Re-Examination of Some Large Early Mesozoic Tetrapod Footprints from the African Collection of Paul Ellenberger

The Late Triassic—Early Jurassic ichnofauna described mainly by Paul Ellenberger from southern Africa (Lesotho) is a valuable window on first phases of dinosaur diversification. Unfortunately, the present taxonomic status of several forms from that ichnofauna is unclear. The revision of this material has been frequently invoked and partially done without reaching many definitive results. Due to the enormous amount of data, a global revision seems at present impossible and must be accomplished in smaller steps. A small number of Ellenberger's ichnogenera including Tetrasauropus, Pseudotetrasauropus, Pentasauropus, Paratetrasauropus, Sauropodopus and Deuterosauropodopus, which different authors have ascribed to basal sauropodomorphs, are here revised in a consistent manner and their attribution to osteological clades is considered. Tetrasauropus and Pseudotetrasauropus are here validated as the only ichnotaxa related to sauropodomorphs. Pentasauropus is retained as valid, and a therapsid trackmaker is suggested. Paratetrasauropus and Sauropodopus are also validated and ascribed to non-dinosaurian trackmakers, and Deuterosauropodopus is synonymized with Sauropodopus.     

Giants on the landscape: modelling the abundance of megaherbivorous dinosaurs of the Morrison Formation (Late Jurassic,western USA)

The ecosystem impact of megaherbivorous dinosaurs of the Morrison Formation would have depended on their abundance (number of animals per unit of habitat area) on the landscape. We constrain Morrison megaherbivore abundance by modelling dinosaur abundance in terms of carrying capacity (K), average body mass (ABM) and animal's energy needs. Two kinds of model are presented: ‘demand-side’ models that estimate K in terms of the aggregate energy demand of the dinosaur community, and ‘supply-side’ models that estimate K in terms of retrodicted primary productivity. Baseline values of K, ABM and energy needs for the models are further derived from comparisons with modern large herbivores, and from the composition of the megaherbivore fauna from a particular stratigraphic interval of the Morrison, but in all models a broad range of fractions and multiples of these baseline parameters are considered. ‘Best-guess’ estimates of Morrison megaherbivore abundance suggest an upper limit of a few hundred animals across all taxa and size classes per square kilometre, and up to a few tens of individuals of large subadults and adults.     

新疆准噶尔盆地侏罗纪石树沟组上部一大型兽脚类跖骨

准噶尔盆地中侏罗世晚期到晚侏罗世早期沉积的石树沟组产出过包括多种兽脚类恐龙在内的大量脊椎动物化石。描述了一件新的采自准噶尔盆地东北缘五彩湾地区石树沟组上部的兽脚类恐龙左第四跖骨标本(IVPPV18060)。通过与其他兽脚类恐龙第四跖骨对比,该标本可归入异特龙超科(Allosauroidea)。它与其他异特龙超科第四跖骨的相似性包括:近端关节面三角形并有一向后方延伸的舌状突出,向外侧轻微弯曲的骨干,位于骨干后方靠近外侧边的半月形肌肉凹陷,三角形骨干横截面。在异特龙超科当中,V18060与准噶尔盆地将军庙地区石树沟组中发现的董氏中华盗龙(Sinraptordongi)最为相似(除了以上相似性,V18060和董氏中华盗龙的相似性还包括远端关节轮廓呈近梯形,骨干后部肌肉凹陷与周围边界呈半开放状态)。但是,V18060和董氏中华盗龙也存在着一些明显的区别:V18060明显比董氏中华盗龙的第四跖骨粗壮,远端内外两髁大小差异相反。这些形态差异可能是由于个体发育或者性双形造成的,也可能代表分类学差异。对比研究和基于第四跖骨形态信息进行的分支系统学分析结果更偏向后一种解释:V18060代表一个不同于董氏中华盗龙但与后者亲缘关系很近的新的兽脚类恐龙。新材料的发现增加了中晚侏罗世石树沟动物群兽脚类恐龙的分异度。在五彩湾和将军庙地区发现不同的中华盗龙类恐龙指示了晚侏罗世早期准噶尔盆地中相邻地区可能存在着一定的生态分异性或者地理隔离。     

No gastric mill in sauropod dinosaurs: new evidence from analysis of gastrolith mass and function in ostriches

