A newly recognized theropod assemblage from the Lewisville Formation (Woodbine Group; Cenomanian) and its implications for understanding Late Cretaceous Appalachian terrestrial ecosystems.

While the terrestrial fossil record of the mid-Cretaceous interval (Aptian to Cenomanian) in North America has been poorly studied, the recent focus on fossil localities from the western United States has offered a more detailed picture of vertebrate diversity, ecosystem dynamics and faunal turnover that took place on the western landmass of Laramidia. This is in stark contrast to the terrestrial record from the eastern landmass of Appalachia, where vertebrate fossils are rare and consist mostly of isolated and fragmentary remains. However, a detailed understanding of these fossil communities during this interval is necessary for comparison of the faunal patterns that developed during the opening of the Western Interior Seaway (WIS). The Woodbine Group of Texas is a Cenomanian age (95-100 mya) deposit consisting of shallow marine, deltaic, and terrestrial communities, which were only recently separated from their western counterparts. These deposits have yielded a wealth of vertebrate remains, yet non-avian theropods are still largely unknown. Recently, multiple localities in the Lewisville Formation of the Woodbine Group have yielded new non-avian theropod material, including numerous isolated teeth and postcranial remains. While largely fragmentary, this material is sufficiently diagnostic to identify the following taxa: a large-bodied carcharodontosaur, a mid-sized tyrannosauroid, a large ornithomimosaur, a large dromaeosaurine, a small dromaeosaurid, a small troodontid, and a small coelurosaur. Some of these groups represent the first occurrence for Appalachia and provide a broader understanding of a newly expanded faunal diversity for the Eastern landmass. The Lewisville Formation theropod fauna is similar in taxonomic composition to contemporaneous deposits in Laramidia, confirming that these groups were widespread across the continent prior to extension of the WIS. The Lewisville Formation documents the transitional nature of Cenomanian coastal ecosystems in Texas while providing additional details on the evolution of Appalachian communities shortly after WIS extension.

High frequencies of theropod bite marks provide evidence for feeding, scavenging, and possible cannibalism in a stressed Late Jurassic ecosystem.

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.

Quantitative heterodonty in Crocodylia: assessing size and shape across modern and extinct taxa.

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.

Claw morphometrics in monitor lizards: Variable substrate and habitat use correlate to shape diversity within a predator guild.

Numerous studies investigate morphology in the context of habitat, and lizards have received particular attention. Substrate usage is often reflected in the morphology of characters associated with locomotion, and, as a result, claws have become well-studied ecomorphological traits linking the two. The Kimberley predator guild of Western Australia consists of 10 sympatric varanid species. The purpose of this study was to quantify claw size and shape in the guild using geometric morphometrics, and determine whether these features correlated with substrate use and habitat. Each species was assigned a Habitat/substrate group based on the substrate their claws interact with in their respective habitat. Claw morphometrics were derived for both wild caught and preserved specimens from museum collections, using a 2D semilandmark analysis. Claw shape significantly separated based on Habitat/substrate group. Varanus gouldii and Varanus panoptes claws were associated with sprinting and extensive digging. Varanus mertensi claws were for shallow excavation. The remaining species' claws reflected specialization for some form of climbing, and differed based on substrate compliance. Varanus glauerti was best adapted for climbing rough sandstone, whereas Varanus scalaris and Varanus tristis had claws ideal for puncturing wood. Phylogenetic signal also significantly influenced claw shape, with Habitat/substrate group limited to certain clades. Positive size allometry allowed for claws to cope with mass increases, and shape allometry reflected a potential size limit on climbing. Claw morphology may facilitate niche separation within this trophic guild, especially when considered with body size. As these varanids are generalist predators, morphological traits associated with locomotion may be more reliable candidates for detecting niche partitioning than those associated directly with diet.

The Dry Season Shuffle: Gorges Provide Refugia for Animal Communities in Tropical Savannah Ecosystems.

