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Global climate patterns fundamentally shape the distribution of species and ecosystems. For example, Bergmann's rule predicts that homeothermic animals, including birds and mammals, inhabiting cooler climates are generally larger than close relatives from warmer climates. The modern world, however, lacks the comparative data needed to evaluate such macroecological rules rigorously. Here, we test for Bergmann's rule in Mesozoic dinosaurs and mammaliaforms that radiated within relatively temperate global climate regimes. We develop a phylogenetic model that accounts for biases in the fossil record and allows for variable evolutionary dispersal rates. Our analysis also includes new fossil data from the extreme high-latitude Late Cretaceous Arctic Prince Creek Formation. We find no evidence for Bergmann's rule in Mesozoic dinosaurs or mammaliaforms, the ancestors of extant homeothermic birds and mammals. When our model is applied to thousands of extant dinosaur (bird) and mammal species, we find that body size evolution remains independent of latitude. A modest temperature effect is found in extant, but not in Mesozoic, birds, suggesting that body size evolution in modern birds was influenced by Bergmann's rule during Cenozoic climatic change. Our study provides a general approach for studying macroecological rules, highlighting the fossil record's power to address longstanding ecological principles., (© 2024. The Author(s).)

Tyrannosaurids were large carnivorous dinosaurs that underwent major changes in skull robusticity and body proportions as they grew, suggesting that they occupied different ecological niches during their life span. Although adults commonly fed on dinosaurian megaherbivores, the diet of juvenile tyrannosaurids is largely unknown. Here, we describe a remarkable specimen of a juvenile Gorgosaurus libratus that preserves the articulated hindlimbs of two yearling caenagnathid dinosaurs inside its abdominal cavity. The prey were selectively dismembered and consumed in two separate feeding events. This predator-prey association provides direct evidence of an ontogenetic dietary shift in tyrannosaurids. Juvenile individuals may have hunted small and young dinosaurs until they reached a size when, to satisfy energy requirements, they transitioned to feeding on dinosaurian megaherbivores. Tyrannosaurids occupied both mesopredator and apex predator roles during their life span, a factor that may have been key to their evolutionary success.

Phytoliths, microscopic deposits of hydrated silica within plants, play a myriad of functional roles in extant tracheophytes - yet their evolutionary origins and the original selective pressures leading to their deposition remain poorly understood. To gain new insights into the ancestral condition of tracheophyte phytolith production and function, phytolith content was intensively assayed in a basal, morphologically conserved tracheophyte: the foxtail clubmoss Lycopodiella alopecuroides. Wet ashing was employed to perform phytolith extractions from every major anatomical region of L. alopecuroides. Phytolith occurrence was recorded, alongside abundance, morphometric information, and morphological descriptions. Phytoliths were recovered exclusively from the microphylls, which were apicodistally silicified into multiphytolith aggregates. Phytolith aggregates were larger and more numerous in anatomical regions engaging in greater evapotranspirational activity. The tissue distribution of L. alopecuroides phytoliths is inconsistent with the expectations of proposed adaptive hypotheses of phytolith evolutionary origin. Instead, it is hypothesized that phytoliths may have arisen incidentally in the L. alopecuroides-like ancestral plant, polymerizing from intraplant silicon accumulations arising via bulk flow and 'leaky' cellular micronutrient channels. This basal, nonadaptive phytolith formation model would provide the evolutionary 'raw material' for later modification into the useful, adaptative, phytolith deposits seen in later-diverging plant clades., (© 2023 The Authors New Phytologist © 2023 New Phytologist Foundation.)

Beavers (Castor) stand out among mammals for their unique capacity to fell trees using their large, ever-growing incisors. This routine consumption of resistant fodder induces prodigious wear in the lower incisors, despite this blunting effect the incisors maintain a remarkably sharp cutting edge. Notably, the enamel edges of their incisors show a highly complex two-part microstructure of which the biomechanical import is unknown. Here, using fracture analysis, nanoindentation, and wear testing on North American beaver (C. canadensis) incisors we test the microstructure's possible contribution to maintaining incisal sharpness. Although comparable in hardness, the inner enamel preferentially fails and readily wears at 2.5 times the rate of the outer enamel. The outer microstructure redirects all fractures in parallel, decreasing fracture coalescence. Conversely, the inner microstructure facilitates crack coalescence increasing the wear rate by isolating layers of enamel prisms that readily fragment. Together these two architectures form a microstructurally driven self-sharpening mechanism contained entirely within the thin enamel shell. Our results demonstrate that enamel microstructures exposed at the occlusal surface can markedly influence both enamel crest shape and surface texture in wearing dentitions. The methods introduced here open the door to exploring the biomechanical functionality and evolution of enamel microstructures throughout Mammalia. STATEMENT OF SIGNIFICANCE: Enamel microstructure varies significantly with the diversity of diets, bite forces, and tooth shapes exhibited by mammals. However, minimal micromechanical exploration of microstructures outside of humans, leaves our understanding of biomechanical functions in a nascent stage. Using biologically informed mechanical testing, we demonstrate that the complex two-part microstructure that comprises the cutting edge of beaver incisors facilitates self-sharpening of the enamel edge. This previously unrecognized mechanism provides critical maintenance to the shape of the incisal edge ensuring continued functionality despite extreme wear incurred during feeding. More broadly, we show how the architecture of prisms and the surrounding interprismatic matrix dictate the propagation of fractures through enamel fabrics and how the pairing of enamel fabrics can result in biologically advantageous functions., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023. Published by Elsevier Ltd.)

