Extinct Relatives Research Articles

Phylogeny of Merlin\u2019s grass (Isoetaceae): revealing an \u201cAmborella syndrome\u201d and the importance of geographic distribution for understanding current and historical diversity

BackgroundMerlin’s grass (Isoetes, Isoetaceae, Lycopsida), is the extant remnant of the isoetalean wood-producing lycopsids that originated during the Paleozoic, possibly in aquatic or boggy habitats. Modern day species are aquatic, semi-aquatic or terrestrial and occur almost worldwide. They display little morphological variation; the lobed corm has helically arranged leaves with internal air channels and basal sporangia. Genetic variation has also proven limited, which has hampered phylogenetic inference. We investigate evolutionary relationships in Isoetes, using molecular data and an extended sample of species compared to previous work, adding species that have never before been included in a phylogenetic study.ResultsOur results reveal an unexpected discovery of an “Amborella syndrome” in Isoetaceae: a single poorly known species is sister to the remaining family. The species, Isoetes wormaldii, is a rare endemic to the Eastern Cape of South Africa. Its leaves are flattened with a rounded point, which sharply contrasts with the awl-shaped leaves of most other species of Isoetes. The remaining species of Isoetes are resolved in five major clades, also indicated in previous work. While the phylogeny shows geographic structure, the patterns are complex. For example, tropical-southern African species occur in at least five clades, and Indian, Australian and Mediterranean species in at least three clades each.ConclusionThe evolutionary and biogeographical history of Isoetes is not easily explained, and may conceivably include ample extinction and a mixture of ancient and more recent processes. Previously shown difficulties with node age estimation increase the problem. The here demonstrated sister-relationship between the phylogenetically, morphologically and genetically distinct Isoetes wormaldii and the remaining family appears to bridge the morphological gap between Isoetes and its extinct relatives, although further studies are needed. Moreover, it shortens the branch length to its living sister genus Selaginella, and may enhance node age estimation in future studies. Isoetes wormaldii is critically endangered, known only from one (to a few) minor populations. Immediate actions need to be taken if we want to prevent this unique species from going extinct.

Size and shape regional differentiation during the development of the spine in the nine‐banded armadillo (Dasypus novemcinctus)

Xenarthrans (armadillos, anteaters, sloths, and their extinct relatives) are unique among mammals in displaying a distinctive specialization of the posterior trunk vertebrae-supernumerary vertebral xenarthrous articulations. This study seeks to understand how xenarthry develops through ontogeny and if it may be constrained to appear within pre-existing vertebral regions. Using three-dimensionalgeometric morphometrics on the neural arches of vertebrae, we explore phenotypic, allometric, and disparity patterns of the different axial morphotypes during the ontogeny of nine-banded armadillos. Shape-based regionalization analyses showed that the adult thoracolumbar column is divided into three regions according to the presence or absence of ribs and the presence or absence of xenarthrous articulations. A three-region division was retrieved in almost all specimens through development, although younger stages (e.g., fetuses, neonates) have more region boundary variability. In size-based regionalization analyses, thoracolumbar vertebrae are separated into two regions: a prediaphragmatic, prexenarthrous region, and a postdiaphragmatic xenarthrous region. We show that posterior thoracic vertebrae grow at a slower rate, while anterior thoracics and lumbars grow at a faster rate relatively, with rates decreasing anteroposteriorly in the former and increasing anteroposteriorly in the latter. We propose that different proportions between vertebrae and vertebral regions might result from differences in growth pattern and timing of ossification.

New Middle Eocene proboscidean from Togo illuminates the early evolution of the elephantiform-like dental pattern.

