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We describe two new osteolaemine crocodylids from the Early and early Middle Miocene of Kenya: Kinyang mabokoensis tax. nov. (Maboko, 15 Ma) and Kinyang tchernovi tax. nov. (Karungu and Loperot, 18 Ma). Additional material referable to Kinyang is known from Chianda and Moruorot. The skull was broad and dorsoventrally deep, and the genus can be diagnosed based on the combined presence of a partial overbite, a subdivided fossa for the lateral collateral ligament on the surangular, and a maxilla with no more than 13 alveoli. Phylogenetic analyses based on morphological and combined morphological and molecular data support a referral of Kinyang to Osteolaeminae, and morphological data alone put the new taxon at the base of Euthecodontini. Some Kinyang maxillae preserve blind pits on the medial caviconchal recess wall. Kinyang co-occurs with the osteolaemine Brochuchus at some localities, and together, they reinforce the phylogenetic disparity between early Neogene osteolaemine-dominated faunas and faunas dominated by crocodylines beginning in the Late Miocene in the Kenya Rift. The causes of this turnover remain unclear, though changes in prevailing vegetation resulting from tectonic and climatic drivers may provide a partial explanation.

Brochuchusis a small crocodylid originally based on specimens from the early Miocene of Rusinga Island, Lake Victoria, Kenya. Here, we report occurrences ofBrochuchusfrom several early and middle Miocene sites. Some are from the Lake Victoria region, and others are in the Lake Turkana Basin. Specimens from the middle Miocene Maboko locality form the basis of a new species,Brochuchus parvidens, which has comparatively smaller maxillary alveoli. Because of the smaller alveoli, the teeth appear to be more widely spaced in the new species. We also provide a revised diagnosis forBrochuchusand its type species,B.pigotti. A phylogenetic analysis supports a close relationship betweenBrochuchusand tube-snoutedEuthecodon, but although relationships among crocodylids appear poorly resolved in the set of optimal trees, this is becauseBrochuchusandEuthecodon, along with early Miocene “Crocodylus”gariepensisfrom the early Miocene of Namibia, jointly adopt two distinct positions—either closely related to the living sharp-nosed crocodile (Mecistops) or to a group including the living dwarf crocodiles (Osteolaemus). Character support for a close relationship withMecistopsis problematic, and we suspect a closer relationship toOsteolaemuswill be recovered with improved sampling, but the results here are ambiguous. In either case,Brochuchusis more closely related to living groups not currently found in East Africa. This material helps constrain the timing of crocodylian faunal turnover in the East African Rift Valley System, with endemic lineages largely being replaced byCrocodylusin the middle or late Miocene possibly in response to regional xerification and the replacement of continuous rainforest cover with open grasslands and savannas.UUID:http://zoobank.org/e6f0b219-5f3e-44e5-bdb9-60a4fae8d126

The “death roll” is an iconic crocodylian behaviour, and yet it is documented in only a small number of species, all of which exhibit a generalist feeding ecology and skull ecomorphology. This has led to the interpretation that only generalist crocodylians can death roll, a pattern which has been used to inform studies of functional morphology and behaviour in the fossil record, especially regarding slender-snouted crocodylians and other taxa sharing this semi-aquatic ambush predator body plan. In order to test this hypothesis, we surveyed death roll behaviour across animals representing all extant crocodylian species. Animals were prompted to death roll using two methods of stimulation: a feeding cue and an escape cue. The feeding cue involved presenting each animal with a bait item, to which resistance would be applied during a biting event. The second cue involved capturing each animal with a rope or catch pole, a standard technique for capturing crocodylians, but one that also often prompts an attempt to escape. All species tested, except Paleosuchus palpebrosus, exhibited the behaviour in response to at least one of the stimuli. This included the following slender-snouted species: Gavialis gangeticus, Tomistoma schlegelii, Mecistops cataphractus, Mecistops leptorhynchus, Crocodylus johnstoni, and Crocodylus intermedius. The patterns of death roll behavior observed in this survey suggest that this behaviour is not novel to any one crocodylian clade, morphotype, or dietary niche. Also, the prevalence of death rolling behaviour across Crocodylia in response to perceived threats indicates that it is not solely, or maybe even primarily, a feeding behaviour, but is also utilised during inter- and intra-specific conflict as a means to escape or injure an opponent. The results of this case study highlight the importance of using multiple modern analogues when attempting to correlate form and function across diverse clades, both living and extinct.