Secondary aquatic adaptations evolved independently more than 30 times from terrestrial vertebrate ancestors 1,2 . For decades, non-avian dinosaurs were believed to be an exception to this pattern. Only a few species have been hypothesized to be partly or predominantly aquatic 3-11 . However, these hypotheses remain controversial 12,13 , largely owing to the difficulty of identifying unambiguous anatomical adaptations for aquatic habits in extinct animals. Here we demonstrate that the relationship between bone density and aquatic ecologies across extant amniotes provides a reliable inference of aquatic habits in extinct species. We use this approach to evaluate the distribution of aquatic adaptations among non-avian dinosaurs. We find strong support for aquatic habits in spinosaurids, associated with a marked increase in bone density, which precedes the evolution of more conspicuous anatomical modifications, a pattern also observed in other aquatic reptiles and mammals 14-16 . Spinosaurids are revealed to be aquatic specialists with surprising ecological disparity, including subaqueous foraging behaviour in Spinosaurus and Baryonyx, and non-diving habits in Suchomimus. Adaptation to aquatic environments appeared in spinosaurids during the Early Cretaceous, following their divergence from other tetanuran theropods during the Early Jurassic 17 ., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

The unexpected discovery of non-avian dinosaurs from Arctic and Antarctic settings has generated considerable debate about whether they had the capacity to reproduce at high latitudes-especially the larger-bodied, hypothetically migratory taxa. Evidence for dinosaurian polar reproduction remains very rare, particularly for species that lived at the highest paleolatitudes (>75°). Here we report the discovery of perinatal and very young dinosaurs from the highest known paleolatitude for the clade-the Cretaceous Prince Creek Formation (PCF) of northern Alaska. These data demonstrate Arctic reproduction in a diverse assemblage of large- and small-bodied ornithischian and theropod species. In terms of overall diversity, 70% of the known dinosaurian families, as well as avialans (birds), in the PCF are represented by perinatal individuals, the highest percentage for any North American Cretaceous formation. These findings, coupled with prolonged incubation periods, small neonate sizes, and short reproductive windows suggest most, if not all, PCF dinosaurs were nonmigratory year-round Arctic residents. Notably, we reconstruct an annual chronology of reproductive events for the ornithischian dinosaurs using refined paleoenvironmental/plant phenology data and new insights into dinosaur incubation periods. Seasonal resource limitations due to extended periods of winter darkness and freezing temperatures placed severe constraints on dinosaurian reproduction, development, and maintenance, suggesting these taxa showed polar-specific life history strategies, including endothermy., Competing Interests: Declaration of interests The authors declare no competing interests., (Crown Copyright © 2021. Published by Elsevier Inc. All rights reserved.)

Body size and body-size shifts broadly impact life-history parameters of all animals, which has made accurate body-size estimates for extinct taxa an important component of understanding their paleobiology. Among extinct crocodylians and their precursors (e.g., suchians), several methods have been developed to predict body size from suites of hard-tissue proxies. Nevertheless, many have limited applications due to the disparity of some major suchian groups and biases in the fossil record. Here, we test the utility of head width (HW) as a broadly applicable body-size estimator in living and fossil suchians. We use a dataset of sexually mature male and female individuals ( n  = 76) from a comprehensive sample of extant suchian species encompassing nearly all known taxa ( n  = 22) to develop a Bayesian phylogenetic model for predicting three conventional metrics for size: body mass, snout-vent length, and total length. We then use the model to estimate size parameters for a select series of extinct suchians with known phylogenetic affinity ( Montsechosuchus , Diplocynodon , and Sarcosuchus ). We then compare our results to sizes reported in the literature to exemplify the utility of our approach for a broad array of fossil suchians. Our results show that HW is highly correlated with all other metrics (all R 2 ≥0.85) and is commensurate with femoral dimensions for its reliably as a body-size predictor. We provide the R code in order to enable other researchers to employ the model in their own research., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.)

