Evolutionary relationships and systematics of Atoposauridae (Crocodylomorpha: Neosuchia): implications for the rise of Eusuchia

Atoposaurids are a group of small-bodied, extinct crocodyliforms, regarded as an important component of Jurassic and Cretaceous Laurasian semi-aquatic ecosystems. Despite the group being known for over 150 years, the taxonomic composition of Atoposauridae and its position within Crocodyliformes are unresolved. Uncertainty revolves around their placement within Neosuchia, in which they have been found to occupy a range of positions from the most basal neosuchian clade to more crownward eusuchians. This problem stems from a lack of adequate taxonomic treatment of specimens assigned to Atoposauridae, and key taxa such as Theriosuchus have become taxonomic ‘waste baskets’. Here, we incorporate all putative atoposaurid species into a new phylogenetic data matrix comprising 24 taxa scored for 329 characters. Many of our characters are heavily revised or novel to this study, and several ingroup taxa have never previously been included in a phylogenetic analysis. Parsimony and Bayesian approaches both recover Atoposauridae as a basal clade within Neosuchia, more stemward than coelognathosuchians, bernissartiids, and paralligatorids. Atoposauridae is a much more exclusive clade than previously recognized, comprising just three genera (Alligatorellus, Alligatorium, and Atoposaurus) that were restricted to the Late Jurassic of western Europe, and went extinct at the Jurassic/Cretaceous boundary. A putative Gondwanan atoposaurid (Brillanceausuchus) is recovered as a paralligatorid. Our results exclude both Montsecosuchus and Theriosuchus from Atoposauridae. Theriosuchus is polyphyletic, forming two groupings of advanced neosuchians. Theriosuchus (restricted to Theriosuchus pusillus, Theriosuchus guimarotae, and Theriosuchus grandinaris) spanned the Middle Jurassic to early Late Cretaceous, and is known from Eurasia and North Africa. Two Cretaceous species previously assigned to Theriosuchus (‘Theriosuchus’ ibericus and ‘Theriosuchus’ sympiestodon) are shown to be nested within Paralligatoridae, and we assign them to the new genus Sabresuchus. The revised phylogenetic placement of Theriosuchus has several implications for our understanding of eusuchian evolution. Firstly, the presence of fully pterygoidean choanae, previously regarded as a defining characteristic of Eusuchia, is not found in some basal members of Eusuchia. However, eusuchians can be distinguished from Theriosuchus and other basal neosuchians in that their choanae are posteriorly positioned, with an anterior margin medial to the posterior edge of the suborbital fenestra. This feature distinguishes eusuchians from Theriosuchus and more basal neosuchians. Secondly, our refined understanding of Theriosuchus implies that this taxon possessed only amphicoelous presacral vertebrae, and therefore fully developed vertebral procoely is likely to have evolved only once in Crocodylomorpha, on the lineage leading to Eusuchia. These and other findings presented herein will provide an important framework for understanding the neosuchian–eusuchian transition

Limb-Bone Scaling Indicates Diverse Stance and Gait in Quadrupedal Ornithischian Dinosaurs

Background The most primitive ornithischian dinosaurs were small bipeds, but quadrupedality evolved three times independently in the clade. The transition to quadrupedality from bipedal ancestors is rare in the history of terrestrial vertebrate evolution, and extant analogues do not exist. Constraints imposed on quadrupedal ornithischians by their ancestral bipedal bauplan remain unexplored, and consequently, debate continues about their stance and gait. For example, it has been proposed that some ornithischians could run, while others consider that none were cursorial. Methodology/Principal Findings Drawing on biomechanical concepts of limb bone scaling and locomotor theory developed for extant taxa, we use the largest dataset of ornithischian postcranial measurements so far compiled to examine stance and gait in quadrupedal ornithischians. Differences in femoral midshaft eccentricity in hadrosaurs and ceratopsids may indicate that hadrosaurs placed their feet on the midline during locomotion, while ceratopsids placed their feet more laterally, under the hips. More robust humeri in the largest ceratopsids relative to smaller taxa may be due to positive allometry in skull size with body mass in ceratopsids, while slender humeri in the largest stegosaurs may be the result of differences in dermal armor distribution within the clade. Hadrosaurs are found to display the most cursorial morphologies of the quadrupedal ornithischian cades, indicating higher locomotor performance than in ceratopsids and thyreophorans. Conclusions/Significance Limb bone scaling indicates that a previously unrealised diversity of stances and gaits were employed by quadrupedal ornithischians despite apparent convergence in limb morphology. Grouping quadrupedal ornithischians together as a single functional group hides this disparity. Differences in limb proportions and scaling are likely due to the possession of display structures such as horns, frills and dermal armor that may have affected the center of mass of the animal, and differences in locomotor behaviour such as migration, predator escape or home range size

