Field of Science

Asperoris mnyama, a New Archisauriform from the Middle Triassic of Tanzania

Nesbitt, S. J., Butler, R. J., and D. J. Gower. 2013. A New Archosauriform (Reptilia: Diapsida) from the Manda Beds (Middle Triassic) of Southwestern Tanzania. PLoS ONE 8(9): e72753. doi:10.1371/journal.pone.0072753


Archosauria and their closest relatives, the non-archosaurian archosauriforms, diversified in the Early and Middle Triassic, soon after the end-Permian extinction. This diversification is poorly documented in most Lower and Middle Triassic rock sequences because fossils of early groups of archosauriforms are relatively rare compared to those of other amniotes. The early Middle Triassic (? late Anisian) Manda beds of southwestern Tanzania form an exception, with early archosaur skeletons being relatively common and preserved as articulated or associated specimens. The Manda archosaur assemblage is exceptionally diverse for the Middle Triassic. However, to date, no non-archosaurian archosauriforms have been reported from these rocks.
Methodology/Principal Findings
Here, we name a new taxon, Asperoris mnyama gen. et sp. nov., from the Manda beds and thoroughly describe the only known specimen. The specimen consists of a well-preserved partial skull including tooth-bearing elements (premaxilla, maxilla), the nasal, partial skull roof, and several incomplete elements. All skull elements are covered in an autapomorphic highly rugose sculpturing. A unique combination of character states indicates that A. mnyama lies just outside Archosauria as a stem archosaur within Archosauriformes, but more precise relationships of A. mnyama relative to other early archosauriform clades (e.g., Erythrosuchidae) cannot be determined currently.
Asperoris mnyama is the first confirmed non-archosaurian archosauriform from the Manda beds and increases the morphological and taxonomic diversity of early archosauriforms known from the Middle Triassic. The direct association of A. mnyama with species referable to Archosauria demonstrates that non-archosaurian archosauriforms were present during the rise and early diversification of Archosauria. Non-archosaurian archosauriforms and archosaurs co-occur in fossil reptile assemblages across Pangaea from the late Early Triassic to the end of the Late Triassic.

The Problem of Dinosaur Origins: Integrating Three Approaches to the Rise of Dinosauria

An excellent essay looking on depth at the question of dinosaur origins. I think the ideas put forth are worthy of dicussion and will be heavily debated, especially since it is written from a North American perspective.

Padian, K., 2013. The problem of dinosaur origins: integrating three approaches to the
rise of Dinosauria. Earth and Environmental Science Transactions of the Royal Society of
Edinburgh First View Article. DOI:

Abstract - The problem of the origin of dinosaurs has historically had three dimensions. The first is the question of whether Dinosauria is monophyletic, and of its relationships to other archosaurs. This
question was plagued from the beginning by a lack of relevant fossils, an historical burden of confusing taxonomic terms and a rudimentary approach to devising phylogenies. The second dimension concerns the functional and ecological adaptations that differentiated dinosaurs from other archosaurs, a question also marred by lack of phylogenetic clarity and testable biomechanical hypotheses. The third dimension comprises the stratigraphic timing of the origin of dinosaurian groups with respect to each other and to related groups, the question of its synchronicity among various geographic regions, and some of the associated paleoenvironmental circumstances. None of these dimensions alone answers the question of dinosaur origins, and they sometimes provide conflicting implications. Since Dinosauria was named, one or another set of questions has historically dominated academic discussion and research. Paradigms have shifted substantially in recent decades, and current evidence suggests that we are due for more such shifts. I suggest two changes in thinking about the beginning of the “Age of Dinosaurs”: first, the event that we call the (phylogenetic) origin of dinosaurs was trivial compared to the origin of Ornithodira; and second, the “Age of Dinosaurs” proper did not begin until the Jurassic. Re-framing our thinking on these issues will improve our understanding of clade dynamics, timing of macroevolutionary events, and the effects of Triassic climate change on terrestrial vertebrates.

