. With an apparent inability to routinely bring down large prey or to consume mechanically demanding skeletal elements, its focus was most likely on tough tissue, which it acquired by actively scavenging the carcasses of freshly dead/freshly killed animals. Mechanically less demanding skeletal elements would have been a secondary food source, ingested and processed mainly in association with muscle and connective tissue.     

Anthropogenic extinction of top carnivores and interspecific animal behaviour: implications of the rapid decoupling of a web involving wolves, bears, moose and ravens

The recent extinction of grizzly bears (Ursus arctos) and wolves (Canis lupus) by humans from 95-99% of the contiguous USA and Mexico in less than 100 years has resulted in dramatically altered and expanded prey communities. Such rampant ecological change and putative ecological instability has not occurred in North American northern boreal zones. This geographical variation in the loss of large carnivores as a consequence of anthropogenic disturbance offers opportunities for examining the potential consequences of extinction on subtle but important ecological patterns involving behaviour and interspecific ecological interactions. In Alaska, where scavengers and large carnivores are associated with carcasses, field experiments involving sound playback simulations have demonstrated that at least one prey species, moose (Alces alces), is sensitive to the vocalizations of ravens (Corvus corax) and may rely on their cues to avoid predation. However, a similar relationship is absent on a predator-free island in Alaska's Cook Inlet and at two sites in the Jackson Hole region of the Rocky Mountains (USA) where grizzly bears and wolves have been extinct for 50-70 years. While prior study of birds and mammals has demonstrated that prey may retain predator recognition capabilities for thousands of years even after predation as a selective force has been relaxed, the results presented here establish that a desensitization in interspecific responsiveness can also occur in less than ten generations. These results affirm (i) a rapid decoupling in behaviour involving prey and scavengers as a consequence of anthropogenic-caused predator-prey disequilibriums, and (ii) subtle, community-level modifications in terrestrial ecosystems where large carnivores no longer exist. If knowledge about ecological and behavioural processes in extant systems is to be enhanced, the potential effects of recently extinct carnivores must be incorporated into current programmes.     

A New Horned Crocodile from the Plio-Pleistocene Hominid Sites at Olduvai Gorge,Tanzania

Background

The fossil record reveals surprising crocodile diversity in the Neogene of Africa, but relationships with their living relatives and the biogeographic origins of the modern African crocodylian fauna are poorly understood. A Plio-Pleistocene crocodile from Olduvai Gorge, Tanzania, represents a new extinct species and shows that high crocodylian diversity in Africa persisted after the Miocene. It had prominent triangular “horns” over the ears and a relatively deep snout, these resemble those of the recently extinct Malagasy crocodile Voay robustus, but the new species lacks features found among osteolaemines and shares derived similarities with living species of Crocodylus.

Methodology/Principal Findings

The holotype consists of a partial skull and skeleton and was collected on the surface between two tuffs dated to approximately 1.84 million years (Ma), in the same interval near the type localities for the hominids Homo habilis and Australopithecus boisei. It was compared with previously-collected material from Olduvai Gorge referable to the same species. Phylogenetic analysis places the new form within or adjacent to crown Crocodylus.

Conclusions/Significance

The new crocodile species was the largest predator encountered by our ancestors at Olduvai Gorge, as indicated by hominid specimens preserving crocodile bite marks from these sites. The new species also reinforces the emerging view of high crocodylian diversity throughout the Neogene, and it represents one of the few extinct species referable to crown genus Crocodylus.