Principal Features of the Mammalian Nasal Cavity were Present in Triassic Eucynodonts

Ruf, I., Maier, W., Rodrigues, P. G., and C. L. Schultz. 2014. Nasal Anatomy of the Non-mammaliaform Cynodont Brasilitherium riograndensis (Eucynodontia, Therapsida) Reveals New Insight into Mammalian Evolution. The Anatomical Record 297: 2018–2030. DOI: 10.1002/ar.23022
Abstract - The mammalian nasal cavity is characterized by a unique anatomy with complex internal features. The evolution of turbinals was correlated with endothermic and macrosmatic adaptations in therapsids and in early mammals, which is still apparent in their twofold function (warming and moistening of air, olfaction). Fossil evidence for the transformation from the nonmammalian to the mammalian nasal cavity pattern has been poor and inadequate. Ossification of the cartilaginous nasal capsule and turbinals seems to be a feature that occurred only very late in synapsid evolution but delicate ethmoidal bones are rarely preserved. Here we provide the first µCT investigation of the nasal cavity of the advanced non-mammaliaform cynodont Brasilitherium riograndensis from the Late Triassic of Southern Brazil, a member of the sister-group of mammaliaforms, in order to elucidate a critical anatomical transition in early mammalian evolution. Brasilitherium riograndensis already had at least partially ossified turbinals as remnants of the nasoturbinal and the first ethmoturbinal are preserved. The posterior nasal septum is partly ossified and contributes to a mesethmoid. The nasal cavity is posteriorly expanded and forms a distinctive pars posterior (ethmoidal recess) that is ventrally separated from the nasopharyngeal duct by a distinct lamina terminalis. Thus, our observations clearly demonstrate that principal features of the mammalian nasal cavity were already present in the sister-group of mammaliaforms.

Morphology and Diversity of the Mandibular Symphysis of Archosauriforms

Excellent study that demonstrates changes in mandibular symphyses through individual clades;  changes that appear to be influenced by changes in dental morphology, which presumably reflect differences in diet. This paper is part of the upcoming volume on early archosaurs.

 

Holliday, C. M., and S. J. Nesbitt. 2013. Morphology and diversity of the mandibular symphysis of archosauriforms In Nesbitt, S. J., Desojo, J. B. and R. B. Irmis (eds) Anatomy, Phylogeny and Palaeobiology of Early Archosaurs and their Kin. Geological Society, London, Special Publications, 379, http://dx.doi.org/10.1144/SP379.2

 

Abstract: Archosauromorphs radiated into numerous trophic niches during the Mesozoic, many of which were accommodated by particular suites of cranial adaptations and feeding behaviours. The mandibular symphysis, the joint linking the mandibles, is a poorly understood craniomandibular joint that may offer significant insight into skull function and feeding ecology. Using comparative data from extant amniotes, we investigated the skeletal anatomy and osteological correlates of relevant soft tissues in a survey of archosauromorph mandibular symphyses. Characters were identified and their evolution mapped using a current phylogeny of archosauriforms with the addition of non-archosauriform archosauromorphs. Extinct taxa with the simple Class I condition (e.g. proterochampsids, ‘rauisuchians’), rugose Class II (aetosaurs, protosuchians, silesaurids) and interdigitating Class III symphyses (e.g. phytosaurs, crocodyliforms) and finally fused Class IV (avians) build the joints in expected ways, although they differ in the contributions of bony elements and Meckel’s cartilage. Optimization of the different classes of symphyses across archosauromorph clades indicates that major iterative transitions from plesiomorphic Class I to derived, rigid Class II–IV symphyses occurred along the lines to phytosaurs, aetosaurs, a subset of poposauroids, crocodyliformes, pterosaurs and birds. These transitions in symphyseal morphology also appear to track changes in dentition and potentially diet.

Two New Temnospondyl Papers - Phylogeny of Major Clades and Suction Feeding in Gerrothorax

Schoch, R. R. 2013. The evolution of major temnospondyl clades: an inclusive phylogenetic analysis. Journal of Systematic Palaeontology DOI:10.1080/14772019.2012.699006http://www.tandfonline.com/doi/full/10.1080/14772019.2012.699006