Medullary bone is an ephemeral type of bone tissue, today found only in sexually mature female birds, that provides a calcium reservoir for eggshell formation. The presence of medullary bone-like tissues in extant birds, pterosaurs, and dinosaurs distantly related to birds shows that caution must be exercised before concluding that fossils bear medullary bone. Here we describe a new specimen of pengornithid enantiornithine from the Lower Cretaceous Jiufotang Formation. Consisting of an isolated left hindlimb, the three-dimensional preservation contrasts with the crushed preservation characteristic of most Jehol specimens. Histological examinations suggest this resulted from the presence of a thick layer of highly vascular bone spanning the medullary cavities of the femur and tibiotarsus, consistent with expectations for medullary bone in extant birds. Micro-computed tomographic scans reveal small amounts of the same tissue extending into the pedal phalanges. We consider the tissue to be homologous to the medullary bone of Neornithines.

Most carnivorous mammals can pulverize skeletal elements by generating tooth pressures between occluding teeth that exceed cortical bone shear strength, thereby permitting access to marrow and phosphatic salts. Conversely, carnivorous reptiles have non-occluding dentitions that engender negligible bone damage during feeding. As a result, most reptilian predators can only consume bones in their entirety. Nevertheless, North American tyrannosaurids, including the giant (13 metres [m]) theropod dinosaur Tyrannosaurus rex stand out for habitually biting deeply into bones, pulverizing and digesting them. How this mammal-like capacity was possible, absent dental occlusion, is unknown. Here we analyzed T. rex feeding behaviour from trace evidence, estimated bite forces and tooth pressures, and studied tooth-bone contacts to provide the answer. We show that bone pulverization was made possible through a combination of: (1) prodigious bite forces (8,526-34,522 newtons [N]) and tooth pressures (718-2,974 megapascals [MPa]) promoting crack propagation in bones, (2) tooth form and dental arcade configurations that concentrated shear stresses, and (3) repetitive, localized biting. Collectively, these capacities and behaviors allowed T. rex to finely fragment bones and more fully exploit large dinosaur carcasses for sustenance relative to competing carnivores.

Birds stand out from other egg-laying amniotes by producing relatively small numbers of large eggs with very short incubation periods (average 11-85 d). This aspect promotes high survivorship by limiting exposure to predation and environmental perturbation, allows for larger more fit young, and facilitates rapid attainment of adult size. Birds are living dinosaurs; their rapid development has been considered to reflect the primitive dinosaurian condition. Here, nonavian dinosaurian incubation periods in both small and large ornithischian taxa are empirically determined through growth-line counts in embryonic teeth. Our results show unexpectedly slow incubation (2.8 and 5.8 mo) like those of outgroup reptiles. Developmental and physiological constraints would have rendered tooth formation and incubation inherently slow in other dinosaur lineages and basal birds. The capacity to determine incubation periods in extinct egg-laying amniotes has implications for dinosaurian embryology, life history strategies, and survivorship across the Cretaceous-Paleogene mass extinction event., Competing Interests: The authors declare no conflict of interest.

The cranial and postcranial anatomy of the basal hadrosaurid dinosaur Eotrachodon orientalis, from the uppermost Santonian of southern Appalachia (southeastern U.S.A.), is described in detail. This animal is the only known pre-Campanian non-lambeosaurine hadrosaurid, and the most complete hadrosauroid known from Appalachia. E. orientalis possesses a mosaic of plesiomorphic and derived characters in the context of Hadrosauroidea. Characters shared with basal hadrosauroids include a short and sloping maxillary ectopterygoid shelf, caudally prominent maxillary jugal process, one functional tooth per alveolus on the maxillary occlusal plane, a jugal rostral process with a shallow caudodorsal margin and medioventrally facing articular facet, a vertical dentary coronoid process with a poorly expanded apex, and tooth crowns with accessory ridges. Derived characters shared with other hadrosaurids include a circumnarial depression compartmented into three fossae (as in brachylophosaurins and Edmontosaurus), a thin everted premaxillary oral margin (as in Gryposaurus, Prosaurolophus, and Saurolophus), and a maxilla with a deep and rostrocaudally extensive rostrodorsal region with a steeply sloping premaxillary margin (as in Gryposaurus). Eotrachodon orientalis differs primarily from the other hadrosauroid from the Mooreville Chalk of Alabama, Lophorhothon atopus, in having a slender and crestless nasal whose caudodorsal margin is not invaded by the circumnarial depression. Hadrosaurus foulkii, the only other known hadrosaurid from Appalachia, is distinct from E. orientalis in having dentary teeth lacking accessory ridges and a dorsally curved shaft of the ischium. A histological section of the tibia of the E. orientalis holotype (MSC 7949) suggests that this individual was actively growing at the time of death and, thus, had the potential to become a larger animal later in development.