Reintroducing the Phytosaur Genus Machaeoroprosopus

I had posted about the taxonomic problem caused by the loss of the holotype specimen Machaeroprosopus validus before with a plea for people to help look for the specimen.  Come to find out a careful reading of the literature and the ICZN demonstrates that the specimen is not as important as previously believed regarding the taxonomic status of the name Machaeroprosopus. It will take awhile for Triassic and phytosaur workers to get used to using the name again and I am sure that some will just flat out refuse, but according to the ICZN the name Machaeoroprosopus is the proper one if the holotype specimen of "Belodon" buceros is truly diagnosable and if the other nominal species of "Pseudopalatus" are referable to the same genus represented by "B." buceros.

Parker, W. G., Hungerbühler, A., and J. W. Martz. 2013. The taxonomic status of the phytosaurs (Archosauriformes) Machaeroprosopus and Pseudopalatus from the Late Triassic of the western United States. Earth and Environmental Science Transactions of the Royal Society of Edinburgh First View Article. DOI:

Abstract - The genus Machaeroprosopus has long been considered invalid because the type specimen of the Late Triassic phytosaur species, M. validus, has been lost. Re-examination of the primary literature regarding the establishment of the Late Triassic phytosaur genus Machaeroprosopus demonstrates that M. buceros is the correct type species, not M. validus. Thus, the genus level name Machaeroprosopus has priority over the genera Pseudopalatus and Arribasuchus and all nominal species should be reassigned. Reassignment of these species to Machaeroprosopus satisfies the requirements of the International Code of Zoological Nomenclature (ICZN) and preserves historical context. The name Pseudopalatinae is retained as the valid clade name for these phytosaurs because its usage falls outside of the ICZN.

New Triassic Papers in the Festschift in Honor of Dr. Wann Langston Jr.

Back in very early 2010 Dr. Ernie Lundelius was honored with a festschift volume.  When I congratulated him on it he lamented that his close friend and colleague Dr. Wann Langston Jr., still did not have a festschift in his honor.  Ernie said they had tried to get one going a few times but nothing had ever come of it. I agreed to help to start another one and after several trials am proud to announce that the first online papers are now available from the Earth and Environmental Transactions of the Royal Society of Edinburgh. The final print volume will be out a bit later.  Unfortunately Dr. Langston passed away earlier this year and did not get to see the completed volume; however he did get to see a compilation of the abstracts.

Dr. Langston was Charles L. Camp's graduate student at the University of California at Berkeley and althogh his dissertation was on Permian vertebrates of North America, one of his firs publications was a description of a new phytosaur from the Upper Triassic of Texas.  Thus it is fitting that there are a few Triassic papers in his festschrift volume.  Two, which I was involved with, are up currently: I'll post the rest when they come online.

Parker, W. G. 2013. Redescription and taxonomic status of specimens of Episcoposaurus and Typothorax, the earliest known aetosaurs (Archosauria: Suchia) from the Upper Triassic of western North America, and the problem of proxy “holotypes”. Earth and Environmental Transactions of the Royal Society of Edinburgh First View Article. DOI:

Abstract - Historic type and referred material of the aetosaurian taxa Typothorax coccinarum, Episcoposaurus horridus and Episcoposaurus haplocerus are redescribed and the non-aetosaurian material identified and removed, a task previously considered “hopeless”. Reexamination of the original material reveals that the holotypes of E. haplocerus and probably T. coccinarum are not diagnosable at the species level and therefore are nomena dubia. The next available names for material referred to these taxa are Desmatosuchus spurensis and E. horridus respectively, although it may be more desirable for reasons of taxonomic stability to attempt to petition for a neotype in the latter case. The redescription of historical specimens is necessary to determine their nomenclatural validity. The use of referred specimens as proxy “type” specimens is problematic, as these referrals were originally made not on the basis of apomorphies, but rather on biostratigraphic and/or geographical assumptions which are inherently circular and cannot be unambiguously supported.