Abstract - Phylogenetic analysis of a large dataset (72 taxa, 212 characters) focuses on the in-group relationships of temnospondyls, the largest lower tetrapod clade. Representatives of all clades and grades were considered, spanning the entire stratigraphical range of temnospondyls from the Early Carboniferous through to the Early Cretaceous. Several major groups are defined phylogenetically (node or branch-based) rather than by apomorphies. The following groups were unequivocally found to be monophyletic: Edopoidea (node), Dvinosauria (stem, excl. Brachyopidae), Dissorophoidea (node), Eryopidae (stem), and Stereospondyli (node). The latter encompass three well-defined, branch-based taxa: Rhinesuchidae, Trematosauria and Capitosauria. Trematosauria (stem) contain Trematosauroidea (node), which includes the classic trematosaurids, metoposaurids, and possibly part of the rhytidosteids (Peltostega) but their in-group relationships remain unsettled; most other short-snouted stereospondyls (chigutisaurids, brachyopids, Laidleria and the plagiosaurids) are probably monophyletic and likely nest in some form with trematosauroids. Capitosauria (stem) include the Capitosauroidea (node) spanned by Parotosuchus and Mastodonsaurus, with the successive stem taxa Edingerella, Benthosuchus, Wetlugasaurus and Watsonisuchus. In all variant analyses, edopoids form the basalmost temnospondyl clade, followed by a potential clade (or grade) of small terrestrial taxa containing Balanerpeton and Dendrerpeton (‘Dendrerpetontidae’). All taxa higher than Edopoidea are suggested to form the monophyletic stem taxon Eutemnospondyli, tax. nov. The remainder of Temnospondyli fall into four robust and undisputed clades: (1) Dvinosauria; (2) Zatracheidae plus Dissorophoidea; (3) Eryopidae; and (4) Stereospondyli. These taxa are together referred to as Rhachitomi (node). Eryopidae and Stereospondylomorpha are probably monophyletic, here referred to as Eryopiformes (tax. nov.). The position of Dissorophoidea + Zatracheidae is still ambiguous; it may either form the sister taxon of Dvinosauria, or nest between Dvinosauria and Eryopiformes, whereas there is no support for Euskelia (Dissorophoidea + Eryopidae) after basal taxa of each clade are better understood.



Witzmann, F. and R. R. Schoch. 2012. Reconstruction of cranial and hyobranchial muscles in the Triassic temnospondyl Gerrothorax provides evidence for akinetic suction feeding. Journal of Morphology DOI: 10.1002/jmor.20113 http://onlinelibrary.wiley.com/doi/10.1002/jmor.20113/abstract

Abstract - The cranial and hyobranchial muscles of the Triassic temnospondyl Gerrothorax have been reconstructed based on direct evidence (spatial limitations, ossified muscle insertion sites on skull, mandible, and hyobranchium) and on phylogenetic reasoning (with extant basal actinopterygians and caudates as bracketing taxa). The skeletal and soft-anatomical data allow the reconstruction of the feeding strike of this bottom-dwelling, aquatic temnospondyl. The orientation of the muscle scars on the postglenoid area of the mandible indicates that the depressor mandibulae was indeed used for lowering the mandible and not to raise the skull as supposed previously and implies that the skull including the mandible must have been lifted off the ground during prey capture. It can thus be assumed that Gerrothorax raised the head toward the prey with the jaws still closed. Analogous to the bracketing taxa, subsequent mouth opening was caused by action of the strong epaxial muscles (further elevation of the head) and the depressor mandibulae and rectus cervicis (lowering of the mandible). During mouth opening, the action of the rectus cervicis muscle also rotated the hyobranchial apparatus ventrally and caudally, thus expanding the buccal cavity and causing the inflow of water with the prey through the mouth opening. The strongly developed depressor mandibulae and rectus cervicis, and the well ossified, large quadrate-articular joint suggest that this action occurred rapidly and that powerful suction was generated. Also, the jaw adductors were well developed and enabled a rapid mouth closure. In contrast to extant caudate larvae and most extant actinopterygians (teleosts), no cranial kinesis was possible in the Gerrothorax skull, and therefore suction feeding was not as elaborate as in these extant forms. This reconstruction may guide future studies of feeding in extinct aquatic tetrapods with ossified hyobranchial apparatus.

Associated Evolution Of Bipedality And Cursoriality Among Triassic Archosaurs

Kubo, T., and M. O. Kubo. 2012. Associated evolution of bipedality and cursoriality among Triassic archosaurs: a phylogenetically controlled evaluation. Paleobiology 38: 474–485. http://www.bioone.org/doi/full/10.1666/11015.1
Abstract - Bipedalism evolved more than twice among archosaurs, and it is a characteristic of basal dinosaurs and a prerequisite for avian flight. Nevertheless, the reasons for the evolution of bipedalism among archosaurs have barely been investigated. Comparative analysis using phylogenetically independent contrasts showed a significant correlation between bipedality (relative length of forelimb) and cursoriality (relative length of metatarsal III) among Triassic archosaurs. This result indicates that, among Triassic archosaurs, bipeds could run faster than quadrupeds. Bipedalism is probably an adaptation for cursoriality among archosaurs, which may explain why bipedalism evolved convergently in the crocodilian and bird lineages. This result also indicates that the means of acquiring cursoriality may differ between archosaurs and mammals.