New material attributable to Deltasuchus motherali, a neosuchian from the Cenomanian of Texas, provides sampling across much of the ontogeny of this species. Detailed descriptions provide information about the paleobiology of this species, particularly with regards to how growth and development affected diet. Overall snout shape became progressively wider and more robust with age, suggesting that dietary shifts from juvenile to adult were not only a matter of size change, but of functional performance as well. These newly described elements provide additional characters upon which to base more robust phylogenetic analyses. The authors provide a revised diagnosis of this species, describing the new material and discussing incidents of apparent ontogenetic variation across the sampled population. The results of the ensuing phylogenetic analyses both situate Deltasuchus within an endemic clade of Appalachian crocodyliforms, separate and diagnosable from goniopholidids and pholidosaurs, herein referred to as Paluxysuchidae. This title is also available as Open Access on Cambridge Core.

Several data loggers deployed to monitor temperature and humidity of Crocodylus acutus (American crocodile) nests in South Florida could not be located after hatching. One badly damaged data logger was retrieved, providing insight into the possible fate of the others. Using a taphonomic approach, we identified numerous indentations, consistent with crocodylian bite marks, and inconsistent with potential mammalian or squamate bites. It seems most likely that the data logger was damaged by the nesting C. acutus rather than during attempted nest predation. Estimated bite forces for reproductive age, female C. acutus exceed the predicted material properties of the data logger’s housing, suggesting that the bites were exploratory in nature. We suggest that data loggers be removed prior to hatching or permit remote data storage.

Bite marks provide direct evidence for trophic interactions and competition in the fossil record. However, variations in paleoecological dynamics, such as trophic relationships, feeding behavior, and food availability, govern the frequency of these traces. Theropod bite marks are particularly rare, suggesting that members of this clade might not often focus on bone as a resource, instead preferentially targeting softer tissues. Here, we present an unusually large sample of theropod bite marks from the Upper Jurassic Mygatt-Moore Quarry (MMQ). We surveyed 2,368 vertebrate fossils from MMQ in this analysis, with 684 specimens (28.885% of the sample) preserving at least one theropod bite mark. This is substantially higher than in other dinosaur-dominated assemblages, including contemporaneous localities from the Morrison Formation. Observed bite marks include punctures, scores, furrows, pits, and striations. Striated marks are particularly useful, diagnostic traces generated by the denticles of ziphodont teeth, because the spacing of these features can be used to provide minimum estimates of trace maker size. In the MMQ assemblage, most of the striations are consistent with denticles of the two largest predators known from the site: Allosaurus and Ceratosaurus. One of the bite marks suggests that a substantially larger theropod was possibly present at the site and are consistent with large theropods known from other Morrison Formation assemblages (either an unusually large Allosaurus or a separate, large-bodied taxon such as Saurophaganax or Torvosaurus). The distribution of the bite marks on skeletal elements, particularly those found on other theropods, suggest that they potentially preserve evidence of scavenging, rather than active predation. Given the relative abundances of the MMQ carnivores, partnered with the size-estimates based on the striated bite marks, the feeding trace assemblage likely preserves the first evidence of cannibalism in Allosaurus.

Existing classifications of snout shape within Crocodylia are supported by functional studies, but ecological surveys often reveal a higher than expected diversity of prey items within putatively specialist groups, and research into bite force and predation behaviour does not always reveal significant differences between snout shape groups. The addition of more distantly related crocodyliforms complicates the ecomorphological signal, because these groups often occupy a larger area of morphospace than the crown group alone. Here, we present an expanded classification of snout shapes and diets across Crocodyliformes, bringing together geometric morphometrics, non-hierarchical cluster analyses, phylogenetic analyses, ancestral state reconstructions, ecological surveys of diet, and feeding traces from the fossil record to build and test predictive models for linking snout shape and function across the clade. When applied to living members of the group, these new classifications partition out based on differences in predator body mass and maximal prey size. When applied to fossils, these classifications predict potential prey items and identify possible examples of scavenging. In a phylogenetic context, these ecomorphs reveal differences in dietary strategies and diversity within major crocodyliform clades. Taken together, these patterns suggest that crocodyliform diversity, in terms of both morphology and diet, has been underestimated.