Africa has played a pivotal role in the evolution of early proboscideans (elephants and their extinct relatives), yet vast temporal and geographical zones remain uncharted on the continent. A long hiatus encompassing most of the Eocene (Ypresian to the Early Priabonian, around 13 Myr timespan) considerably hampers our understanding of the early evolutionary history of the group. It is notably the case with the origin of its most successful members, the Elephantiformes, i.e. all elephant-like proboscideans most closely related to modern elephants. Here, we describe a proboscidean lower molar discovered in Lutetian phosphate deposits from Togo, and name a new genus and species, Dagbatitherium tassyi. We show that Dagbatitherium displays several elephantiform dental characteristics such as a three-layered Schmelzmuster, the presence of a mesoconid, transversely enlarged buccal cusps and the individualization of a third lophid closely appressed to a minute distal cingulid. Dagbatitherium represents a stem Elephantiformes, pushing back the origin of the group by about 10 Myr, i.e. a third of its currently known evolutionary history. More importantly, Dagbatitherium potentially unlocks the puzzle of the origin of the unique elephantiform tooth crown organization by bridging a critical temporal and morphological gap between early bunodont incipiently bilophodont proboscidean taxa and more derived elephantiforms.

Giant Mesozoic coelacanths (Osteichthyes, Actinistia) reveal high body size disparity decoupled from taxic diversity

The positive correlation between speciation rates and morphological evolution expressed by body size is a macroevolutionary trait of vertebrates. Although taxic diversification and morphological evolution are slow in coelacanths, their fossil record indicates that large and small species coexisted, which calls into question the link between morphological and body size disparities. Here, we describe and reassess fossils of giant coelacanths. Two genera reached up to 5 m long, placing them among the ten largest bony fish that ever lived. The disparity in body size adjusted to taxic diversity is much greater in coelacanths than in ray-finned fishes. Previous studies have shown that rates of speciation and rates of morphological evolution are overall low in this group, and our results indicate that these parameters are decoupled from the disparity in body size in coelacanths. Genomic and physiological characteristics of the extant Latimeria may reflect how the extinct relatives grew to such a large size. These characteristics highlight new evolutionary traits specific to these “living fossils”.

Ancient DNA from bulk bone reveals past genetic diversity of vertebrate fauna on Kangaroo Island, Australia

It is indisputable that much of Australia's modern biodiversity decline was triggered by European settlement. However, the driver(s) of pre-European extinctions and extirpations are more challenging to identify, particularly on islands where animals are faced with the additional pressures imposed by isolation. Kangaroo Island, South Australia, has been identified as a potential haven for the reintroduction of endangered animals, but the genetic relationship between reintroduction candidates on the mainland and their extinct relatives on Kangaroo Island is poorly understood. Here, we present a late Pleistocene to mid-Holocene genetic record from Kangaroo Island based on bulk bone metabarcoding of two thousand bone fragments excavated from Kelly Hill Cave. We detect 33 species of bird, reptile, fish and mammal and report additional intra-specific genetic diversity in Kangaroo Island's now extinct population of spotted-tailed quoll. Furthermore, we provide genetic evidence that the Eastern grey kangaroo formerly inhabited Kangaroo Island. Taken together, these data establish a more complete baseline of local biodiversity against which reintroduction programmes and bushfire recovery measures can be evaluated, which is more important than ever in light of the recent fires that devastated much of Kangaroo Island's flora and fauna.

Brain organoids with Neanderthal genes

Human Evolution The genomes of Neanderthals and modern humans are overall very similar. To understand the impact of genetic variants that are specific to modern humans, Trujillo et al. performed a genome-wide analysis to identify 61 coding variants in protein-coding genes. Identifying the gene encoding the RNA-binding protein NOVA1 as a top candidate for functional analyses, they introduced the archaic gene variant into human pluripotent stem cells and generated brain organoids. These organoids showed alterations in gene expression and splicing as well as morphology and synaptogenesis, suggesting that this method could be used to explore other genetic changes that underlie the phenotypic traits separating our species from extinct relatives. Science , this issue p. [eaax2537][1] [1]: /lookup/doi/10.1126/science.aax2537

Low elbow mobility indicates unique forelimb posture and function in a giant extinct marsupial.