Herbivorous reptiles rarely evolve occluding dentitions that allow for the mastication (chewing) of plant matter. Conversely, most herbivorous mammals have occluding teeth with complex tissue architectures that self-wear to complex morphologies for orally processing plants. Dinosaurs stand out among reptiles in that several lineages acquired the capacity to masticate. In particular, the horned ceratopsian dinosaurs, among the most successful Late Cretaceous dinosaurian lineages, evolved slicing dentitions for the exploitation of tough, bulky plant matter. We show how Triceratops, a 9-m-long ceratopsian, and its relatives evolved teeth that wore during feeding to create fullers (recessed central regions on cutting blades) on the chewing surfaces. This unique morphology served to reduce friction during feeding. It was achieved through the evolution of a complex suite of osseous dental tissues rivaling the complexity of mammalian dentitions. Tribological (wear) properties of the tissues are preserved in ~66-million-year-old teeth, allowing the creation of a sophisticated three-dimensional biomechanical wear model that reveals how the complexes synergistically wore to create these implements. These findings, along with similar discoveries in hadrosaurids (duck-billed dinosaurs), suggest that tissue-mediated changes in dental morphology may have played a major role in the remarkable ecological diversification of these clades and perhaps other dinosaurian clades capable of mastication.

Background: Crocodilians have dominated predatory niches at the water-land interface for over 85 million years. Like their ancestors, living species show substantial variation in their jaw proportions, dental form and body size. These differences are often assumed to reflect anatomical specialization related to feeding and niche occupation, but quantified data are scant. How these factors relate to biomechanical performance during feeding and their relevance to crocodilian evolutionary success are not known., Methodology/principal Findings: We measured adult bite forces and tooth pressures in all 23 extant crocodilian species and analyzed the results in ecological and phylogenetic contexts. We demonstrate that these reptiles generate the highest bite forces and tooth pressures known for any living animals. Bite forces strongly correlate with body size, and size changes are a major mechanism of feeding evolution in this group. Jaw shape demonstrates surprisingly little correlation to bite force and pressures. Bite forces can now be predicted in fossil crocodilians using the regression equations generated in this research., Conclusions/significance: Critical to crocodilian long-term success was the evolution of a high bite-force generating musculo-skeletal architecture. Once achieved, the relative force capacities of this system went essentially unmodified throughout subsequent diversification. Rampant changes in body size and concurrent changes in bite force served as a mechanism to allow access to differing prey types and sizes. Further access to the diversity of near-shore prey was gained primarily through changes in tooth pressure via the evolution of dental form and distributions of the teeth within the jaws. Rostral proportions changed substantially throughout crocodilian evolution, but not in correspondence with bite forces. The biomechanical and ecological ramifications of such changes need further examination.

Differential scaling of musculoskeletal traits leads to differences in performance across ontogeny and ultimately determines patterns of resource use during development. Because musculoskeletal growth of the feeding system facilitates high bite-force generation necessary to overcome the physical constraints of consuming more durable prey, durophagous taxa are well suited for investigations of the scaling relationships between musculoskeletal growth, bite-force generation and dietary ontogeny. To elucidate which biomechanical factors are responsible for allometric changes in bite force and durophagy, we developed and experimentally tested a static model of bite-force generation throughout development in the durophagous turtle Sternotherus minor. Moreover, we quantified the fracture properties of snails found in the diet to evaluate the relationship between bite force and the forces required to process durable prey. We found that (1) the static bite-force model accurately predicts the ontogenetic scaling of bite forces, (2) bite-force positive allometry is accomplished by augmenting muscle size and muscle pennation, and (3) the rupture forces of snails found in the diet show a similar scaling pattern to bite force across ontogeny. These results indicate the importance of muscle pennation for generating high bite forces while maintaining muscle size and provide empirical evidence that the allometric patterns of musculoskeletal growth in S. minor are strongly linked to the structural properties of their primary prey.

Background: Tyrannosaurus rex was one of the largest terrestrial carnivores of all time, and consequently its ecology and diet have been the focus of much discussion. However, there is little direct evidence of diet or feeding habits in this species., Methodology/principal Findings: Examination of museum collections has revealed four specimens of Tyrannosaurus rex that bear tooth marks made by large, carnivorous dinosaurs. Because Tyrannosaurus is the only large carnivore known from the Late Maastrichtian of western North America, we infer that Tyrannosaurus made these tooth marks., Conclusions/significance: The marks are interpreted as feeding traces and these fossils therefore record instances of cannibalism. Given that this behavior has a low preservation potential, cannibalism seems to have been a surprisingly common behavior in Tyrannosaurus, and this behavior may have been relatively common in carnivorous dinosaurs.