Captive broad snouted crocodylians are generally thought to have wider, shorter rostra than their wild counterparts. Interpreted to reflect morphological change in response to the conditions of captivity, this qualitative pattern could affect the utility of these animals in a variety of fields of research. However, due to relative ease of access and availability of life history data, captive animals are often utilized in actualistic research. Thus, this issue should be addressed in more detail. Here we explore snout shape variation between captive and wild members of Alligator mississippiensis using two-dimensional (2D) morphometric techniques. Several landmark schemesare used to assess the utility of different aspects of morphology in distinguishing the groups. While statistical analyses consistently differentiated between the groups, the area of morphospace occupied by wild members of A. mississippiensis generally overlapped with the larger area encompassing the captive specimens. This indicates that the captive condition is not as uniform as previously thought and instead encompasses a large spectrum of morphologies, ranging from the stereotypical broad, shortened snouts to outlines that are indistinguishable from the wild morphotype. These results align well with the interpretation that this change reflects an extreme example of ecophenotypy, since ranched, farmed, or zoo organisms are held in an array of enclosures, ranging from indoor, climate controlled pens to outdoor, more natural areas. This variation in environments should be reflected in different reactions to the animals' surroundings, resulting in a broad spectrum of morphotypes. While wild specimens are still preferred, especially for fine scale analyses, these results indicate that not all captive members of A. mississippiensis exhibit the extreme morphological alterations often cited in the literature. Weighing the conditions in which the animals are held and exploring the possibility of morphological differences against the benefits of using captive specimens should be part of any actualistic study. J. Morphol. 277:866-878, 2016. © 2016 Wiley Periodicals, Inc.

New discoveries at the Arlington Archosaur Site (AAS), a Cenomanian (Late Cretaceous) locality in north-central Texas, are filling gaps in our knowledge of mid-Cretaceous Appalachian ecosystems, which remain poorly characterized. The AAS is notable because it preserves a diverse crocodyliform record. As seen in other sites that preserve four or more crocodyliform taxa, the species present at the AAS exhibit different snout shapes and body sizes, indicating that this high diversity of sympatric species was likely sustainable due to niche partitioning. Here we describe Scolomastax sahlsteini gen. et sp. nov., a new species of crocodyliform from the AAS, currently known from a partial right mandibular ramus. This species differs from other crocodyliforms in possessing features associated with durophagy or omnivory, including a shortened mandible, reduced tooth count, heterodonty, a dorsally expanded surangular, and enlarged attachments for jaw adductor muscles. Our phylogenetic analysis places this new taxon within Eusuchia as a member of Paralligatoridae and sister taxon to Paralligator gradilifrons. Scolomastax sahlsteini extends the record of paralligatorids into the Late Cretaceous of North America. This discovery represents the first appearance of this clade on the poorly known landmass of Appalachia, supporting a biogeographic connection between North America and Asia in the Early Cretaceous prior to completion of the Western Interior Seaway. However, relationships among other endemic crocodyliforms and tree instability within Paralligatoridae suggest further analysis is needed to resolve phylogenetic and biogeographic relationships (http://zoobank.org/urn:lsid:zoobank.org:pub:DC114471-6687-4BB5-8FAE-96F7278B1DAF). Anat Rec, 303:801-812, 2020. © 2019 Wiley Periodicals, Inc.