Martz, J. W., Mueller, B., Nesbitt, S. J., Stocker, M. R., Parker, W. G., Atanassov, M., Fraser, N., Weinbaum, J., and J. R. Lehane. 2013. A taxonomic and biostratigraphic re-evaluation of the Post Quarry vertebrate assemblage from the Cooper Canyon Formation (Dockum Group, Upper Triassic) of southern Garza County, western Texas. Earth and Environmental Transactions of the Royal Society of Edinburgh First View Article. DOI:

Abstract - The Post Quarry, within the lower part of the type section of the Upper Triassic Cooper Canyon Formation in southern Garza County, western Texas, contains a remarkably diverse vertebrate assemblage. The Post Quarry has produced: the small temnospondyl Rileymillerus cosgriffi; the metoposaurid Apachesaurus gregorii; possible dicynodonts and eucynodonts; a clevosaurid sphenodontian; non-archosauriform archosauromorphs (Trilophosaurus dornorum, simiosaurians, and possibly Malerisaurus); the phytosaur Leptosuchus; several aetosaurs (Calyptosuchus wellesi, Typothorax coccinarum, Paratypothorax, and Desmatosuchus smalli); the poposauroid Shuvosaurus inexpectatus (“Chatterjeea elegans”); the rauisuchid Postosuchus kirkpatricki; an early crocodylomorph; several dinosauromorphs (the lagerpetid Dromomeron gregorii, the silesaurid Technosaurus smalli, a herrerasaurid, and an early neotheropod); and several enigmatic small diapsids. Revised lithostratigraphic correlations of the lower Cooper Canyon Formation with the Tecovas Formation, the occurrence of Leptosuchus, and the overall composition of the assemblage indicate that the Post Quarry falls within the Adamanian biozone, and not the Revueltian biozone. Stratigraphic subdivision of the Adamanian biozone may be possible, and the Post Quarry may be correlative with the upper part of the Adamanian biozone in Arizona. The age of the Post Quarry assemblage is possibly late Lacian or earliest Alaunian (late early Norian or earliest middle Norian), between 220 and 215 Ma.

Paleoecology of the Early Triassic Dinwoody Formation: Insights on Recovery After the End-Permain Extinction

Hofmann, R., Hautmann, M. and H. Bucher. 2013. A New Paleoecological Look at the Dinwoody Formation (Lower Triassic, Western USA): Intrinsic Versus Extrinsic Controls on Ecosystem Recovery After the End-Permian Mass Extinction Journal of Paleontology 87:854-880. 2013  doi:

Abstract - The Dinwoody Formation of the western United States represents an important archive of Early Triassic ecosystems in the immediate aftermath of the end-Permian mass extinction. We present a systematic description and a quantitative paleoecological analysis of its benthic faunas in order to reconstruct benthic associations and to explore the temporal and spatial variations of diversity, ecological structure and taxonomic composition throughout the earliest Triassic of the western United States. A total of 15 bivalve species, two gastropod species, and two brachiopod species are recognized in the study area. The paleoecological analysis shows that the oldest Dinwoody communities are characterized by low diversity, low ecological complexity and high dominance of few species. We suggest that this low diversity most likely reflects the consequences of the mass extinction in the first place and not necessarily the persistence of environmental stress. Whereas this diversity pattern persists into younger strata of the Dinwoody Formation in outer shelf environments, an increase in richness, evenness and guild diversity occurred around the Griesbachian–Dienerian boundary in more shallow marine habitats. This incipient recovery towards the end of the Griesbachian is in accordance with observations from other regions and thus probably represents an interregional signal. In contrast to increasing richness within communities (alpha-diversity), beta-diversity remained low during the Griesbachian and Dienerian in the study area. This low beta-diversity reflects a wide environmental and geographical range of taxa during the earliest Triassic, indicating that the increase of within-habitat diversity has not yet led to significant competitive exclusion. We hypothesize that the well-known prevalence of generalized taxa in post-extinction faunas is primarily an effect of reduced competition that allows species to exist through the full range of their fundamental niches, rather than being caused by unusual and uniform environmental stress.

Morphological and Biomechanical Disparity of Crocodile-line Archosaurs Following the End-Triassic Extinction

This article is open access. I'm assuming that in using 'crurotarsan' they are using Paul Sereno's original definition and not the clade including Avemetatarsalia as recovered by Nesbitt (2011). 