Tyrannosaurs, the group of dinosaurian carnivores that includes Tyrannosaurus rex and its closest relatives, are icons of prehistory. They are also the most intensively studied extinct dinosaurs, and thanks to large sample sizes and an influx of new discoveries, have become ancient exemplar organisms used to study many themes in vertebrate paleontology. A phylogeny that includes recently described species shows that tyrannosaurs originated by the Middle Jurassic but remained mostly small and ecologically marginal until the latest Cretaceous. Anatomical, biomechanical, and histological studies of T. rex and other derived tyrannosaurs show that large tyrannosaurs could not run rapidly, were capable of crushing bite forces, had accelerated growth rates and keen senses, and underwent pronounced changes during ontogeny. The biology and evolutionary history of tyrannosaurs provide a foundation for comparison with other dinosaurs and living organisms.

The extremes of dinosaur body size have long fascinated scientists. The smallest (<1 m length) known dinosaurs are carnivorous saurischian theropods, and similarly diminutive herbivorous or omnivorous ornithischians (the other major group of dinosaurs) are unknown. We report a new ornithischian dinosaur, Fruitadens haagarorum, from the Late Jurassic of western North America that rivals the smallest theropods in size. The largest specimens of Fruitadens represent young adults in their fifth year of development and are estimated at just 65-75 cm in total body length and 0.5-0.75 kg body mass. They are thus the smallest known ornithischians. Fruitadens is a late-surviving member of the basal dinosaur clade Heterodontosauridae, and is the first member of this clade to be described from North America. The craniodental anatomy and diminutive body size of Fruitadens suggest that this taxon was an ecological generalist with an omnivorous diet, thus providing new insights into morphological and palaeoecological diversity within Dinosauria. Late-surviving (Late Jurassic and Early Cretaceous) heterodontosaurids are smaller and less ecologically specialized than Early (Late Triassic and Early Jurassic) heterodontosaurids, and this ecological generalization may account in part for the remarkable 100-million-year-long longevity of the clade.

Ornithomimosaurs (ostrich-mimic dinosaurs) are a common element of some Cretaceous dinosaur assemblages of Asia and North America. Here, we describe a new species of ornithomimosaur, Beishanlong grandis, from an associated, partial postcranial skeleton from the Aptian-Albian Xinminpu Group of northern Gansu, China. Beishanlong is similar to another Aptian-Albian ornithomimosaur, Harpymimus, with which it shares a phylogenetic position as more derived than the Barremian Shenzhousaurus and as sister to a Late Cretaceous clade composed of Garudimimus and the Ornithomimidae. Beishanlong is one of the largest definitive ornithomimosaurs yet described, though histological analysis shows that the holotype individual was still growing at its death. Together with the co-eval and sympatric therizinosaur Suzhousaurus and the oviraptorosaur Gigantraptor, Beishanlong provides evidence for the parallel evolution of gigantism in separate lineages of beaked and possibly herbivorous coelurosaurs within a short time span in Central Asia.

Tyrannosaurid theropods are characterized by a generalized body plan, and all well-known taxa possess deep and robust skulls that are optimized for exerting powerful bite forces. The fragmentary Late Cretaceous Alioramus appears to deviate from this trend, but its holotype and only known specimen is incomplete and poorly described. A remarkable new tyrannosaurid specimen from the Maastrichtian (Late Cretaceous) of Mongolia, including a nearly complete and well-preserved skull and an extensive postcranium, represents a new species of Alioramus, Alioramus altai. This specimen conclusively demonstrates that Alioramus is a small, gracile, long-snouted carnivore that deviates from other tyrannosaurids in its body plan and presumably its ecological habits. As such, it increases the range of morphological diversity in one of the most familiar extinct clades. Phylogenetic analysis places Alioramus deep within the megapredatory Tyrannosauridae, and within the tyrannosaurine subclade that also includes Tarbosaurus and Tyrannosaurus. Both pneumatization and ornamentation are extreme compared with other tyrannosaurids, and the skull contains eight discrete horns. The new specimen is histologically aged at nine years old but is smaller than other tyrannosaurids of similar age. Despite its divergent cranial form, Alioramus is characterized by a similar sequence of ontogenetic changes as the megapredatory Tyrannosaurus and Albertosaurus, indicating that ontogenetic change is conservative in tyrannosaurids.