Crocodylians are known to consume and modify bones, but actualistic observations of their bite marks have been limited to forensic case studies and surveys of two taxa: Crocodylus niloticus and Crocodylus porosus. To further explore patterns of crocodylian bite mark expression, we conducted a survey of traces left by Alligator mississippiensis. We compared the results to pre-existing crocodylian datasets regarding the potentially diagnostic traits of bisected marks, hook scores, and a lack of furrows. Mark type did not correlate with vital statistics of the sampled animals or collections protocol. Bisected marks were found in rates similar to those seen in one previous survey of C. niloticus, and rates of hook scoring and bone breakage were higher. These traces were all present in higher rates than those reported in C. porosus. Unlike results seen in Crocodylus, furrows were identified in the A. mississippiensis samples. Hook scores were also identified, but recent surveys of non-crocodylian taxa have sho...

Heterodonty in Crocodylia and closely related taxa has not been defined quantitatively, as the teeth rarely have been measured. This has resulted in a range of qualitative descriptors, with little consensus on the condition of dental morphology in the clade. The purpose of this study is to present a method for the quantification of both size- and shape-heterodonty in members of Crocodylia. Data were collected from dry skeletal and fossil specimens of 34 crown crocodylians and one crocodyliform, resulting in 21 species total. Digital photographs were taken of each tooth and the skull, and the margins of both were converted into landmarks and semilandmarks. We expressed heterodonty through Foote’s morphological disparity, and a principal components analysis quantified shape variance. All specimens sampled were heterodont to varying degrees, with the majority of the shape variance represented by a ‘caniniform’ to ‘molariform’ transition. Heterodonty varied significantly between positions; size undulated whereas shape was significantly linear from mesial to distal. Size and shape appeared to be primarily decoupled. Skull shape correlated significantly with tooth shape. High size-heterodonty often correlated with relatively large caniniform teeth, reflecting a prioritization of securing prey. Large, highly molariform, distal teeth may be a consequence of high-frequency durophagy combined with prey size. The slender-snouted skull shape correlated with a caniniform arcade with low heterodonty. This was reminiscent of other underwater-feeding tetrapods, as they often focus on small prey that requires minimal processing. Several extinct taxa were very molariform, which was associated with low heterodonty. The terrestrial peirosaurid shared similarities with large modern crocodylian taxa, but may have processed prey differently. Disparity measures can be inflated or deflated if certain teeth are absent from the tooth row, and regression analysis may not best apply to strongly slender-snouted taxa. Nevertheless, when these methods are used in tandem they can give a complete picture of crocodylian heterodonty. Future researchers may apply our proposed method to most crocodylian specimens with an intact enough tooth row regardless of age, species, or rearing conditions, as this will add rigor to many life history studies of the clade.

Captive broad snouted crocodylians are generally thought to have wider, shorter rostra than their wild counterparts. Interpreted to reflect morphological change in response to the conditions of captivity, this qualitative pattern could affect the utility of these animals in a variety of fields of research. However, due to relative ease of access and availability of life history data, captive animals are often utilized in actualistic research. Thus, this issue should be addressed in more detail. Here we explore snout shape variation between captive and wild members of Alligator mississippiensis using two-dimensional (2D) morphometric techniques. Several landmark schemesare used to assess the utility of different aspects of morphology in distinguishing the groups. While statistical analyses consistently differentiated between the groups, the area of morphospace occupied by wild members of A. mississippiensis generally overlapped with the larger area encompassing the captive specimens. This indicates that the captive condition is not as uniform as previously thought and instead encompasses a large spectrum of morphologies, ranging from the stereotypical broad, shortened snouts to outlines that are indistinguishable from the wild morphotype. These results align well with the interpretation that this change reflects an extreme example of ecophenotypy, since ranched, farmed, or zoo organisms are held in an array of enclosures, ranging from indoor, climate controlled pens to outdoor, more natural areas. This variation in environments should be reflected in different reactions to the animals' surroundings, resulting in a broad spectrum of morphotypes. While wild specimens are still preferred, especially for fine scale analyses, these results indicate that not all captive members of A. mississippiensis exhibit the extreme morphological alterations often cited in the literature. Weighing the conditions in which the animals are held and exploring the possibility of morphological differences against the benefits of using captive specimens should be part of any actualistic study. J. Morphol. 277:866-878, 2016. © 2016 Wiley Periodicals, Inc.

