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Megaherbivorous dinosaurs were exceptionally diverse on the Late Cretaceous island continent of Laramidia, and a growing body of evidence suggests that this diversity was facilitated by dietary niche partitioning. We test this hypothesis using the fossil megaherbivore assemblage from the Dinosaur Park Formation (upper Campanian) of Alberta as a model. Comparative tooth morphology and wear, including the first use of quantitative dental microwear analysis in the context of Cretaceous palaeosynecology, are used to infer the mechanical properties of the foods these dinosaurs consumed. The phylliform teeth of ankylosaurs were poorly adapted for habitually processing high-fibre plant matter. Nevertheless, ankylosaur diets were likely more varied than traditionally assumed: the relatively large, bladed teeth of nodosaurids would have been better adapted to processing a tougher, more fibrous diet than the smaller, cusp-like teeth of ankylosaurids. Ankylosaur microwear is characterized by a preponderance of pits and scratches, akin to modern mixed feeders, but offers no support for interspecific dietary differences. The shearing tooth batteries of ceratopsids are much better adapted to high-fibre herbivory, attested by their scratch-dominated microwear signature. There is tentative microwear evidence to suggest differences in the feeding habits of centrosaurines and chasmosaurines, but statistical support is not significant. The tooth batteries of hadrosaurids were capable of both shearing and crushing functions, suggestive of a broad dietary range. Their microwear signal overlaps broadly with that of ankylosaurs, and suggests possible dietary differences between hadrosaurines and lambeosaurines. Tooth wear evidence further indicates that all forms considered here exhibited some degree of masticatory propaliny. Our findings reveal that tooth morphology and wear exhibit different, but complimentary, dietary signals that combine to support the hypothesis of dietary niche partitioning. The inferred mechanical and dietary patterns appear constant over the 1.5 Myr timespan of the Dinosaur Park Formation megaherbivore chronofauna, despite continual species turnover.

Megaherbivorous dinosaur coexistence on the Late Cretaceous island continent of Laramidia has long puzzled researchers, owing to the mystery of how so many large herbivores (6-8 sympatric species, in many instances) could coexist on such a small (4-7 million km(2)) landmass. Various explanations have been put forth, one of which-dietary niche partitioning-forms the focus of this study. Here, we apply traditional morphometric methods to the skulls of megaherbivorous dinosaurs from the Dinosaur Park Formation (upper Campanian) of Alberta to infer the ecomorphology of these animals and to test the niche partitioning hypothesis. We find evidence for niche partitioning not only among contemporaneous ankylosaurs, ceratopsids, and hadrosaurids, but also within these clades at the family and subfamily levels. Consubfamilial ceratopsids and hadrosaurids differ insignificantly in their inferred ecomorphologies, which may explain why they rarely overlap stratigraphically: interspecific competition prevented their coexistence.

The Hangenberg extinction has been hypothesized as a first order event in vertebrate evolution; however, information on the earliest Carboniferous vertebrate fauna, crucial in evaluating biodiversity changes, is scarce. Post-extinction recovery has been suggested as the driver of ray-finned fish (actinopterygian) richness increase and differentiation in the Carboniferous. Under this model, actinopterygian postcranial morphology differentiates in the second stage of their radiation. Here, we report on a platysomid occurrence from the Tournaisian of Nova Scotia, Canada. Despite long-standing taxonomic issues with deep-bodied actinopterygians, this specimen represents the earliest known occurrence of one such fish. Its presence in the earliest Carboniferous indicates that actinopterygians were already postcranially differentiated in the aftermath of the Hangenberg. Moreover, this specimen suggests that earliest Carboniferous actinopterygians used multiple locomotory modes; recent data from later Carboniferous taxa suggest that actinopterygian locomotory modes proliferated throughout the Carboniferous. Taken together, these data suggest that early Carboniferous actinopterygians were morphologically, ecologically, and functionally diverse.

The lack of fossil tetrapod bearing deposits in the earliest Carboniferous ('Romer's Gap') has provoked some recent discussions regarding the proximal cause, with three explanations being offered: environmental, taphonomic, and collection failure. One of the few, and earliest, windows into this time is the locality of Blue Beach exposed in the Tournaisian deposits at Horton Bluff lying along the Avon River near Hantsport, Nova Scotia, Canada. This locality has long been known but, because the fossils were deposited in high energy settings they are almost always disarticulated, so the fauna has not been described in detail. Recent intensive collection has revealed a diverse assemblage of material, including for the first time associated elements, which permits an evaluation of the faunal constituents at the locality. Although not diagnosable to a fine taxonomic level, sufficient apomorphies are present to identify representatives from numerous clades known from more complete specimens elsewhere. The evidence suggests a diverse fauna was present, including whatcheeriids and embolomeres. A single humerus previously had been attributed to a colosteid, but there is some uncertainty with this identification. Additional elements suggest the presence of taxa otherwise only known from the late Devonian. Depositional biases at the locality favor tetrapod fossils from larger individuals, but indirect evidence from trackways and tantalizing isolated bones evidences the presence of small taxa that remain to be discovered. The fossils from Blue Beach demonstrate that when windows into the fauna of 'Romer's Gap' are found a rich diversity of tetrapods will be shown to be present, contra arguments that suggested this hiatus in the fossil record was due to extrinsic factors such as atmospheric oxygen levels. They also show that the early tetrapod fauna is not easily divisible into Devonian and Carboniferous faunas, suggesting that some tetrapods passed through the end Devonian extinction event unaffected.