Skull Mechanics of Capitosaurs (Amphibia: Temnospondyli)

Fortuny, J., Marcé-Nogué, J., Gil, L. and Galobart, À. 2012. Skull Mechanics and the Evolutionary Patterns of the Otic Notch Closure in Capitosaurs (Amphibia: Temnospondyli). The Anatomical Record (advance online publication) doi: 10.1002/ar.22486 http://onlinelibrary.wiley.com/doi/10.1002/ar.22486/abstract

Abstract - Capitosaurs were among the largest amphibians that have ever lived. Their members displayed an amphibious lifestyle. We provide new information on functional morphology data, using finite element analysis (FEA) which has palaeoecological implications for the group. Our analyses included 17 taxa using (2D) plate models to test four loading cases (bilateral, unilateral and lateral bitings and skull raising system simulation). Our results demonstrates that, when feeding, capitosaurs concentrated the stress at the circumorbital region of the capitosaur skull and cranial sutures probably played a key role in dissipating and absorbing the stress generated during biting. Basal members (as Wetlugasaurus) were probably less specialized forms, while during Middle- and Late Triassic the group radiated into different ecomorphotypes with closed otic notch forms (as Cyclotosaurus) resulting in the strongest skulls during biting. Previous interpretations discussed a trend from an open to closed otic notch associated with lateral repositioning of the tabular horns, but the analysis of the skull-raising system reveals that taxa exhibiting posteriorly directed tabular horns display similar results during skull raising to those of closed otic notch taxa. Our results suggest that various constraints besides otic notch morphology, such as the elongation of the tabular horns, snout length, skull width and position, and size of the orbits affect the function of the skull. On the light of our results, capitosaur skull showed a trend to reduce the stresses and deformation during biting. Capitosaurs could be considered crocodilian analogues as they were top-level predators in fluvial and brackish Triassic ecosystems.

Postcranial Skeletal Pneumaticity in Triassic Archosaurs

Butler, R. J., Barrett, P. M., and D. J. Gower. 2012. Reassessment of the evidence for postcranial skeletal pneumaticity in Triassic archosaurs, and the early evolution of the avian respiratory system. PLoS ONE 7(3): e34094. doi:10.1371/journal.pone.0034094


Abstract - Uniquely among extant vertebrates, birds possess complex respiratory systems characterised by the combination of small, rigid lungs, extensive pulmonary air sacs that possess diverticula that invade (pneumatise) the postcranial skeleton, unidirectional ventilation of the lungs, and efficient crosscurrent gas exchange. Crocodilians, the only other living archosaurs, also possess unidirectional lung ventilation, but lack true air sacs and postcranial skeletal pneumaticity (PSP). PSP can be used to infer the presence of avian-like pulmonary air sacs in several extinct archosaur clades (non-avian theropod dinosaurs, sauropod dinosaurs and pterosaurs). However, the evolution of respiratory systems in other archosaurs, especially in the lineage leading to crocodilians, is poorly documented. Here, we use mCT-scanning to investigate the vertebral anatomy of Triassic archosaur taxa, from both the avian and crocodilian lineages as well as non-archosaurian diapsid outgroups. Our results confirm previous suggestions that unambiguous evidence of PSP (presence of internal pneumatic cavities linked to the exterior by foramina) is found only in bird-line (ornithodiran) archosaurs. We propose that pulmonary air sacs were present in the common ancestor of Ornithodira and may have been subsequently lost or reduced in some members of the clade (notably in ornithischian dinosaurs). The development of these avian-like respiratory features might have been linked to inferred increases in activity levels among ornithodirans. By contrast, no crocodile-line archosaur (pseudosuchian) exhibits evidence for unambiguous PSP, but many of these taxa possess the complex array of vertebral laminae and fossae that always accompany the presence of air sacs in ornithodirans. These laminae and fossae are likely homologous with those in ornithodirans, which suggests the need for further investigation of the hypothesis that a reduced, or non-invasive, system of pulmonary air sacs may be have been present in these taxa (and secondarily lost in extant crocodilians) and was potentially primitive for Archosauria as a whole.

Further Evidence of Burrowing in Procolophonid Parareptiles

MacDougall, M. J., Modesto, S. P., and J. Botha-Brink. 2012. The postcranial skeleton of the Early Triassic parareptile Sauropareion anoplus, with a discussion of possible life history. Acta Palaeontologica Polonica in press. Available online 27 Feb 2012 doi: http://dx.doi.org/10.4202/app.2011.0099.