A new taxon of neosuchian crocodyliform, Deltasuchus motherali, gen. et. sp. nov., is described on the basis of a partial skull recovered from the Arlington Archosaur Site within the Upper Cretaceous (Cenomanian) Woodbine Formation of north central Texas. This productive locality represents a delta plain ecosystem preserving a diverse coastal fauna including lungfish, turtles, dinosaurs (ornithopods and theropods), and crocodyliforms. Prior to this discovery, the only identified crocodyliforms from the Woodbine Formation had been the longirostrine taxa Terminonaris and Woodbinesuchus. This new taxon is differentiated from other known crocodyliforms by the presence of dual pseudocanines on both the dentary and maxilla;; anterior and posterior rami of jugal comparable in depth; anterolaterally facing margin on the dorsal portion of the postorbital; contact between the descending process of the postorbital and the ectopterygoid; and a large, deep fossa on the ventral surface of the quadrate. Phylogenetic analysis recovers D. motherali as the sister taxon to Paluxysuchus newmani from the Lower Cretaceous Twin Mountains Formation of Texas. This clade lies within Neosuchia basal to Goniopholididae + Eusuchia. The associated cranial elements of this new crocodyliform represent a large, broad snouted individual, an ecomorphotype often associated with the semi-aquatic ambush predator niche in this clade, and one not previously reported from the formation.

Hypotheses of feeding behaviors and community structure are testable with rare direct evidence of trophic interactions in the fossil record (e.g., bite marks). We present evidence of four predation, scavenging, and/or interspecific fighting events involving two large paracrocodylomorphs (=‘rauisuchians’) from the Upper Triassic Chinle Formation (∼220–210 Ma). The larger femur preserves a rare history of interactions with multiple actors prior to and after death of this ∼8–9-m individual. A large embedded tooth crown and punctures, all of which display reaction tissue formed through healing, record evidence of a failed attack on this individual. The second paracrocodylomorph femur exhibits four unhealed bite marks, indicating the animal either did not survive the attack or was scavenged soon after death. The combination of character states observed (e.g., morphology of the embedded tooth, ‘D’-shaped punctures, evidence of bicarination of the marking teeth, spacing of potentially serial marks) indicates that large phytosaurs were actors in both cases. Our analysis of these specimens demonstrates phytosaurs targeted large paracrocodylomorphs in these Late Triassic ecosystems. Previous distinctions between ‘aquatic’ and ‘terrestrial’ Late Triassic trophic structures were overly simplistic and built upon mistaken paleoecological assumptions; we show they were intimately connected at the highest trophic levels. Our data also support that size cannot be the sole factor in determining trophic status. Furthermore, these marks provide an opportunity to start exploring the seemingly unbalanced terrestrial ecosystems from the Late Triassic of North America, in which large carnivores far outnumber herbivores in terms of both abundance and diversity.

Crocodyliforms serve as important taphonomic agents, accumulating and modifying vertebrate remains. Previousdiscussions of Mesozoic crocodyliform feeding in terrestrial and riverine ecosystems have often focused on larger taxa and their interactions with equally large dinosaurian prey. However, recent evidence suggests that the impact of smaller crocodyliforms on their environments should not be discounted. Here we present direct evidence of feeding by a small crocodyliform on juvenile specimens of a ‘hypsilophodontid’ dinosaur from the Upper Cretaceous (Campanian) Kaiparowits Formation of southern Utah. Diagnostic crocodyliform bite marks present on a left scapula and a right femur, as well as a partial probable crocodyliform tooth crown (ovoid in cross-section) preserved within a puncture on the right femur, comprise the bulk of the feeding evidence. Computed tomography scans of the femoral puncture reveal impact damage to the surrounding bone and that the distal tip of the embedded tooth was missing prior to the biting event. This is only the second reported incidence of a fossil crocodyliform tooth being found embedded directly into prey bone. These bite marks provide insight into the trophic interactions of the ecosystem preserved in the Kaiparowits Formation. The high diversity of crocodyliforms within this formation may have led to accentuated niche partitioning, which seems to have included juvenile dinosaurian prey.