A Late Triassic Vertebrate Paleontology Extravaganza - ‘Late Triassic Terrestrial Biotas and the Rise of Dinosaurs’ Special Issue

It's out. The special volume in Earth and Environmental Science Transactions of the Royal Society of Edinburgh on Late Triassic Terrestrial Biotas and the Rise of Dinosaurs. These papers stem from the Society of Vertebrate Paleontology Symposium at the 2009 annual meeting in Bristol, U. K.  I've included a list of the papers and their abstracts below. There is an amazing amount of information here on Late Triassic vertebrates and early dinosaurs.  It will take me awhile to get through the papers, but for now I'd like to say it is an honor to be included in this volume.  I think Randy, Richard, Max, and the staff of EESTRSE did an excellent job in the production of this volume and I hope everyone who has an interest in Late Triassic terrestrial biotas enjoys the volume and finds the papers interesting and useful.

Issue cover featuring artwork by Jeff Martz.

Table of Contents - Volume 101 - Special Issue 3-4 (Late Triassic Terrestrial Biotas and the Rise of Dinosaurs)

Abstract - The early Mesozoic records an important transition in the history of the Earth’s terrestrial ecosystems. As they recovered from the largest known mass extinction (the end-Permian event), organisms in these ecosystems transitioned to new forms that eventually evolved into the classic Mesozoic biotas, and laid the foundations for many groups still flourishing today (Fraser 2006; Irmis & Whiteside 2010; Sues & Fraser 2010). All of this was set against a backdrop of dynamic climatic and physical events that shaped these biotas. This early Mesozoic terrestrial transition reached its culmination in many ways during the Late Triassic, when ecosystems had largely recovered from the end-Permian extinction, but had not yet been affected by the end-Triassic mass extinction (Fraser & Sues this volume). Thus, we see a combination of taxa, with some groups that would not survive the end of the Triassic living alongside early representatives of lineages that flourished later in the Mesozoic (e.g., Fraser 2006; Irmis et al. 2007; Brusatte et al. 2008; Sues & Fraser 2010, this volume) and in some cases are still diverse today. Just one example of this transition, recorded during the Late Triassic, is the origin and diversification of non-avian dinosaurs, the iconic representatives of Mesozoic terrestrial ecosystems (Brusatte et al. 2010; Langer et al. 2010). Although small and rare components of their respective biotas when they first evolved ∼231 Ma, dinosaurs were abundant and had a near-worldwide distribution by the beginning of the Jurassic Period (∼201·3 Ma).

Abstract - The first appearance of dinosaurs during the early Late Triassic coincided with marked faunal changes in terrestrial ecosystems. Most of the major groups of extant tetrapods (or their proximate sister-taxa), including mammaliaforms, crocodyliform archosaurs, lepidosaurs and turtles, also first appeared in the fossil record during the Late Triassic. On the other hand, a number of Palaeozoic ‘holdovers’, such as procolophonid parareptiles, dicynodont therapsids and many groups of temnospondyls, vanished near or at the end of the Triassic. The tempo and mode of this faunal turnover have long been debated, but there has been growing acceptance of a rather sudden event, although the precise dating of such an event remains controversial. However, new discoveries have cast doubt on this assumption. The persistence of non-dinosaurian dinosauromorphs alongside dinosaurs well into Norian times hints at a more protracted turnover. New data on Triassic insect assemblages indicate that turnover among insects may also have been more protracted and possibly not co-incident with the faunal changes among tetrapods. Future work directed toward improved absolute age assessments for major faunal assemblages will be critical for a better understanding of the transition from therapsid-dominated to dinosaur-dominated communities during the early Mesozoic.

Abstract - The Newark-APTS established a high-resolution framework for the Late Triassic and Early Jurassic. Palaeomagnetic polarity correlations to marine sections show that stage-level correlations of continental sequences were off by as much as 10 million years. New U–Pb ages show the new correlations and the Newark basin astrochronology to be accurate. Correlation of Newark-APTS to the Chinle Formation/Dockum Group, Glen Canyon Group, Fleming Fjord Formation and Ischigualasto Formation led to the following conclusions: (1) there are no unequivocal Carnian-age dinosaurs; (2) the Norian Age was characterised by a slowly increasing saurischian diversity but no unequivocal ornithischians; (3) there was profound Norian and Rhaetian continental provinciality; (4) the classic Chinle-, Germanic- and Los Colorados-type assemblages may have persisted to the close of the Rhaetian; (5) the distinct genus-level biotic transition traditionally correlated with the marine Carnian–Norian is in fact mid-Norian in age and within published error of the Manicouagan impact; (6) the end-Triassic marine and continental extinctions as seen in eastern North America were contemporaneous; and (7) compared to Triassic communities, Hettangian and Sinemurian age terrestrial communities were nearly globally homogenous and of low diversity. Consequently, the complex emerging picture of dinosaur diversification demands biostratigraphically-independent geochronologies in each of the faunally-important regions.