The Early Permian recumbirostran lepospondyl Rhynchonkos stovalli has been identified as a possible close relative of caecilians due to general similarities in skull shape as well as similar robustness of the braincase, a hypothesis that implies the polyphyly of extant lissamphibians. In order to better assess this phylogenetic hypothesis, we studied the morphology of the holotype and three specimens previously attributed to R. stovalli. With the use of micro-computed x-ray tomography (μCT) we are able to completely describe the external and internal cranial morphology of these specimens, dramatically revising our knowledge of R. stovalli and recognizing two new taxa, Aletrimyti gaskillae gen et sp. n. and Dvellacanus carrolli gen et sp. n. The braincases of R. stovalli, A. gaskillae, and D. carrolli are described in detail, demonstrating detailed braincase morphology and new information on the recumbirostran supraoccipital bone. All three taxa show fossorial adaptations in the braincase, sutural articulations of skull roof bones, and in the lower jaw, but variation in cranial morphology between these three taxa may reflect different modes of head-first burrowing behaviors and capabilities. We revisit the homology of the supraoccipital, median anterior bone, and temporal bone of recumbirostrans, and discuss implications of alternate interpretations of the homology of these elements. Finally, we evaluate the characteristics previously used to unite Rhynchonkos stovalli with caecilians in light of these new data. These proposed similarities are more ambiguous than previous descriptions suggest, and result from the composite nature of previous descriptions, ambiguities in external morphology, and functional convergence between recumbirostrans and caecilians for head-first burrowing.

Recumbirostran 'microsaurs,' a group of early tetrapods from the Late Carboniferous and Early Permian, are the earliest known example of adaptation to head-first burrowing in the tetrapod fossil record. However, understanding of the diversity of fossorial adaptation within the Recumbirostra has been hindered by poor anatomical knowledge of the more divergent forms within the group. Here we report the results of μCT study of Quasicaecilia texana, a poorly-known recumbirostran with a unique, broad, shovel-like snout. The organization of the skull roof and braincase of Quasicaecilia is found to be more in line with that of other recumbirostrans than previously described, despite differences in overall shape. The braincase is found to be broadly comparable to Carrolla craddocki, with a large presphenoid that encompasses much of the interorbital septum and the columella ethmoidalis, and a single compound ossification encompassing the sphenoid, otic, and occipital regions. The recumbirostran braincase conserves general structure and topology of braincase regions and cranial nerve foramina, but it is highly variable in the number of ossifications and their extent, likely associated with the reliance on braincase ossifications to resist compression during sediment compaction and mechanical manipulation by epaxial and hypaxial musculature. Expansion of the deep ventral neck musculature in Quasicaecilia, autapomorphic among recumbirostrans, may reflect unique biomechanical function, and underscores the importance of future attention to the role of the cervical musculature in contextualizing the origin and evolution of fossoriality in recumbirostrans.

The scant fossil record of caecilians has obscured the origin and evolution of this lissamphibian group. Eocaecilia micropodia from the Lower Jurassic of North America remains the only stem-group caecilian with an almost complete skull preserved. However, this taxon has been controversial, engendering re-evaluation of traits considered to be plesiomorphic for extant caecilians. Both the validity of the placement of E. micropodia as a stem caecilian and estimates of the plesiomorphic condition of extant caecilians have been questioned. In order to address these issues, the braincase of E. micropodia was examined via micro-computed tomography. The braincase is considered to be a more reliable phylogenetic indicator than peripheral regions of the skull. These data reveal significant new information, including the possession of an ossified nasal septum, ossified anterior wall of the sphenethmoid, long anterolateral processes on the sphenethmoid, and paired olfactory nerve foramina, which are known only to occur in extant caecilians; the latter are possibly related to the evolution of the tentacle, a caecilian autapomorphy. A phylogenetic analysis that included 64 non-amniote taxa and 308 characters represents the first extensive test of the phylogenetic affinities of E. micropodia. The results place E. micropodia securely on the stem of extant caecilians, representing a clade within Temnospondyli that is the sister taxon to batrachians plus Gerobatrachus. Ancestral character state reconstruction confirms the braincase of E. micropodia to be largely representative of the plesiomorphic condition of extant caecilians. Additionally, the results refine the context within which the evolution of the caecilian form can be evaluated. The robust construction and pattern of the dermal skull of E. micropodia is interpreted as symplesiomorphic with advanced dissorophoid temnospondyls, rather than being autapomorphic in its robust construction. Together these data increase confidence in incorporating E. micropodia into discussions of caecilian evolution.