Background: Archaeopteryx is the oldest and most primitive known bird (Avialae). It is believed that the growth and energetic physiology of basalmost birds such as Archaeopteryx were inherited in their entirety from non-avialan dinosaurs. This hypothesis predicts that the long bones in these birds formed using rapidly growing, well-vascularized woven tissue typical of non-avialan dinosaurs., Methodology/principal Findings: We report that Archaeopteryx long bones are composed of nearly avascular parallel-fibered bone. This is among the slowest growing osseous tissues and is common in ectothermic reptiles. These findings dispute the hypothesis that non-avialan dinosaur growth and physiology were inherited in totality by the first birds. Examining these findings in a phylogenetic context required intensive sampling of outgroup dinosaurs and basalmost birds. Our results demonstrate the presence of a scale-dependent maniraptoran histological continuum that Archaeopteryx and other basalmost birds follow. Growth analysis for Archaeopteryx suggests that these animals showed exponential growth rates like non-avialan dinosaurs, three times slower than living precocial birds, but still within the lowermost range for all endothermic vertebrates., Conclusions/significance: The unexpected histology of Archaeopteryx and other basalmost birds is actually consistent with retention of the phylogenetically earlier paravian dinosaur condition when size is considered. The first birds were simply feathered dinosaurs with respect to growth and energetic physiology. The evolution of the novel pattern in modern forms occurred later in the group's history.

Theropods have traditionally been assumed to have lost manual digits from the lateral side inward, which differs from the bilateral reduction pattern seen in other tetrapod groups. This unusual reduction pattern is clearly present in basal theropods, and has also been inferred in non-avian tetanurans based on identification of their three digits as the medial ones of the hand (I-II-III). This contradicts the many developmental studies indicating II-III-IV identities for the three manual digits of the only extant tetanurans, the birds. Here we report a new basal ceratosaur from the Oxfordian stage of the Jurassic period of China (156-161 million years ago), representing the first known Asian ceratosaur and the only known beaked, herbivorous Jurassic theropod. Most significantly, this taxon possesses a strongly reduced manual digit I, documenting a complex pattern of digital reduction within the Theropoda. Comparisons among theropod hands show that the three manual digits of basal tetanurans are similar in many metacarpal features to digits II-III-IV, but in phalangeal features to digits I-II-III, of more basal theropods. Given II-III-IV identities in avians, the simplest interpretation is that these identities were shared by all tetanurans. The transition to tetanurans involved complex changes in the hand including a shift in digit identities, with ceratosaurs displaying an intermediate condition.

The repeated discovery of adult dinosaurs in close association with egg clutches leads to speculation over the type and extent of care exhibited by these extinct animals for their eggs and young. To assess parental care in Cretaceous troodontid and oviraptorid dinosaurs, we examined clutch volume and the bone histology of brooding adults. In comparison to four archosaur care regressions, the relatively large clutch volumes of Troodon, Oviraptor, and Citipati scale most closely with a bird-paternal care model. Clutch-associated adults lack the maternal and reproductively associated histologic features common to extant archosaurs. Large clutch volumes and a suite of reproductive features shared only with birds favor paternal care, possibly within a polygamous mating system. Paternal care in both troodontids and oviraptorids indicates that this care system evolved before the emergence of birds and represents birds' ancestral condition. In extant birds and over most adult sizes, paternal and biparental care correspond to the largest and smallest relative clutch volumes, respectively.

The timing of sexual maturation in non-avian dinosaurs is not known. In extant squamates and crocodilians it occurs in conjunction with the initial slowing of growth rates as adult size is approached. In birds (living dinosaurs) on the other hand, reproductive activity begins well after somatic maturity. Here we used growth line counts and spacing in all of the known brooding non-avian dinosaurs to determine the stages of development when they perished. It was revealed that sexual maturation occurred well before full adult size was reached-the primitive reptilian condition. In this sense, the life history and physiology of non-avian dinosaurs was not like that of modern birds. Palaeobiological ramifications of these findings include the potential to deduce reproductive lifespan, fecundity and reproductive population sizes in non-avian dinosaurs, as well as aid in the identification of secondary sexual characteristics.

Fossil evidence for changes in dinosaurs near the lineage leading to birds and the origin of flight has been sparse. A dinosaur from Mongolia represents the basal divergence within Dromaeosauridae. The taxon's small body size and phylogenetic position imply that extreme miniaturization was ancestral for Paraves (the clade including Avialae, Troodontidae, and Dromaeosauridae), phylogenetically earlier than where flight evolution is strongly inferred. In contrast to the sustained small body sizes among avialans throughout the Cretaceous Period, the two dinosaurian lineages most closely related to birds, dromaeosaurids and troodontids, underwent four independent events of gigantism, and in some lineages size increased by nearly three orders of magnitude. Thus, change in theropod body size leading to flight's origin was not unidirectional.