Abstract - The skeletal anatomy of the Early Triassic (Induan) procolophonid reptile Sauropareion anoplus is described on the basis of three partial skeletons from Vangfontein, Middelburg District, South Africa. Together these three specimens preserve the large majority of the pectoral and pelvic girdles, articulated forelimbs and hindlimbs, and all but the caudal portion of the vertebral column, elements hitherto undescribed. Our phylogenetic analysis of the Procolophonoidea is consonant with previous work, positing S. anoplus as the sister taxon to a clade composed of all other procolophonids exclusive of Coletta seca. Previous studies have suggested that procolophonids were burrowers, and this seems to have been the case for S. anoplus, based on comparisons with characteristic skeletal anatomy of living digging animals, such as the presence of a spade-shaped skull, robust phalanges, and large unguals.

Disparity and Convergence in Bipedal Archosaur Locomotion

Bates, K. T., and E. R. Schachner. 2011. Disparity and convergence in bipedal archosaur locomotion. Journal of the Royal Society Interface (advance online publication) doi: 10.1098/​rsif.2011.0687

Abstract - This study aims to investigate functional disparity in the locomotor apparatus of bipedal archosaurs. We use reconstructions of hindlimb myology of extant and extinct archosaurs to generate musculoskeletal biomechanical models to test hypothesized convergence between bipedal crocodile-line archosaurs and dinosaurs. Quantitative comparison of muscle leverage supports the inference that bipedal crocodile-line archosaurs and non-avian theropods had highly convergent hindlimb myology, suggesting similar muscular mechanics and neuromuscular control of locomotion. While these groups independently evolved similar musculoskeletal solutions to the challenges of parasagittally erect bipedalism, differences also clearly exist, particularly the distinct hip and crurotarsal ankle morphology characteristic of many pseudosuchian archosaurs. Furthermore, comparative analyses of muscle design in extant archosaurs reveal that muscular parameters such as size and architecture are more highly adapted or optimized for habitual locomotion than moment arms. The importance of these aspects of muscle design, which are not directly retrievable from fossils, warns against over-extrapolating the functional significance of anatomical convergences. Nevertheless, links identified between posture, muscle moments and neural control in archosaur locomotion suggest that functional interpretations of osteological changes in limb anatomy traditionally linked to postural evolution in Late Triassic archosaurs could be constrained through musculoskeletal modelling.

Evolution of the Dinosauriform Respiratory Apparatus: New Evidence from the Postcranial Axial Skeleton

Schachner, E. R., Farmer, C. G., McDonald, A. T., and P. Dodson. 2011. Evolution of the dinosauriform respiratory apparatus: new evidence from the postcranial axial skeleton. The Anatomical Record Early View. DOI: 10.1002/ar.21439

Abstract - Examination of the thoracic rib and vertebral anatomy of extant archosaurs indicates a relationship between the postcranial axial skeleton and pulmonary anatomy. Lung ventilation in extant crocodilians is primarily achieved with a hepatic piston pump and costal rotation. The tubercula and capitula of the ribs lie on the horizontal plane, forming a smooth thoracic ‘‘ceiling’’ facilitating movement of the viscera. Although the parietal pleura is anchored to the dorsal thoracic wall, the dorsal visceral pleura exhibits a greater freedom of movement. The air sac system and lungs of birds are associated with bicapitate ribs with a ventrally positioned capitular articulation, generating a rigid and furrowed rib cage that minimizes dorsoventral changes in volume in the dorsal thorax. The thin walled bronchi are kept from collapsing by fusion of the lung to the thorax on all sides. Data from this study suggest a progression from a dorsally rigid, heterogeneously partitioned, multichambered lung in basal dinosauriform archosaurs towards the small entirely rigid avian-style lung that was likely present in saurischian dinosaurs, consistent with a constant volume cavum pulmonale, thin walled parabronchi, and distinct air sacs. There is no vertebral evidence for a crocodilian hepatic piston pump in any of the taxa reviewed. The evidence for both a rigid lung and unidirectional airflow in dinosauriformes raises the possibility that these animals had a highly efficient lung relative to other Mesozoic vertebrates, which may have contributed to their successful radiation during this time period.

Pelvic and Hindlimb Myology of the Basal Archosaur Poposaurus gracilis

Schachner, E. R., Manning, P. L., and P. Dodson. 2011. Pelvic and hindlimb myology of the basal archosaur Poposaurus gracilis (Archosauria: Poposauroidea). Journal of Morphology. doi: 10.1002/jmor.10997. [Epub ahead of print]