Re-assessment of the Late Jurassic eusauropod dinosaur Hudiesaurus sinojapanorum Dong, 1997, from the Turpan Basin, China, and the evolution of hyperrobust antebrachia in sauropods

Hudiesaurus sinojapanorum is a Late Jurassic sauropod from northwestern China that was erected on the basis of a cervicodorsal vertebra, four teeth, and a nearly complete forelimb. However, re-evaluation of this material, and comparisons with other taxa, indicate that there are few grounds for regarding these specimens as congeneric. Consequently, although we retain the vertebra as the holotype specimen of Hudiesaurus, the forelimb is assigned to a new taxon—Rhomaleopakhus turpanensis, gen. et sp. nov. The teeth previously referred to Hudiesaurus are poorly preserved but resemble those of several other ‘core Mamenchisaurus-like taxa’ (CMTs) from East Asia, such as Mamenchisaurus sinocanadorum. Phylogenetic analyses confirm that Hudiesaurus is a CMT and the sister taxon of Xinjiangtitan. Despite some uniquely shared features, their large size, and close geographic provenance, Hudiesaurus and Xinjiangtitan are retained as distinct genera based on their stratigraphic separation and numerous anatomical differences. Rhomaleopakhus is also shown to be a CMT in all analyses, being most closely related to Chuanjiesaurus and Analong. We link the convergent evolution of robust antebrachia and an enlarged olecranon in CMTs, titanosaurs, and some ornithischians (e.g., ceratopsids) to a more flexed orientation of the forearm, an enhanced role for the forelimb in locomotion, and an anterior shift in the whole-body center of mass. CMTs and titanosaurs potentially converged on a feeding strategy in which the ability to increase browse height via bipedal rearing was sacrificed in return for more efficient locomotion that improved travel between patchily distributed food sources

How to date a crocodile – estimation of neosuchian clade ages and a comparison of four time-scaling methods

Clade ages within the crocodylomorph clade Neosuchia have long been debated. Molecular and morphological studies have yielded remarkably divergent results. Despite recent advances, there has been no comprehensive relative comparison of the major time calibration methods available to estimate clade ages based on morphological data. We used four methods (cal3, extended Hedman, smoothed ghost lineage analysis (sGLA) and the fossilized birth–death model (FBD)) to date clade ages derived from a published crocodylomorph supertree and a new neosuchian phylogeny. All time-scaling methods applied here agree on the origination of Neosuchia during the Late Triassic or Early Jurassic, and the presence of the major extant eusuchian groups (Crocodyloidea, Gavialoidea, Alligatoroidea and Caimaininae) by the end of the Late Cretaceous. The number of distinct lineages present before the K/Pg boundary is less certain, with support for two competing scenarios in which Crocodylinae, Tomistominae and Diplocynodontinae either: (1) diverged from other eusuchian lineages before the K/Pg boundary; or (2) evolved during a ‘burst’ of diversification after the K/Pg event. Cal3 and FBD proved to be the most suitable methods for time-scaling phylogenetic trees dominated by fossil taxa. Extended Hedman estimates are substantially older than the others, with larger standard deviations and a strong sensitivity to taxon sampling and topological changes; sGLA has similar problems. We conclude that a detailed understanding of phylogenetic relationships, tree reconstruction methods, and good taxonomic coverage (in particular the inclusion of the oldest taxon in each clade) is essential when evaluating the results of such dating analyses