In the wet-dry tropics, animal species face the major challenges of acquiring food, water or shelter during an extended dry season. Although large and conspicuous animals such as ungulates and waterfowl migrate to wetter areas during this time, little is known of how smaller and more cryptic animal species with less mobility meet these challenges. We fenced off the entire entrance of a gorge in the Australian tropical savanna, offering the unique opportunity to determine the composition and seasonal movement patterns of the small vertebrate community. The 1.7 km-long fence was converted to a trapline that was deployed for 18-21 days during the early dry season in each of two years, and paired traps on both sides of the fence allowed us to detect the direction of animal movements. We predicted that semi-aquatic species (e.g., frogs and turtles) would move upstream into the wetter gorge during the dry season, while more terrestrial species (e.g., lizards, snakes, mammals) would not. The trapline captured 1590 individual vertebrates comprising 60 species. There was a significant bias for captures on the outside of the fence compared to the inside for all species combined (outside/inside = 5.2, CI = 3.7-7.2), for all vertebrate classes, and for specific taxonomic groups. The opposite bias (inside/outside = 7.3, N= 25) for turtles during the early wet season suggested return migration heading into the wet season. Our study revealed that the small vertebrate community uses the gorge as a dry season refuge. The generality of this unreplicated finding could be tested by extending this type of survey to tropical savannahs worldwide. A better understanding of how small animals use the landscape is needed to reveal the size of buffer zones around wetlands required to protect both semi-aquatic and terrestrial fauna in gorges in tropical savannah woodland, and thus in ecosystems in general.

Illustrating ontogenetic change in the dentition of the Nile monitor lizard, Varanus niloticus: a case study in the application of geometric morphometric methods for the quantification of shape-size heterodonty.

Many recent attempts have been made to quantify heterodonty in non-mammalian vertebrates, but the majority of these are limited to Euclidian measurements. One taxon frequently investigated is Varanus niloticus, the Nile monitor. Juveniles possess elongate, pointed teeth (caniniform) along the entirety of the dental arcade, whereas adults develop large, bulbous distal teeth (molariform). The purpose of this study was to present a geometric morphometric method to quantify V. niloticus heterodonty through ontogeny that may be applied to other non-mammalian taxa. Data were collected from the entire tooth row of 19 dry skull specimens. A semilandmark analysis was conducted on the outline of the photographed teeth, and size and shape were derived. Width was also measured with calipers. From these measures, sample ranges and allometric functions were created using multivariate statistical analyses for each tooth position separately, as well as overall measures of heterodonty for each specimen based on morphological disparity. The results confirm and expand upon previous studies, showing measurable shape-size heterodonty in the species with significant differences at each tooth position. Tooth size increases with body size at most positions, and the allometric coefficient increases at more distal positions. Width shows a dramatic increase at the distal positions with ontogeny, often displaying pronounced positive allometry. Dental shape varied in two noticeable ways, with the first composing the vast majority of shape variance: (i) caniniformy vs. molariformy and (ii) mesially leaning, 'rounded' apices vs. distally leaning, 'pointed' apices. The latter was twice as influential in the mandible, a consequence of host bone shape. Mesial teeth show no significant shape change with growth, whereas distal teeth change significantly due primarily to an increase in molariformy. Overall, heterodonty increases with body size concerning both tooth size and shape, but shape heterodonty changes in the mandible are much less pronounced. Although it is unclear to what degree V. niloticus specializes in hard prey items (durophagy), previous studies of varanid feeding behavior, along with research on analogous durophagous vertebrates, indicate a division of labor along the tooth row in adults, due to a possible transition to at least a partial durophagous niche. The geometric morphometric method proposed here, although not without its own limitations, may be ideal for use with a number of dental morphotypes in the future.

Learning style preferences and academic success of preclinical allied health students.

Student learning style modality preferences, in preclinical classes, were assessed using the visual-aural-read/write-kinesthetic (VARK) inventory. Preferences were assessed for 137 preclinical students, including those in nursing, physician's assistant, physical therapy, athletic training, and natural science programs using the online VARK inventory. All classes contained a majority of multimodal and a significantly high proportion of kinesthetic learners. No correlations were noted between modality preference strength and assessment performance in general biology classes; significant correlations were discovered for kinesthetic preference among the same cohort in subsequent human anatomy (negative correlation) and general physiology (positive correlation) classes. Assessment performance of nursing students in an anatomy and physiology class resulted in correlations with aural (negative correlation) and visual (positive correlation) preference strengths. Study findings are used to evaluate the efficacy of non-omnimodal delivery of content-focused science classes, before the students have developed the background knowledge or skills required to contextualize the learning.