Abstract - Recent stratigraphic revisions of the Upper Triassic Chinle Formation of Petrified Forest National Park, in conjunction with precise and accurate documentation of fossil tetrapod occurrences, clarified the local biostratigraphy, with regional and global implications. A significant overlap between Adamanian and Revueltian faunas is rejected, as is the validity of the Lamyan sub-land vertebrate faunachron. The Adamanian–Revueltian boundary can be precisely placed within the lower Jim Camp Wash beds of the Sonsela Member and thus does not occur at the hypothesised Tr-4 unconformity. This mid-Norian faunal turnover, may coincide with a floral turnover, based on palynology studies and also on sedimentological evidence of increasing aridity. Available age constraints bracketing the turnover horizon are consistent with the age of the Manicouagan impact event. The rise of dinosaurs in western North America did not correspond to the Adamanian–Revueltian transition, and overall dinosauromorph diversity seems to have remained at a constant level across it. The paucity of detailed Late Triassic vertebrate biostratigraphic data and radioisotopic dates makes it currently impossible to either support or reject the existence of globally synchronous Late Triassic extinctions for tetrapods.
5) Sulej, T., Bronowicz, R., Tałanda, M., and G. Niedźwiedzki. A new dicynodont–archosaur assemblage from the Late Triassic (Carnian) of Poland.

Abstract - This paper reports a new assemblage from the Late Triassic (mid–late Carnian) at Woźniki near Częstochowa (Poland). The Woźniki vertebrate assemblage is similar to that of Lisowice–Lipie Śląskie, a new locality bearing vertebrates from latest Triassic (latest Norian–early Rhaetian) strata of southern Poland, in the presence of dicynodonts, shark spines, plagiosaurs and a cyclotosaur, but conchostracans and bivalves are similar to those from the Krasiejów site (late Carnian). The most complete specimen from Woźniki belongs to a dicynodont, and consists of cranial and postcranial bones of a single individual. It demonstrates that large dicynodonts were part of the Late Triassic vertebrate assemblage in Central Europe. Numerous tetrapod tracks and traces are associated with skeletal fossils at Woźniki.

Abstract - The two major lineages of extant archosaurs, crocodylians and avians, diverged in the Triassic, but the details and timing of this event are incompletely understood. Fragmentary and phylogenetically uninformative specimens, in addition to poor temporal constraints on rock units from the Early and Middle Triassic, typify obstacles in identifying early archosaurs. This paper re-describes the partial skeleton of the only known specimen of Xilousuchus sapingensis Wu, 1981 from the Early Triassic Heshanggou Formation in north-central China. Originally assigned to the non-archosaurian archosauriform clade Proterosuchidae, an extensive phylogenetic analysis posits X. sapingensis as a crown-group archosaur within Suchia, thus making this taxon the unequivocally oldest known member of Archosauria. The age and phylogenetic position of X. sapingensis indicate that many archosaurs, including all major clades of non-archosaurian archosauriforms, the avianline, ornithosuchids, aetosaurs and paracrocodylomorph lineages, must have diverged by the end of the Early Triassic. X. sapingensis is part of a possible clade of sail-backed poposauroids that were common components of archosaur assemblages during the Early to Middle Triassic.