Abstract - The discovery of a largely complete and well preserved specimen of Poposaurus gracilis has provided the opportunity to generate the first phylogenetically based reconstruction of pelvic and hindlimb musculature of an extinct nondinosaurian archosaur. As in dinosaurs, multiple lineages of basal archosaurs convergently evolved parasagittally erect limbs. However, in contrast to the laterally projecting acetabulum, or "buttress erect" hip morphology of ornithodirans, basal archosaurs evolved a very different, ventrally projecting acetabulum, or "pillar erect" hip. Reconstruction of the pelvic and hindlimb musculotendinous system in a bipedal suchian archosaur clarifies how the anatomical transformations associated with the evolution of bipedalism in basal archosaurs differed from that of bipedal dinosaurs and birds. This reconstruction is based on the direct examination of the osteology and myology of phylogenetically relevant extant taxa in conjunction with osteological correlates from the skeleton of P. gracilis. This data set includes a series of inferences (presence/absence of a structure, number of components, and origin/insertion sites) regarding 26 individual muscles or muscle groups, three pelvic ligaments, and two connective tissue structures in the pelvis, hindlimb, and pes of P. gracilis. These data provide a foundation for subsequent examination of variation in myological orientation and function based on pelvic and hindlimb morphology, across the basal archosaur lineage leading to extant crocodilians.

Biomechanical modeling of Coelophysis bauri

Jasinski, S. E. 2011. Biomechanical modeling of Coelophysis bauri: Possible feeding methods and behavior of a Late Triassic theropod. New Mexico Museum of Natural History and Science, Bulletin 53:195-201.

ABSTRACT—Coelophysis bauri is a small theropod dinosaur from the Late Triassic of the southwestern United States. The Whitaker (= Ghost Ranch)quarry, which preserves hundreds to perhaps thousands of individuals, has led to many hypotheses about its behavior and feeding strategies. The beam theory allows for a quantitative approach to reconstruction of the feeding mechanism of Coelophysis. The Coelophysis mandible behaves as a simple lever, similar to that of Varanus komodoensis, and thus is more inclined to produce slashing bites. The middentary region of the Coelophysis mandible has the greatest labiolingual strength, suggesting that this would have been the region most important to live prey capture. Comparisons of the juvenile and adult specimens of Coelophysis bauri show remarkable similarities in the relative mandibular strengths, suggesting juveniles would have been apt predators. Further study of the biomechanics of theropod dinosaurs such as Coelophysis bauri may lead to a better understanding of how they lived and behaved.

A New Nearly Complete Articulated Skeleton of the Bipedal Pseudosuchian Poposaurus gracilis from the Upper Triassic of Utah

Back in 2003 I was fortunate to be part of a research team that discovered a mostly complete articulated skeleton of Poposaurus gracilis from the Chinle Formation of Utah. Uncovering such an amazing specimen is definitely one of the highlights of my career, especially the moment when we realized how amazingly complete and articulated the specimen was. Below is a photo of the articulated hind limb and pelvis at that moment during the excavation.

I've seen the specimen a couple of times at Yale in the process of preparation and the specimen is definitely a beauty. Furthermore is is extremely important because prior to this find P. gracilis was only known from the pelvis, hind limbs, and some vertebrae.  Unfortunately, the skull of this new specimen was unrecoverable, but the rest provides much information including an articulated forelimb and the pes (foot). I should also add that we met a lot of good friends on this trip.

The first paper out on this specimen is not a detailed osteological study but rather introduces the specimen and provides a detailed determination of the proposed locomotor abilities of this taxon, concluding from numerous lines of evidence (phylogenetic, extant phylogenetic bracketing, morphological) that Poposaurus was an obligatory biped. Morphological support for bipedality in P. gracilis includes a fully erect posture, relatively short forelimbs, elongate pes, five sacral vertebrae, partially open acetabulum, and an elongate tail. Gauthier et al. also provide an in-depth argument that the development of bipedality in Archosauria represents a major evolutionary innovation in vertebrate history.

Finally this paper gives a preview of Sterling Nesbitt's upcoming (in press, out very very shortly) extremely detailed phylogenetic analysis of the Archosauria (from his PhD dissertation).  Sterling's analysis provides some pretty surprising (and well-supported) placements of various taxa that will turn some heads and cause a good bit of discussion, especially regarding the revised definitions of some beloved (by some) clade names.

Although I am told that the Poposaurus paper is officially out, it is not up yet on either the Yale Peabody Museum website or the BioOne site, but should be soon. I'll post the link as soon as it does. In the meantime you can read more about the initial find here. I'm also looking forward to the complete osteological description of this amazng specimen.

Gauthier, J. A., Nesbitt, S. J., Schachner, E. R., Bever, G. S., and W. G. Joyce. 2011. The bipedal stem crocodilian Poposaurus gracilis: inferring function in fossils and innovation in archosaur locomotion. Bulletin of the Peabody Museum of Natural History 52:107-126.