Direct evidence of prey choice in carnivorous dinosaurs is rare in the fossil record. The most celebrated example pertains to purported stomach contents in the carnivorous dinosaur Coelophysis bauri, which besides revealing prey choice, also points to cannibalistic behaviour as being commonplace (Colbert 1989, 1995). Here, we test this hypothesis by conducting the first comprehensive anatomical and histological examination of the famed Coelophysis 'cannibals'. The results unequivocally show that the gut contents derive from early crocodylomorphs rather than juveniles of Coelophysis. These findings suggest that this taxon is not cannibalistic and bring into question the commonality of this behaviour among non-avian dinosaurs.

The size and age structures for four assemblages of North American tyrannosaurs-Albertosaurus, Tyrannosaurus, Gorgosaurus, and Daspletosaurus-reveal a pronounced, bootstrap-supported pattern of age-specific mortality characterized by relatively high juvenile survivorship and increased mortality at midlife and near the maximum life span. Such patterns are common today in wild populations of long-lived birds and mammals. Factors such as predation and entrance into the breeding population may have influenced tyrannosaur survivorship. This survivorship pattern can explain the rarity of juvenile specimens in museum collections.

The tyrannosauroid fossil record is mainly restricted to Cretaceous sediments of Laurasia, although some very fragmentary Jurassic specimens have been referred to this group. Here we report a new basal tyrannosauroid, Guanlong wucaii gen. et sp. nov., from the lower Upper Jurassic of the Junggar Basin, northwestern China. G. wucaii is the oldest known tyrannosauroid and shows several unexpectedly primitive pelvic features. Nevertheless, the limbs of G. wucaii share several features with derived coelurosaurs, and it possesses features shared by other coelurosaurian clades. This unusual combination of character states provides an insight into the poorly known early radiation of the Coelurosauria. Notably, the presumed predatory Guanlong has a large, fragile and highly pneumatic cranial crest that is among the most elaborate known in any non-avian dinosaur and could be comparable to some classical exaggerated ornamental traits among vertebrates.

Long-standing controversy surrounds the question of whether living bird lineages emerged after non-avian dinosaur extinction at the Cretaceous/Tertiary (K/T) boundary or whether these lineages coexisted with other dinosaurs and passed through this mass extinction event. Inferences from biogeography and molecular sequence data (but see ref. 10) project major avian lineages deep into the Cretaceous period, implying their 'mass survival' at the K/T boundary. By contrast, it has been argued that the fossil record refutes this hypothesis, placing a 'big bang' of avian radiation only after the end of the Cretaceous. However, other fossil data--fragmentary bones referred to extant bird lineages--have been considered inconclusive. These data have never been subjected to phylogenetic analysis. Here we identify a rare, partial skeleton from the Maastrichtian of Antarctica as the first Cretaceous fossil definitively placed within the extant bird radiation. Several phylogenetic analyses supported by independent histological data indicate that a new species, Vegavis iaai, is a part of Anseriformes (waterfowl) and is most closely related to Anatidae, which includes true ducks. A minimum of five divergences within Aves before the K/T boundary are inferred from the placement of Vegavis; at least duck, chicken and ratite bird relatives were coextant with non-avian dinosaurs.

How evolutionary changes in body size are brought about by variance in developmental timing and/or growth rates (also known as heterochrony) is a topic of considerable interest in evolutionary biology. In particular, extreme size change leading to gigantism occurred within the dinosaurs on multiple occasions. Whether this change was brought about by accelerated growth, delayed maturity or a combination of both processes is unknown. A better understanding of relationships between non-avian dinosaur groups and the newfound capacity to reconstruct their growth curves make it possible to address these questions quantitatively. Here we study growth patterns within the Tyrannosauridae, the best known group of large carnivorous dinosaurs, and determine the developmental means by which Tyrannosaurus rex, weighing 5,000 kg and more, grew to be one of the most enormous terrestrial carnivorous animals ever. T. rex had a maximal growth rate of 2.1 kg d(-1), reached skeletal maturity in two decades and lived for up to 28 years. T. rex's great stature was primarily attained by accelerating growth rates beyond that of its closest relatives.