Abstract - Saltopus elginensis, reported in 1910 from the yellow sandstones of the Lossiemouth Sandstone Formation (Late Triassic) of Morayshire, NE Scotland, has long been controversial. It was described first as a theropod dinosaur, but others disagreed. Reanalysis of the type, and only, specimen using casts from the natural rock moulds, as well as X-rays and CT scans, has revealed new anatomical data not available to previous researchers. Saltopus was a small, 800–1000 mm-long biped, whose tail made up more than half its length. It is an avemetatarsalian because it has elongated and tightly bunched metatarsals, the tibia is longer than the femur, the calcaneal tuber is rudimentary or absent, and metatarsal II is equal to or longer than metatarsal IV; a unique assemblage of characters diagnosing this clade. Saltopus is a dinosauromorph on the basis of the reduced fingers IV and V, the saddle-shaped dorsal margin of the iliac blade, and the articulation of sacral rib 1 close to the front of the iliac blade. Saltopus is a dinosauriform on the basis of the trochanteric shelf and lesser trochanter on the proximal end of the femur, the waisted sacral ribs, and perhaps the rod-like and straight pubis. However, it lacks all apomorphies of Dinosauria, retaining for example the primitive condition of two sacral vertebrae. Cladistic analyses place Saltopus within Dinosauromorpha and Dinosauriformes, and between the basal dinosauriform Pseudolagosuchus and the derived clade consisting of Silesauridae and Dinosauria, so making it one of a radiation of small pre-dinosaurian bipedal archosaurs in the Triassic found so far in North and South America and in Europe.

Abstract - The dinosaur record of the Santa Maria beds of Rio Grande do Sul (Mid–Late Triassic; south Brazil) includes the herrerasaur Staurikosaurus pricei, and two basal members of the sauropodomorph lineage: Saturnalia tupiniquim and Unaysaurus tolentinoi. The most enigmatic of the saurischian taxa is, however, Guaibasaurus candelariensis, previously regarded as either a basal theropod or a basal sauropodomorph. This study provides a detailed anatomical revision of all specimens originally referred to G. candelariensis by Bonaparte and co-authors, including its type-series and a more recently excavated partial postcranium. Although coming from different sites, these specimens share a unique combination of traits, and at least one possible autapomorphic feature of the pelvis, which support the inclusivity and uniqueness of the species. G. candelariensis was a medium-sized (nearly 2 m long) biped with an intriguing mix of plesiomorphic and derived (eusaurischian/theropod) features. Phylogenetic studies reveal weak support for the nesting of G. candelariensis within Theropoda, but this affinity is bolstered by various traits it shares with neotheropods. The Norian age of G. candelariensis corroborates previous studies that suggest the continuous radiation of more basal dinosauromorphs until the end of the Triassic, after the appearance of the three main dinosaur clades.

Abstract - The beginning of dinosaur evolution is currently known based on a handful of highly informative Gondwanan outcrops of Ischigualastian age (late Carnian–early Norian). The richest Triassic dinosaur records of the southern continents are those of South America and South Africa, with taxonomically diverse faunas, whereas faunas from India and central Africa are more poorly known. Here, the known diversity of Gondwanan Triassic dinosaurs is increased with new specimens from central India, which allow a more comprehensive characterisation of these dinosaur assemblages. Five dinosauriform specimens are reported from the probable late Norian–earliest Rhaetian Upper Maleri Formation, including two new sauropodomorph species, the non-plateosaurian Nambalia roychowdhurii and the plateosaurian Jaklapallisaurus asymmetrica, a guaibasaurid and two basal dinosauriforms. The Lower Dharmaram Formation, probably latest Norian–Rhaetian in age, includes basal sauropodomorph and neotheropod remains, providing the second record of a Triassic Gondwanan neotheropod. The currently available evidence suggests that the oldest known Gondwanan dinosaur assemblages (Ischigualastian) were not homogeneous, but more diverse in South America than in India. In addition, the Upper Maleri and Lower Dharmaram dinosaur assemblages resemble purported coeval South American and European beds in the presence of basal sauropodomorphs. Accordingly, the current available evidence of the Triassic beds of the Pranhita–Godavari Basin suggests that dinosaurs increased in diversity and abundance during the late Norian to Rhaetian in this region of Gondwana.

10) Porro, L. B., Butler, R. J., Barrett, P. M., Moore-Fay, S., and R. L. Abel. New heterodontosaurid specimens from the Lower Jurassic of southern Africa and the early ornithischian dinosaur radiation.

Abstract - Heterodontosaurids are poorly understood early ornithischian dinosaurs with extensive geographic and stratigraphic ranges. The group is best known from the Lower Jurassic upper ‘Stormberg Group’ (upper Elliot and Clarens formations) of southern Africa, previously represented by at least three distinct species and ten described specimens. This paper describes four additional heterodontosaurid specimens from southern Africa. A partial skull of a large individual of Heterodontosaurus tucki (NM QR 1788) is approximately 70 longer than that of the type specimen of Heterodontosaurus, and provides new information on allometric changes in mandibular morphology during growth in this taxon. It is the largest known heterodontosaurid cranial specimen, representing an individual approximately 1·75 metres in length, and perhaps 10 kg in body mass. NHMUK R14161 is a partial skull that appears to differ from all other heterodontosaurids on the basis of the proportions of the dentaries, and may represent an unnamed new taxon. Two additional partial skulls (NHMUK RU C68, NHMUK RU69) are referred to cf. Lycorhinus. At least four, and possibly five or more, heterodontosaurid species are present in the upper ‘Stormberg’. This high diversity may have been achieved by dietary niche partitioning, and suggests an adaptive radiation of small-bodied ornithischians following the end Triassic extinctions.