An amphibian evolution: Bridging the gap The origin of the amphibians (frogs, salamanders and caecilians) is one of the most controversial questions in vertebrate evolution, because of the large morphological and temporal gaps between today's amphibians and the extinct fossil forms. The discovery of an unusually complete Palaeozoic amphibian from the Early Permian of Texas has now helped to fill that gap. The new fossil has the overall appearance of a temnospondyl — an archaic amphibian — but boasts many characteristic features seen in modern frogs, toads and salamanders. A phylogenetic analysis splits the modern amphibia into two groups, separating at some time before 330 million years ago, with frogs, toads and salamanders related to temnospondyls, and caecilians more closely related to the lepospondyls, another group of archaic amphibians. Anderson and colleagues describe a fossil amphibian from the Early Permian of Texas that has the chassis of a temnospondyl but with the addition of many features seen in modern frogs, toads and salamanders. A phylogenetic analysis splits modern Amphibia into two groups, with frogs, toads and salamanders related to temnospondyls, and caecilians to leopspndyls. The origin of extant amphibians (Lissamphibia: frogs, salamanders and caecilians) is one of the most controversial questions in vertebrate evolution, owing to large morphological and temporal gaps in the fossil record 1 , 2 , 3 . Current discussions focus on three competing hypotheses: a monophyletic origin within either Temnospondyli 4 , 5 , 6 , 7 or Lepospondyli 8 , 9 , 10 , or a polyphyletic origin with frogs and salamanders arising among temnospondyls and caecilians among the lepospondyls 11 , 12 , 13 , 14 , 15 , 16 . Recent molecular analyses are also controversial, with estimations for the batrachian (frog–salamander) divergence significantly older than the palaeontological evidence supports 17 , 18 . Here we report the discovery of an amphibamid temnospondyl from the Early Permian of Texas that bridges the gap between other Palaeozoic amphibians and the earliest known salientians 19 , 20 and caudatans 21 from the Mesozoic. The presence of a mosaic of salientian and caudatan characters in this small fossil makes it a key taxon close to the batrachian (frog and salamander) divergence. Phylogenetic analysis suggests that the batrachian divergence occurred in the Middle Permian, rather than the late Carboniferous as recently estimated using molecular clocks 18 , 22 , but the divergence with caecilians corresponds to the deep split between temnospondyls and lepospondyls, which is congruent with the molecular estimates.

Elasmosaurid plesiosaurian remains have been documented from non-marine to paralic (fluvial to estuarine) sediments of the upper Campanian Dinosaur Park Formation (DPF) of southern Alberta since 1898. Despite this long collection history, this material has received relatively little research attention, largely due to the highly fragmentary nature of most recovered specimens. However, this assemblage is significant, as it constitutes a rare occurrence of plesiosaurian remains in a non-marine depositional environment. This study reports on a recently collected and prepared specimen, which represents the most complete elasmosaurid yet collected from the DPF. This specimen preserves the trunk region, the base of the neck and tail, a partial fore and hind limb, and tooth, and is sufficiently complete to be assigned as the holotype of a new genus and species. This new taxon is diagnosed by a distinctive character state combination including a boomerang-shaped clavicular arch with acute anterior process, convex anterolateral margin, deeply embayed posterior margin, and pronounced ventral keel, together with the presence of 22 dorsal vertebrae, and the anterior dorsal centra bearing a ventral notch. The DPF plesiosaurian fossils were recovered from both estuarine/bay and fluvial palaeochannel sediments. The holotype skeleton represents an osteologically mature individual with an estimated body length of around 5 m, although the largest referred DPF elasmosaurid might have been closer to 7 m, which is considerably larger than other plesiosaurians reported from non-marine deposits. This suggests small-body lengths relative to typical elasmosaurids from marine settings, but is consistent with other plesiosaurians recovered from non-marine sediments. The identification of a distinct elasmosaurid taxon in the DPF might be evidence of niche-partitioning among the predominantly oceanic members of the ubiquitous plesiosaurian clade.

Lysorophians are a group of early tetrapods with extremely elongate trunks, reduced limbs, and highly reduced skulls. Since the first discovery of this group, general similarities in outward appearance between lysorophians and some modern lissamphibian orders (specifically Urodela and Gymnophiona) have been recognized, and sometimes been the basis for hypotheses of lissamphibian origins. We studied the morphology of the skull, with particular emphasis on the neurocranium, of a partial growth series of the lysorophian Brachydectes newberryi using x-ray micro-computed tomography (μCT). Our study reveals similarities between the braincase of Brachydectes and brachystelechid recumbirostrans, corroborating prior work suggesting a close relationship between these taxa. We also describe the morphology of the epipterygoid, stapes, and quadrate in this taxon for the first time. Contra the proposals of some workers, we find no evidence of expected lissamphibian synapomorphies in the skull morphology in Brachydectes newberryi, and instead recognize a number of derived amniote characteristics within the braincase and suspensorium. Morphology previously considered indicative of taxonomic diversity within Lysorophia may reflect ontogenetic rather than taxonomic variation. The highly divergent morphology of lysorophians represents a refinement of morphological and functional trends within recumbirostrans, and is analogous to morphology observed in many modern fossorial reptiles.