Polished pebbles occasionally found within skeletons of giant herbivorous sauropod dinosaurs are very likely to be gastroliths (stomach stones). Here, we show that based on feeding experiments with ostriches and comparative data for relative gastrolith mass in birds, sauropod gastroliths do not represent the remains of an avian-style gastric mill. Feeding experiments with farm ostriches showed that bird gastroliths experience fast abrasion in the gizzard and do not develop a polish. Relative gastrolith mass in sauropods (gastrolith mass much less than 0.1% of body mass) is at least an order of magnitude less than that in ostriches and other herbivorous birds (gastrolith mass approximates 1% of body mass), also arguing against the presence of a gastric mill in sauropods. Sauropod dinosaurs possibly compensated for their limited oral processing and gastric trituration capabilities by greatly increasing food retention time in the digestive system. Gastrolith clusters of some derived theropod dinosaurs (oviraptorosaurs and ornithomimosaurs) compare well with those of birds, suggesting that the gastric mill evolved in the avian stem lineage.     

Avian-like breathing mechanics in maniraptoran dinosaurs

In 1868 Thomas Huxley first proposed that dinosaurs were the direct ancestors of birds and subsequent analyses have identified a suite of 'avian' characteristics in theropod dinosaurs. Ossified uncinate processes are found in most species of extant birds and also occur in extinct non-avian maniraptoran dinosaurs. Their presence in these dinosaurs represents another morphological character linking them to Aves, and further supports the presence of an avian-like air-sac respiratory system in theropod dinosaurs, prior to the evolution of flight. Here we report a phylogenetic analysis of the presence of uncinate processes in Aves and non-avian maniraptoran dinosaurs indicating that these were homologous structures. Furthermore, recent work on Canada geese has demonstrated that uncinate processes are integral to the mechanics of avian ventilation, facilitating both inspiration and expiration. In extant birds, uncinate processes function to increase the mechanical advantage for movements of the ribs and sternum during respiration. Our study presents a mechanism whereby uncinate processes, in conjunction with lateral and ventral movements of the sternum and gastral basket, affected avian-like breathing mechanics in extinct non-avian maniraptoran dinosaurs.     

Fossil quality and naming dinosaurs

The intense interest in dinosaurs through the past 30 years might have led to an increase in poor practice in naming new species. A review of the data shows that the reverse is the case. For 130 years, from the 1820s to the 1950s, most new species of dinosaurs were based on scrappy and incomplete material. After 1960, the majority of new species have been based on complete skulls or skeletons, and sometimes on materials from several individuals. This switch in the quality of type specimens corresponds to the recent explosive renaissance of interest in dinosaurs, during which the number of new species named per year has risen, from three or four in the 1950s, to thirty or more today. The pattern of specimen quality varies by continent, with the highest proportion of new species based on good material in North America, then Asia, then South America, then Africa and finally Europe. This ranking reflects a complex pattern of perhaps overstudy in Europe, immensely rich reserves of new dinosaur materials in North America and Asia, and a relative paucity in South America and Africa.     

Protracted growth impedes the detection of sexual dimorphism in non‐avian dinosaurs

Evidence for sexual dimorphism is extremely limited in the non‐avian dinosaurs despite their high diversity and disparity, and despite the fact that dimorphism is very common in vertebrate lineages of all kinds. Using body‐size data from both Alligator mississippiensis and Rhea americana, which phylogenetically bracket the dinosaurs, we demonstrate that even when there is strong dimorphism in a species, random sampling of populations of individuals characterized by sustained periods of growth (as in the alligator and most dinosaurs) can result in the loss of this signal. Dimorphism may be common in fossil taxa but very hard to detect without ontogenetic age control and large sample sizes, both of which are hampered by the limitations of the fossil record. Signal detection may be further hindered by Type III survivorship, whereby increased mortality among the young favours the likelihood that they will be sampled (unless predation or taphonomic bias against small size acts against this). These, and other considerations relating to behaviour and ecology, provide powerful reasons to suggest that sexual dimorphism in dinosaurs may be very difficult to detect in almost all currently available samples. Similar issues are likely also to be applicable to many fossil reptiles, or animals more generally.     