Abstract - We introduce a spectacular new specimen of a Late Triassic stem crocodilian identified as Poposaurus gracilis. It is part of a poorly known group, Poposauroidea, that, because of its striking similarities with contemporaneous stem avians (“dinosaurs”), has long puzzled archosaur paleontologists. Observed vertebrate locomotor behaviors, together with exceptional preservation of distinctive anatomical clues in this fossil, enable us to examine locomotor evolution in light of new advances in phylogenetic relationships among Triassic archosaurs. Because this stem crocodilian is unambiguously an archosaur, a diapsid, a tetrapod and a choanate sarcopterygian, we can safely infer major components of its locomotor behavior. These inferences, together with form-function constraints, suggest that P. gracilis was a fleet-footed, obligately erect-postured, striding biped. That behavior seems to have been superimposed on the ancestral archosaur’s innovative locomotor repertoire, which includes the capacity to “high walk.” These novelties persist in a recognizable form in archosaurs for at least 245 million years and are widely distributed across Earth’s surface in diverse ecological settings. They thus qualify as evolutionary innovations regardless of significant differences in diversification rates among extant diapsid reptiles.

New From Jeff Martz - Poposaurus in Color; and the Question of Bipedality in Animals with a Crocodile-normal Ankle Type

Jeff Martz has recently colo(u)rized his Poposaurus gracilis reconstruction and here it is in its new glory.

When I see reconstructions of pseudosuchians in a bipedal stance I always reflect upon the question of whether or not it is functionally possible for an animal with a crocodile-normal ankle type to actually be bipedal, especially an obligate biped.  I've perused the literature but have found very little to address this problem. I'm certain that Emma Schachner dealt with this in her recently completed PhD on the new equisitely preserved Poposaurus specimen from the Chinle Formation in Utah, but unfortunately I've not had a chance to borrow and read a copy.  Her 2009 SVP abstract suggests that myological developments differ between bipedal pseudosuchians and bipedal dinosaurs, suggesting that these myological changes compensate for the alternate ankle structures.

Crocodylian-like Trunk Bracing System in Armored Basal Tetrapods

Buchwitz, M., Witzmann, F., Voigt, S. and Golubev, V. 2011. Osteoderm microstructure indicates the presence of a crocodylian-like trunk bracing system in a group of armoured basal tetrapods. Acta Zoologica (Stockholm) 00:1–21. DOI: 10.1111/j.1463-6395.2011.00502.x

Abstract - The microstructure of dorsal osteoderms referred to the chroniosuchid taxa Chroniosuchus, Chroniosaurus, Madygenerpeton and cf. Uralerpeton is compared to existing data on the bystrowianid chroniosuchian Bystrowiella and further tetrapods. Chroniosuchid osteoderms are marked by thin internal and relatively thick external cortices that consist of lowly vascularised parallel-fibred bone. They are structured by growth marks and, in case of Madygenerpeton, by lines of arrested growth. The cancellous middle region is marked by a high degree of remodelling and a primary bone matrix of parallel-fibred bone that may include domains of interwoven structural fibres. Whereas the convergence of Bystrowiella and chroniosuchid osteoderms is not confirmed by our observations, the internal cortex of the latter displays a significant peculiarity: It contains distinct bundles of shallowly dipping Sharpey’s fibres with a cranio- or caudoventral orientation. We interpret this feature as indicative for the attachment of epaxial muscles which spanned several vertebral segments between the medioventral surface of the osteoderms and the transversal processes of the thoracic vertebrae. This finding endorses the hypothesis that the chroniosuchid osteoderm series was part of a crocodylian-like trunk bracing system that supported terrestrial locomotion. According to the measured range of osteoderm bone compactness, some chroniosuchian species may have had a more aquatic lifestyle than others.

Postcranial Skeletal Pneumaticity in Basal Sauropodomorphs

Yates, A. M., Wedel, M. J., and M. F. Bonnan. 2011. The early evolution of postcranial skeletal pneumaticity in sauropodomorph dinosaurs. Acta Palaeontologica Polonica in press available online 07 Mar 2011 doi:10.4202/app.2010.0075

Abstract - Postcranial skeletal pneumaticity (PSP) is present in a range of basal sauropodomorphs spanning the basal sauropodomorph–sauropod transition. We describe the PSP of five taxa, Plateosaurus engelhardti, Eucnemesaurus fortis, Aardonyx celestae, Antetonitrus ingenipes and an unnamed basal sauropod from Spion Kop, South Africa (hereafter referred to as the Spion Kop sauropod). The PSP of Plateosaurus is apparently sporadic in its occurrence and has only been observed in very few specimens, in which it is of very limited extent, affecting only the posterior cervical vertebrae and possibly the mid dorsals in one specimen. The PSP of Eucnemesaurus, Aardonyx, Antetonitrus and the Spion Kop sauropod consists of subfossae (fossa-within-fossa structures) that excavate the vertices of the posterior infradiapophyseal fossae of the posterior dorsal vertebrae. These subfossae range from simple shallow depressions (Eucnemesaurus) to deep, steep-sided, internally subdivided and asymmetrically developed chambers (Antetonitrus). The middle and anterior dorsal vertebrae of these taxa lack PSP, demonstrating that abdominal air sacs were the source of the invasive diverticula. The presence of pneumatic features within the infradiapophyseal fossae suggest that the homologous fossae of more basal saurischians and dinosauriforms were receptacles that housed pneumatic diverticula. We suggest that it is probable that rigid non-compliant lungs ventilated by compliant posterior air sacs evolved prior to the origination of Dinosauria.