Did dinosaurs grow in a manner similar to extant reptiles, mammals or birds, or were they unique? Are rapid avian growth rates an innovation unique to birds, or were they inherited from dinosaurian precursors? We quantified growth rates for a group of dinosaurs spanning the phylogenetic and size diversity for the clade and used regression analysis to characterize the results. Here we show that dinosaurs exhibited sigmoidal growth curves similar to those of other vertebrates, but had unique growth rates with respect to body mass. All dinosaurs grew at accelerated rates relative to the primitive condition seen in extant reptiles. Small dinosaurs grew at moderately rapid rates, similar to those of marsupials, but large species attained rates comparable to those of eutherian mammals and precocial birds. Growth in giant sauropods was similar to that of whales of comparable size. Non-avian dinosaurs did not attain rates like those of altricial birds. Avian growth rates were attained in a stepwise fashion after birds diverged from theropod ancestors in the Jurassic period.

Dinosaur dentine exhibits growth lines that are tens of micrometers in width. These laminations are homologous to incremental lines of von Ebner found in extant mammal and crocodilian teeth (i.e., those of amniotes). The lines likely reflect daily dentine formation, and they were used to infer tooth development and replacement rates. In general, dinosaur tooth formation rates negatively correlated with tooth size. Theropod tooth replacement rates negatively correlated with tooth size, which was due to limitations in the dentine formation rates of their odontoblasts. Derived ceratopsian and hadrosaurian dinosaurs retained relatively rapid tooth replacement rates through ontogeny. The evolution of dental batteries in hadrosaurs and ceratopsians can be explained by dentine formation constraints and rapid tooth wear. In combination with counts of shed dinosaur teeth, tooth replacement rate data can be used to assess population demographics of Mesozoic ecosystems. Finally, it is of historic importance to note that Richard Owen appears to have been the first to observe incremental lines of von Ebner in dinosaurs more than 150 years ago.

Two diets were formulated to contain 30% NDF (DM basis) but differed in source of NDF and type of concentrate. One diet contained approximately 70% alfalfa silage and 30% corn grain (89% of NDF from forage) and the other diet contained about 50% alfalfa silage and 50% barley grain (65% of NDF from forage). Diets were fed to 12 cows in a short production trial, a total collection digestion experiment, and a rate of passage study. Cows fed the alfalfa-corn diet produced more 4% FCM (22.4 vs. 20.7 kg/d) than did cows fed the alfalfa-barley diet. Intake of DM (3.7% of BW), NE1 (35 Mcal), and NDF (1.1% of BW) were similar between diets, but due to differences in diet composition, intake of ADF was greater by cows fed alfalfa-corn than by cows fed alfalfa-barley. Digestibilities of DM, NDF, and CP were not affected by diet, but hemicellulose digestion was higher and cellulose and ADF digestibilities were lower by cows fed alfalfa-barley diets. Cows fed alfalfa-corn diet had a higher ruminal acetate to propionate molar ratio. No effect of diet was observed on turnover kinetics of the forage, concentrate, or liquid markers. These data show that forage to grain ratio and source of starch must be considered prior to recommending that diets be balanced for NDF.

This study identifies and analyzes factors under a hospital's control that can affect its market share. The study, utilizing the Multiplicative Competitive Interaction model, specifically focuses on determining the market share for each hospital within a geographic area, as opposed to the total demand for hospital services within an area. The results indicate that the effect of the number of physician affiliations on hospital patient share is statistically significant. The article investigates the variables that affect the level of physician affiliation. Besides physician affiliation, hospital location, a PROFILE factor based on a composite of a number of variables, and the proportion of affiliated physicians who are not affiliated elsewhere have a significant impact on each hospital's market share. The variables examined resulted in an R2 = 0.901 for individual hospital patient market share.

Dried distillers grains produced from a mix of 65% barley and 35% corn were evaluated in digestion and lactation experiments. Dried barley distillers grains had 56% NDF, 29% CP, 3% amino acid N, 2.5% NDIN (55% of total N), and 1.8% ADIN (39% of total N). Wet barley distillers grains had 38% NDF, 27% CP, 2.7% amino acid N, .5% NDIN (12% of total N), and .8% ADIN (19% of total N). Digestibility of DM and N was similar among lactating dairy cows fed diets containing approximately 25% corn silage DM, 15% alfalfa silage DM, 15% alfalfa hay DM, plus varying amounts of a corn-barley concentrate mix and supplemental CP from soybean meal, barley distillers grains, or from 1:1 mixture of soybean meal and barley distillers grains. Digestibility of ADIN, NDF, and ADF increased with increasing amounts of barley distillers grains in the diet. Similar diets were fed to 60 Holstein cows for 84 d in a lactation experiment. Source of supplemental protein did not affect milk production (22.5 kg/d), FCM (20.4 kg/d), milk fat percent (3.6%), or DM intake (19.0 kg/d). Milk protein percent was decreased by feeding barley distillers grains. It was concluded that barley distillers grains were an acceptable protein source for dairy cows and that ADIN and NDF might not be appropriate measures of the nutritional value of this product.