Ten more years of discovery: revisiting the quality of the sauropodomorph dinosaur fossil record

Spatiotemporal changes in fossil specimen completeness can bias our understanding of a group's evolutionary history. The quality of the sauropodomorph fossil record was assessed a decade ago, but the number of valid species has since increased by 60%, and 17% of the taxa from that study have since undergone taxonomic revision. Here, we assess how 10 years of additional research has changed our outlook on the group's fossil record. We quantified the completeness of all 307 sauropodomorph species currently considered valid using the skeletal completeness metric, which calculates the proportion of a complete skeleton preserved for each taxon. Taxonomic and stratigraphic age revisions, rather than new species, are the drivers of the most significant differences between the current results and those of the previous assessment. No statistical differences appeared when we use our new dataset to generate temporal completeness curves based solely on taxa known in 2009 or 1999. We now observe a severe drop in mean completeness values across the Jurassic–Cretaceous boundary that never recovers to pre‐Cretaceous levels. Explaining this pattern is difficult, as we find no convincing evidence that it is related to environmental preferences or body size changes. Instead, it might result from: (1) reduction of terrestrial fossil preservation space due to sea level rise; (2) ecological specificities and relatively high diagnosability of Cretaceous species; and/or (3) increased sampling of newly explored sites with many previously unknown taxa. Revisiting patterns in this manner allows us to test the longevity of conclusions made in previous quantitative studies

Decoupling of morphological disparity and taxic diversity during the adaptive radiation of anomodont therapsids

Adaptive radiations are central to macroevolutionary theory. Whether triggered by acquisition of new traits or ecological opportunities arising from mass extinctions, it is debated whether adaptive radiations are marked by initial expansion of taxic diversity or of morphological disparity (the range of anatomical form). If a group rediversifies following a mass extinction, it is said to have passed through a macroevolutionary bottleneck, and the loss of taxic or phylogenetic diversity may limit the amount of morphological novelty that it can subsequently generate. Anomodont therapsids, a diverse clade of Permian and Triassic herbivorous tetrapods, passed through a bottleneck during the end-Permian mass extinction. Their taxic diversity increased during the Permian, declined significantly at the Permo–Triassic boundary and rebounded during the Middle Triassic before the clade's final extinction at the end of the Triassic. By sharp contrast, disparity declined steadily during most of anomodont history. Our results highlight three main aspects of adaptive radiations: (i) diversity and disparity are generally decoupled; (ii) models of radiations following mass extinctions may differ from those triggered by other causes (e.g. trait acquisition); and (iii) the bottleneck caused by a mass extinction means that a clade can emerge lacking its original potential for generating morphological variety

Best value and workplace partnership in local government

Purpose – This paper explores employee experiences concerning job security/insecurity, workload, job satisfaction and employee involvement in the aftermath of Best Value reviews in a local authority. Design/methodology/approach – Using a mix of quantitative and qualitative data collection techniques employees’ experiences of Best Value reviews in a local authority are compared and contrasted with council staff employed elsewhere in the authority to establish the extent to which workplace partnership principles have taken hold under a Best Value regime. Findings – Little evidence of positive outcomes was found from partnership at work under a Best Value regime. The constraints imposed by central government, under which managers in the public sector operate, contributed significantly to partnership at work remaining little more than a hollow shell. Originality/value – This paper provides a recent in-depth case study of the experience of workplace partnership, which was developed not discrete from but as part of the Best Value modernisation programme in a local authority

Partnership with and without trade unions in the UK financial services: filling or fuelling the representation gap?

Partnership theory proposes that an appropriate integration of direct and indirect employee participation mutually benefits workers and company. This study explores the putative employee voice gains and the risks for union effectiveness by comparing employees' evaluation of partnership practices at two financial service companies with nonunion and union employee representation respectively