Unique Carapace Structure in a Triassic Tetrapod

Wow, and I thought that aetosaur carapaces were complex. Osteoderms provide more than defense and display.  They also provide support and stabilization for the axial skeleton. Choniosuchians seem to take this to a unique level.

Buchwitz, M. and S. Voigt. 2010. Peculiar carapace structure of a Triassic chroniosuchian implies evolutionary shift in trunk flexibility. Journal of Vertebrate Paleontology 30:1697-1708. DOI: 10.1080/02724634.2010.521685
Abstract - Dermal ossifications are widespread in Permian and Triassic tetrapods, but only members of the Chroniosuchia possess a series of dorsal osteoderms with a complex plate-to-plate articulation mechanism in addition to a contact between each osteoderm and its associated vertebral spine. The stratigraphically youngest chroniosuchid, Madygenerpeton pustulatus, from the Triassic of Kyrgyzstan provides new insight on the function of the chroniosuchian osteoderm system. Osteoderms of M. pustulatus are broad, peaked-roof-shaped to arched, with enlarged posterodorsal and anteroventral articulation facets bearing unique sets of concentric rail-like ridges and furrows. Supplementing the multiple-overlap chroniosuchian type articulation, the interlocking ridges and furrows confined the relative motion of two neighboring osteoderms to a rotation in slightly oblique and curved contact planes. Given the significant lateral narrowing of the dorsal ornamented non-overlap area, the horizontal component of the plate-to-plate rotation angles could reach up to 7.5◦, enabling more extensive lateral flexion of the trunk than in other chroniosuchids. Considering functional analogs, the chroniosuchian osteoderm system probably stabilized the vertebral column against shearing, torsion, tension, and compression loads and thus facilitated terrestrial locomotion at the expense of trunk flexibility. With its particular morphology, the carapace of M. pustulatus, however, was more suitable for locomotion styles featuring lateral body undulation than the carapaces of Permian chroniosuchids. We interpret this speciality as a secondary adaptation to an aquatic habitat.

New Species of the Late Triassic Venomous Archosauriform Uatchitodon

Mitchell, J. S., Heckert, A. B., and H.-D. Sues. 2010. Grooves to tubes: evolution of the venom delivery system in a Late Triassic “reptile”. Naturwissenschaften, online first. DOI 10.1007/s00114-010-0729-0

Abstract - Venom delivery systems occur in a wide range of extant and fossil vertebrates and are primarily based on oral adaptations. Teeth range from unmodified (Komodo dragons) to highly specialized fangs similar to hypodermic needles (protero- and solenoglyphous snakes). Developmental biologists have documented evidence for an infolding pathway of fang evolution, where the groove folds over to create the more derived condition. However, the oldest known members of venomous clades retain the same condition as their extant relatives, resulting in no fossil evidence for the transition. Based on a comparison of previously known specimens with newly discovered teeth from North Carolina, we describe a new species of the Late Triassic archosauriform Uatchitodon and provide detailed analyses that provide evidence for both venom conduction and document a complete structural series from shallow grooves to fully enclosed tubular canals. While known only from teeth, Uatchitodon is highly diagnostic in possessing compound serrations and for having two venom canals on each tooth in the dentition. Further, although not a snake, Uatchitodon sheds light on the evolutionary trajectory of venom delivery systems in amniotes and provide solid evidence for venom conduction in archosaur-line diapsids.

Supplemental data available here.

The Role of the Calcaneal 'Heel' in Basal Archosaurs

Sullivan, C. 2010. The role of the calcaneal 'heel' as a propulsive lever in basal archosaurs and extant monitor lizards. Journal of Vertebrate Paleontology 30:1422-1432. 