Stubb, T. L., Pierce, S. E., Rayfield, E. J., and P. S. L. Anderson. 2013. Morphological and biomechanical disparity of crocodile-line archosaurs following the end-Triassic extinction. Proceedings of the Royal Society B 280 no. 1770 20131940 doi: 10.1098/rspb.2013.1940
Abstract - Mesozoic crurotarsans exhibited diverse morphologies and feeding modes, representing considerable ecological diversity, yet macroevolutionary patterns remain unexplored. Here, we use a unique combination of morphological and biomechanical disparity metrics to quantify the ecological diversity and trophic radiations of Mesozoic crurotarsans, using the mandible as a morpho-functional proxy. We recover three major trends. First, the diverse assemblage of Late Triassic crurotarsans was morphologically and biomechanically disparate, implying high levels of ecological variation; but, following the end-Triassic extinction, disparity declined. Second, the Jurassic radiation of marine thalattosuchians resulted in very low morphological disparity but moderate variation in jaw biomechanics, highlighting a hydrodynamic constraint on mandibular form. Third, during the Cretaceous terrestrial radiations of neosuchians and notosuchians, mandibular morphological variation increased considerably. By the Late Cretaceous, crocodylomorphs evolved a range of morphologies equalling Late Triassic crurotarsans. By contrast, biomechanical disparity in the Cretaceous did not increase, essentially decoupling from morphology. This enigmatic result could be attributed to biomechanical evolution in other anatomical regions (e.g. cranium, dentition or postcranium), possibly releasing the mandible from selective pressures. Overall, our analyses reveal a complex relationship between morphological and biomechanical disparity in Mesozoic crurotarsans that culminated in specialized feeding ecologies and associated lifestyles.

New Information on the Temnospondyl Calamops paludosus from the Upper Triassic of Pennsylvania

Sues, H.-D., and R. R. Schoch. 2013. Anatomy and phylogenetic relationships of Calamops paludosus (Temnospondyli, Stereospondyli) from the Triassic of the Newark Basin, Pennsylvania. Journal of Vertebrate Paleontology 33:1061-1070. DOI:10.1080/02724634.2013.759120

Abstract - The holotype of the large temnospondyl Calamops paludosus is the oldest known tetrapod fossil from the Triassic of the Newark basin in Pennsylvania. Although it is usually placed in Metoposauridae, its affinities have remained unknown since its original description because the unique specimen had never been prepared. Preparation and casting of the specimen, which comprises three pieces of a left mandibular ramus, now permits detailed anatomical description of the jaw and assessment of its affinities. Calamops paludosus is a valid taxon of trematosauroid temnospondyls that can be diagnosed by several autapomorphies. It represents one of the geologically youngest known records of long-snouted trematosaurs and the first record of these temnospondyls from the Late Triassic of North America.

A New Semionotiform Fish from the Upper Triassic Chinle Formation of Utah

Gibson, S. Z. 2013. A new hump-backed ginglymodian fish (Neopterygii, Semionotiformes) from the Upper Triassic Chinle Formation of southeastern Utah. Journal of Vertebrate Paleontology 33:1037-1050. DOI:10.1080/02724634.2013.758125

Abstract - A new species of hump-backed semionotiform fish, Lophionotus sanjuanensis, gen et sp. nov., is described based on specimens recently and previously collected from the Upper Triassic Church Rock Member of the Chinle Formation of southeastern Utah. It is characterized by a deep body with a large postcranial hump, and dense tuberculation on the posterodorsal margin of the skull that continues into the dorsal ridge and dorsolateral flank scales. The vertical preoperculum bears a short and broad paddle-like ventral process. The infraorbital series expands ventral to the suborbital and contacts the anterior ramus of the preoperculum, although this character has also been observed in other deep-bodied semionotiform taxa. This taxon represents the first newly described semionotiform fish species from the western United States in over 45 years, and adds to knowledge of Triassic fishes biodiversity.