11) Brusatte, S. L., Benton, M. J., Lloyd, G. T., Ruta, M., and S. C. Wang. Macroevolutionary patterns in the evolutionary radiation of archosaurs (Tetrapoda: Diapsida).

Abstract - The rise of archosaurs during the Triassic and Early Jurassic has been treated as a classic example of an evolutionary radiation in the fossil record. This paper reviews published studies and provides new data on archosaur lineage origination, diversity and lineage evolution, morphological disparity, rates of morphological character change, and faunal abundance during the Triassic–Early Jurassic. The fundamental archosaur lineages originated early in the Triassic, in concert with the highest rates of character change. Disparity and diversity peaked later, during the Norian, but the most significant increase in disparity occurred before maximum diversity. Archosaurs were rare components of Early–Middle Triassic faunas, but were more abundant in the Late Triassic and pre-eminent globally by the Early Jurassic. The archosaur radiation was a drawn-out event and major components such as diversity and abundance were discordant from each other. Crurotarsans (crocodile-line archosaurs) were more disparate, diverse, and abundant than avemetatarsalians (bird-line archosaurs, including dinosaurs) during the Late Triassic, but these roles were reversed in the Early Jurassic. There is no strong evidence that dinosaurs outcompeted or gradually eclipsed crurotarsans during the Late Triassic. Instead, crurotarsan diversity decreased precipitously by the end-Triassic extinction, which helped usher in the age of dinosaurian dominance. 

12) Barrett, P. M., Butler, R. J., and S. J. Nesbitt. The roles of herbivory and omnivory in early dinosaur evolution.

Abstract - Herbivorous and omnivorous dinosaurs were rare during the Carnian stage of the Late Triassic. By contrast, the succeeding Norian stage witnessed the rapid diversification of sauropodomorphs and the rise of the clade to ecological dominance. Ornithischians, by contrast, remained relatively rare components of dinosaur assemblages until much later in the Mesozoic. The causes underlying the differential success of ornithischians and sauropodomorphs remain unclear, but might be related to trophic specialisation. Sauropodomorphs replaced an established herbivore guild consisting of rhynchosaurs, aetosaurs and basal synapsids, but this faunal turnover appears to have been opportunistic and cannot be easily attributed to either competitive interactions or responses to floral change. Consideration of diversity patterns and relative abundance suggests that the ability to eat plants might have been a major factor promoting sauropodomorph success, but that it was less important in the early evolution of Ornithischia. On the basis of current evidence it is difficult to determine the diet of the ancestral dinosaur and scenarios in which omnivory or carnivory represent the basal condition appear equally likely.

13) Irmis, R. B. Evaluating hypotheses for the early diversification of dinosaurs.

Abstract - Many hypotheses have been proposed for the rise of dinosaurs, but their early diversification remains poorly understood. This paper examines the occurrences, species diversity and abundance of early dinosaurs at both regional and global scales to determine patterns of their early evolutionary history. Four main patterns are clear: (1) sauropodomorph dinosaurs became abundant during the late Norian–Rhaetian of Gondwana and Europe; (2) Triassic dinosaurs of North America have low species diversity and abundance until the beginning of the Jurassic; (3) sauropodomorphs and ornithischians are absent in the Triassic of North America; and (4) ornithischian dinosaurs maintain low species diversity, relative abundance and small body size until the Early Jurassic. No one hypothesis fully explains these data. There is no evidence for a Carnian–Norian extinction event, but sauropodomorphs did become abundant during the Norian in some assemblages. No clear connection exists between palaeoenvironment and early dinosaur diversity, but environmental stress at the Triassic–Jurassic boundary is consistent with changes in North American dinosaur assemblages. Elevated growth rates in dinosaurs are consistent with the gradual phyletic increase in body size. This study demonstrates that early dinosaur diversification was a complex process that was geographically diachronous and probably had several causes.

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