Biotic and environmental dynamics through the Late Jurassic–Early Cretaceous transition: evidence for protracted faunal and ecological turnover

The Late Jurassic to Early Cretaceous interval represents a time of environmental upheaval and cataclysmic events, combined with disruptions to terrestrial and marine ecosystems. Historically, the Jurassic/Cretaceous (J/K) boundary was classified as one of eight mass extinctions. However, more recent research has largely overturned this view, revealing a much more complex pattern of biotic and abiotic dynamics than has previously been appreciated. Here, we present a synthesis of our current knowledge of Late Jurassic–Early Cretaceous events, focusing particularly on events closest to the J/K boundary. We find evidence for a combination of short‐term catastrophic events, large‐scale tectonic processes and environmental perturbations, and major clade interactions that led to a seemingly dramatic faunal and ecological turnover in both the marine and terrestrial realms. This is coupled with a great reduction in global biodiversity which might in part be explained by poor sampling. Very few groups appear to have been entirely resilient to this J/K boundary ‘event’, which hints at a ‘cascade model’ of ecosystem changes driving faunal dynamics. Within terrestrial ecosystems, larger, more‐specialised organisms, such as saurischian dinosaurs, appear to have suffered the most. Medium‐sized tetanuran theropods declined, and were replaced by larger‐bodied groups, and basal eusauropods were replaced by neosauropod faunas. The ascent of paravian theropods is emphasised by escalated competition with contemporary pterosaur groups, culminating in the explosive radiation of birds, although the timing of this is obfuscated by biases in sampling. Smaller, more ecologically diverse terrestrial non‐archosaurs, such as lissamphibians and mammaliaforms, were comparatively resilient to extinctions, instead documenting the origination of many extant groups around the J/K boundary. In the marine realm, extinctions were focused on low‐latitude, shallow marine shelf‐dwelling faunas, corresponding to a significant eustatic sea‐level fall in the latest Jurassic. More mobile and ecologically plastic marine groups, such as ichthyosaurs, survived the boundary relatively unscathed. High rates of extinction and turnover in other macropredaceous marine groups, including plesiosaurs, are accompanied by the origin of most major lineages of extant sharks. Groups which occupied both marine and terrestrial ecosystems, including crocodylomorphs, document a selective extinction in shallow marine forms, whereas turtles appear to have diversified. These patterns suggest that different extinction selectivity and ecological processes were operating between marine and terrestrial ecosystems, which were ultimately important in determining the fates of many key groups, as well as the origins of many major extant lineages. We identify a series of potential abiotic candidates for driving these patterns, including multiple bolide impacts, several episodes of flood basalt eruptions, dramatic climate change, and major disruptions to oceanic systems. The J/K transition therefore, although not a mass extinction, represents an important transitional period in the co‐evolutionary history of life on Earth.     

The Fossil Record of Vertebrate Tracks in Mexico

The fossil record of vertebrate tracks in Mexico is taxonomically rich and geographically diverse, and ranges in age from Middle Jurassic to Pleistocene and possibly Holocene. Middle Jurassic saurischian dinosaur tracks from Oaxaca represent the oldest record of vertebrate tracks in Mexico. Tracks attributed to Late Jurassic theropods and ornithopods are known from Michoacán. Theropod tracks of supposed Jurassic age are known from Durango. Lower Cretaceous Saurischian and ornithopod tracks are known from southern Puebla. Central Mexico has yielded Late Cretaceous hadrosaurid and sauropod tracks (Puebla), and tracks belonging to Theropoda and Ornithopoda (El Aguaje, Michoacan). In Coahuila several tracksites occur within the Cerro del Pueblo Formation and include tracks of mesoreptiles, pterosaurs, small to large theropods, bipedal and quadrupedal ornithopods, birds and possibly a mammal. A tracksite in the Olmos Formation includes tracks of turtles, crocodilians, medium-sized theropods, small ornithopods, and birds. Bird tracks of supposed Eocene age are known from Sinaloa. Two important Pleistocene ichnofaunas occur in the states of Puebla (Tepexi de Rodríguez) and Jalisco (San Juan de los Lagos), and include tracks of camelids, artiodactyls, small ungulates, elephants, large felids and birds. Pleistocene human tracks occur in Coahuila and Jalisco. Mexico's track record provides important insights into vertebrate diversity, paleoecology, and paleoenvironments. Given the rate of new discoveries since 1998, there is no doubt that new sites await to be found.