A New Protorosaur from the Middle Triassic of China

Fraser, N. C., Rieppel, O., and L. Chun. 2013. A long-snouted protorosaur from the Middle Triassic of southern China. Journal of Vertebrate Paleontology 33:1120-1126 DOI:10.1080/02724634.2013.764310

Abstract -
A new protorosaur is described on the basis of a single specimen from the Ladinian of southern China. Although it has been greatly crushed, it still preserves clear details of the skull and axial skeleton. It possesses a neck that is longer than the trunk and is similar to tanystropheids in having 12 or 13 cervicals. Unusual among protorosaurs, the new form has an elongate snout. It also lacks a clear thyroid fenestra, although there is a slight separation of the pubis and ischium close to the pubic symphysis. The new form adds to the growing diversity and disparity of protorosaur taxa from the Middle Triassic of southern China.

Robotic Tanystropheus to Promote Marine Biology Conservation

Bone Diagenesis Study of Early Triassic Cynodonts from Argentina

Previtera, E., D'Angelo, J. A., and A. C. Mancuso. 2013. Preliminary chemometric study of bone diagenesis in Early Triassic cynodonts from Mendoza, Argentina. Ameghiniana 50:460-468.

Abstract - The non-mammalian therapsids dominated the terrestrial ecosystems during the Late Paleozoic–Early Mesozoic. The cynodonts have been studied from a taxonomic, osteologic and morphological perspective. However, taphonomy using chemometrics has been barely explored. This report includes a rib and an appendicular bone of cynodonts from the Puesto Viejo Group (Mendoza, Argentina). These fossils are studied for the first time using SEM-EDX. Semi-quantitative data derived from SEM-EDX spectra is evaluated by principal component analysis to gain new insights regarding the different diagenetic pathways of bone microstructure. The multivariate model supports the distinction of different sampled areas (bone, transition zone and rock matrix), in terms of chemical parameters. Differentiation is based mainly on varying contents of Ca, P, F, Si, Al, K, O, Mn and Fe. Variable concentrations of Fe and Mn could be related to different facies (floodplain and crevasse splay). These results along with thin section petrographical analysis confirm –in one of the cases– the substitution of hydroxyapatite by fluorapatite in the bone microstructure. Fossil-diagenetic processes observed herein include substitution, fracturing, brittle deformation and different permineralization events. Permineralization stages during burial history include infilling of vascular canals, trabeculae and fractures with hematite, manganite and calcite. The presence of calcite and iron enrichment indicates local reducing conditions below water-table during precipitation. This chemometric approach to the study of Triassic cynodont remains proved useful for assessing the chemical changes in bone microstructure.

Resumen -  Los terápsidos no-mamaliformes dominaron los ecosistemas terrestres durante el Paleozoico Tardío–Mesozoico Temprano. Los cinodontes han sido estudiados desde una perspectiva taxonómica, osteológica y morfológica. Sin embargo, análisis tafonómicos usando quimiometría son escasos. Esta contribución incluye una costilla y un hueso apendicular de cinodontes del Grupo Puesto Viejo (Mendoza, Argentina). Estos fósiles se analizaron por primera vez usando SEM-EDX. Los datos semi-cuantitativos derivados de SEM-EDX se evaluaron mediante análisis de componentes principales para obtener nuevos conocimientos sobre las diferentes vías diagenéticas de la microestructura ósea. El modelo multivariado apoya la distinción de las diferentes áreas de la muestra (hueso, zona de transición y matriz rocosa), en términos de parámetros químicos. La diferenciación se basa principalmente en diferentes contenidos de Ca, P, F, Si, Al, K, O, Mn y Fe. Concentraciones variables de Fe y Mn pueden asociarse a las diferentes facies (planicie de inundación y depósitos de desbordamiento). Estos resultados, junto con los análisis petrográficos de secciones delgadas confirman en uno de los casos la sustitución de la hidroxiapatita por fluorapatita en la microestructura ósea. Los procesos fósildiageneticos observados incluyen sustitución, fracturación, deformación frágil y diferentes eventos de permineralización. Las etapas de permineralización incluyen rellenos de hematita, manganita y calcita, en canales vasculares, trabéculas y fracturas, durante el enterramiento. La presencia de calcita y de hierro indica condiciones reductoras locales bajo el nivel freático durante la precipitación. Este enfoque quimiométrico para el estudio de cinodontes triásicos demostró ser útil para evaluar los cambios químicos de la microestructura ósea.