Abstract - The structurally complex ankles of Triassic archosaurs (Reptilia: Diapsida) have been repeatedly described, and tarsal characters have played an important role in analyses of archosaur phylogeny. Morphological variations in the ankle joint undoubtedly had a major impact on locomotion, but the functional implications of many ankle features remain poorly understood. This paper investigates the function of one such structure, the prominent and highly distinctive lateral to posterolateral calcaneal process that occurs throughout basal archosaurs and is homologous to the heel-like calcaneal tuber of pseudosuchians. A morphologically analogous lateral process occurs in extant varanid lizards, and X-ray rotoscopic analysis of the limb movements of savannah monitors (Varanus exanthematicus) indicates that the lateral process rotates into a vertical orientation as the limb retracts. As the varanid lateral process approaches the vertical, it becomes increasingly effective in adding to the propulsive moment arm of m. peroneus longus. A computer model of the ankle was compared to modified versions in which the lateral process and the associated proximal expansion of the fifth metatarsal were eliminated, and this virtual experiment demonstrated that the combination of the two structures increases the peak propulsive moment arm of the varanid m. peroneus longus by over 26%. The lateral calcaneal process of basal archosaurs probably acted similarly, enhancing the ability of the peroneus musculature to contribute to propulsion during walking. Like the calcaneal tuber of pseudosuchian archosaurs, the primitive lateral process was a lever that contributed to locomotion by allowing ankle plantarflexion to generate a greater propulsive moment.

Postosuchus was Even Bigger Than we Thought

Holliday, C. M., Ridgely, R. C., Sedlmayr, J. C., and L. M. Witmer. 2010. Cartilaginous Epiphyses in Extant Archosaurs and Their Implications for Reconstructing Limb Function in Dinosaurs. PLoS ONE 5(9): e13120. doi:10.1371/journal.pone.0013120

Abstract - Extinct archosaurs, including many non-avian dinosaurs, exhibit relatively simply shaped condylar regions in their appendicular bones, suggesting potentially large amounts of unpreserved epiphyseal (articular) cartilage. This “lost anatomy” is often underappreciated such that the ends of bones are typically considered to be the joint surfaces, potentially having a major impact on functional interpretation. Extant alligators and birds were used to establish an objective basis for inferences about cartilaginous articular structures in such extinct archosaur clades as non-avian dinosaurs. Limb elements of alligators, ostriches, and other birds were dissected, disarticulated, and defleshed. Lengths and condylar shapes of elements with intact epiphyses were measured. Limbs were subsequently completely skeletonized and the measurements repeated. Removal of cartilaginous condylar regions resulted in statistically significant changes in element length and condylar breadth. Moreover, there was marked loss of those cartilaginous structures responsible for joint architecture and congruence. Compared to alligators, birds showed less dramatic, but still significant changes. Condylar morphologies of dinosaur limb bones suggest that most non-coelurosaurian clades possessed large cartilaginous epiphyses that relied on the maintenance of vascular channels that are otherwise eliminated early in ontogeny in smaller-bodied tetrapods. A sensitivity analysis using cartilage correction factors (CCFs) obtained from extant taxa indicates that whereas the presence of cartilaginous epiphyses only moderately increases estimates of dinosaur height and speed, it has important implications for our ability to infer joint morphology, posture, and the complicated functional movements in the limbs of many extinct archosaurs. Evidence suggests that the sizes of sauropod epiphyseal cartilages surpassed those of alligators, which account for at least 10% of hindlimb length. These data suggest that large cartilaginous epiphyses were widely distributed among non-avian archosaurs and must be considered when making inferences about locomotor functional morphology in fossil taxa.

The Digital Plateosaurus II

Mallison, H. 2010. The digital Plateosaurus II: An assessment of the range of motion of the limbs and vertebral column and of previous reconstructions using a digital skeletal mount. Acta Palaeontologica Polonica 55:433–458.

Abstract - Scientific literature and museum exhibits are full of explicit and implicit claims about the possible postures and motion ranges of dinosaurs. For the example of the prosauropod Plateosaurus engelhardti I assessed the motion range of limbs and vertebral column in a CAD program using a 3D virtual skeletal mount. The range of motion of the forelimb is very limited, allowing the grasping of objects placed directly ventrally and ventrolaterally of the anterior torso. The manus is adapted for grasping. The powerful fore limb can barely reach in front of the shoulder, making a quadrupedal walking cycle impractical. Only a digitigrade pose of the pes with a steeply held metatarsus is feasible, and the morphology of the stylopodium and zeugopodium indicates a slightly flexed limb posture. Hind limb protraction and retraction are limited
by the pelvic architecture. The neck has significant mobility both dorsoventrally and laterally, but blocks torsion. The dorsal vertebral column is flexible to a degree similar to the neck, mainly in the anterior half, but blocks torsion totally in the anterior and posterior thirds. The anterior dorsals are similar in shape to the posterior cervicals and significantly increase the motion range of the neck. The tail is highly flexible due to its large number of elements, showing more lateral than dorsoventral mobility. These results are compared to reconstruction drawings and museum skeletal mounts, highlighting a pattern of errors specific to certain widely used reconstruction methods.