The effects of biting and pulling on the forces generated during feeding in the Komodo dragon (Varanus komodoensis).

In addition to biting, it has been speculated that the forces resulting from pulling on food items may also contribute to feeding success in carnivorous vertebrates. We present an in vivo analysis of both bite and pulling forces in Varanus komodoensis, the Komodo dragon, to determine how they contribute to feeding behavior. Observations of cranial modeling and behavior suggest that V. komodoensis feeds using bite force supplemented by pulling in the caudal/ventrocaudal direction. We tested these observations using force gauges/transducers to measure biting and pulling forces. Maximum bite force correlates with both body mass and total body length, likely due to increased muscle mass. Individuals showed consistent behaviors when biting, including the typical medial-caudal head rotation. Pull force correlates best with total body length, longer limbs and larger postcranial motions. None of these forces correlated well with head dimensions. When pulling, V. komodoensis use neck and limb movements that are associated with increased caudal and ventral oriented force. Measured bite force in Varanus komodoensis is similar to several previous estimations based on 3D models, but is low for its body mass relative to other vertebrates. Pull force, especially in the ventrocaudal direction, would allow individuals to hunt and deflesh with high success without the need of strong jaw adductors. In future studies, pull forces need to be considered for a complete understanding of vertebrate carnivore feeding dynamics.

A functional explanation for denticulation in theropod dinosaur teeth.

The serrated, or denticulated, ziphodont teeth of theropod dinosaurs display variability in their extent of denticulation. The functional model proposed here tests the hypothesis that denticles will not exist in areas that do not frequently contact the substrate. This area, defined as the "dead-space," is determined by the direction the tooth moves through the fleshy substrate. The extent of denticulation, as well as the dead-space dimensions, is measured from photographs of 235 isolated and in situ theropod teeth, to determine a meaningful relationship between the two variables. Both Euclidean and geometric morphometric methods are employed, and the data are expressed in bivariate and ordination plots. The model predicts the direction of tooth movement through the curvature of the tip/apex. Tooth position and taxon are considered. The results show that the mesial margin is usually partially denticulated, while the distal margin is usually totally denticulated. Curved teeth have large dead-spaces, and tend to be less denticulated mesially. Straighter teeth are more extensively denticulated, to the point where they became symmetrical. The mesial denticulation is determined by the dead-space, and dictated by the substrate contact. The dead-space almost always predicted less extensive denticulation; a consequence of the model's limitations. Tooth curvature increases with a more distal position, due to rotation based on the proximity to the hinge. Denticulation indicates that theropods used a distally oriented puncture to modify the substrate, similar to modern analogues. Although there is little taxonomic variation, Troodontidae show unique and extreme degrees of mesial denticulation.

Cranial performance in the Komodo dragon (Varanus komodoensis) as revealed by high-resolution 3-D finite element analysis.

The Komodo dragon (Varanus komodoensis) displays a unique hold and pull-feeding technique. Its delicate 'space-frame' skull morphology differs greatly from that apparent in most living large prey specialists and is suggestive of a high degree of optimization, wherein use of materials is minimized. Here, using high-resolution finite element modelling based on dissection and in vivo bite and pull data, we present results detailing the mechanical performance of the giant lizard's skull. Unlike most modern predators, V. komodoensis applies minimal input from the jaw muscles when butchering prey. Instead it uses series of actions controlled by postcranial muscles. A particularly interesting feature of the performance of the skull is that it reveals considerably lower overall stress when these additional extrinsic forces are added to those of the jaw adductors. This remarkable reduction in stress in response to additional force is facilitated by both internal and external bone anatomy. Functional correlations obtained from these analyses also provide a solid basis for the interpretation of feeding ecology in extinct species, including dinosaurs and sabre-tooth cats, with which V. komodoensis shares various cranial and dental characteristics.