Postcranial Anatomy and Phylogenetic Relationships of the Sauropodomorph Mussaurus patagonicus

Otero, A. and D. Pol. 2013. Postcranial anatomy and phylogenetic relationships of Mussaurus patagonicus (Dinosauria: Sauropodomorpha). Journal of Vertebrate Paleontology 33:1138-1168.

Abstract - The transition from basal sauropodomorphs to sauropods is one of the most dramatic evolutionary transformations in the history of dinosaurs. Constituent taxa of this transition were recorded mainly in South Africa and South America, and to a lesser extent in North America. We describe here the postcranial anatomy of four specimens of basal sauropodomorphs from the Late Triassic of Patagonia, Argentina, and identify them as adult individuals of Mussaurus patagonicus. The material is composed of one subadult and three adult specimens and was originally identified as Plateosaurus. The completeness of the material provides more complete knowledge of this taxon and allows us to introduce aspects of basal sauropodomorph anatomy that were poorly understood until now, such as the configuration and arrangement of the distal carpal elements. The phylogenetic relationships of Mussaurus patagonicus are tested through a cladistic analysis of basal sauropodomorphs based on the anatomy of these specimens rather than on the post-hatchling and juvenile specimens previously known for this taxon. Mussaurus is recovered as a non-sauropod anchisaurian, being the sister group of Aardonyx plus more derived sauropodomorphs and is depicted outside the ‘quadrupedal clade,’ given the presence of plesiomorphic features such as a humerus/femur ratio <0.8, a curved femoral shaft in lateral view, and a nearly circular femoral midshaft cross-section. Mussaurus patagonicus adds new and valuable information that helps to clarify the core of the basal sauropodomorph-sauropod transition.

A New Silesaurid from the Middle Triassic of Zambia and Evidence for Rapid Diversification of Silesauridae

Peecook, B. R., Sidor, C. A., Nesbitt, S. J., Smith, R. M. H., Steyer, J. S., and K. D. Angielczyk. 2013. A new silesaurid from the upper Ntawere Formation of Zambia (Middle Triassic) demonstrates the rapid diversification of Silesauridae (Avemetatarsalia, Dinosauriformes) Journal of Vertebrate Paleontology 33:1127-1137 DOI:10.1080/02724634.2013.755991 

Abstract - Recent discoveries have shown that non-dinosaurian dinosauromorphs were morphologically diverse, globally distributed, and have a stratigraphic range extending into the Upper Triassic. Silesauridae, the sister group to Dinosauria, contains at least seven species. Here we describe Lutungutali sitwensis, gen. et sp. nov., the first silesaurid from the upper portion of the Ntawere Formation of the Luangwa Basin, Zambia. The upper Ntawere Formation has been correlated with subzone C of the Cynognathus Assemblage Zone of the Karoo Basin in South Africa and the Lifua Member of the Manda beds in the Ruhuhu Basin in Tanzania, both of which are considered Anisian in age and the latter has yielded the silesaurid Asilisaurus kongwe. The results of our phylogenetic analysis, including a new pelvic character, allies Lutungutali with Upper Triassic silesaurids such as Silesaurus, Sacisaurus, and Eucoelophysis rather than with the possibly coeval Asilisaurus. The Zambian silesaurid shares a laterally oriented brevis fossa on the ilium and a transversely thin ischium in cross-section with Upper Triassic forms. Silesaurids were more diverse during their early evolution in the Anisian than previously suspected. Lutungutali and Asilisaurus are the two oldest known members of the bird-line archosaurs represented by body fossils. Together they show that a subclade of bird-line archosaurs was diversifying soon after its origin, building further support for the rapid diversification of Archosauria in the wake of the Permo-Triassic extinction.