Joint mobility is a key factor in determining the functional capacity of tetrapod limbs, and is important in palaeobiological reconstructions of extinct animals. Recent advances have been made in quantifying osteological joint mobility using virtual computational methods; however, these approaches generally focus on the proximal limb joints and have seldom been applied to fossil mammals. Palorchestes azael is an enigmatic, extinct ~1000kg marsupial with no close living relatives, whose functional ecology within Australian Pleistocene environments is poorly understood. Most intriguing is its flattened elbow morphology, which has long been assumed to indicate very low mobility at this important joint. Here, we tested elbow mobility via virtual range of motion (ROM) mapping and helical axis analysis, to quantitatively explore the limits of Palorchestes' elbow movement and compare this with their living and extinct relatives, as well as extant mammals that may represent functional analogues. We find that Palorchestes had the lowest elbow mobility among mammals sampled, even when afforded joint translations in addition to rotational degrees of freedom. This indicates that Palorchestes was limited to crouched forelimb postures, something highly unusual for mammals of this size. Coupled flexion and abduction created a skewed primary axis of movement at the elbow, suggesting an abducted forelimb posture and humeral rotation gait that is not found among marsupials and unlike that seen in any large mammals alive today. This work introduces new quantitative methods and demonstrates the utility of comparative ROM mapping approaches, highlighting that Palorchestes' forelimb function was unlike its contemporaneous relatives and appears to lack clear functional analogues among living mammals.

A symmoriiform from the Late Devonian of Morocco demonstrates a derived jaw function in ancient chondrichthyans

The Palaeozoic record of chondrichthyans (sharks, rays, chimaeras, extinct relatives) and thus our knowledge of their anatomy and functional morphology is poor because of their predominantly cartilaginous skeletons. Here, we report a previously undescribed symmoriiform shark, Ferromirum oukherbouchi, from the Late Devonian of the Anti-Atlas. Computed tomography scanning reveals the undeformed shape of the jaws and hyoid arch, which are of a kind often used to represent primitive conditions for jawed vertebrates. Of critical importance, these closely fitting cartilages preclude the repeatedly hypothesized presence of a complete gill between mandibular and hyoid arches. We show that the jaw articulation is specialized and drives mandibular rotation outward when the mouth opens, and inward upon closure. The resultant eversion and inversion of the lower dentition presents a greater number of teeth to prey through the bite-cycle. This suggests an increased functional and ecomorphological disparity among chondrichthyans preceding and surviving the end-Devonian extinctions.

Vertebral pneumaticity is correlated with serial variation in vertebral shape in storks.

Birds and their ornithodiran ancestors are unique among vertebrates in exhibiting air-filled sinuses in their postcranial bones, a phenomenon called postcranial skeletal pneumaticity. The factors that account for serial and interspecific variation in postcranial skeletal pneumaticity are poorly understood, although body size, ecology, and bone biomechanics have all been implicated as influencing the extent to which pneumatizing epithelia invade the skeleton and induce bone resorption. Here, I use high-resolution computed-tomography to holistically quantify vertebral pneumaticity in members of the neognath family Ciconiidae (storks), with pneumaticity measured as the relative volume of internal air space. These data are used to describe serial variation in extent of pneumaticity and to assess whether and how pneumaticity varies with the size and shape of a vertebra. Pneumaticity increases dramatically from the middle of the neck onwards, contrary to previous predictions that cervical pneumaticity should decrease toward the thorax to maintain structural integrity as the mass and bending moments of the neck increase. Although the largest vertebrae sampled are also the most pneumatic, vertebral size cannot on its own account for serial or interspecific variation in extent of pneumaticity. Vertebral shape, as quantified by three-dimensional geometric morphometrics, is found to be significantly correlated with extent of pneumaticity, with elongate vertebrae being less pneumatic than craniocaudally short and dorsoventrally tall vertebrae. Considered together, the results of this study are consistent with the hypothesis that shape- and position-specific biomechanics influence the amount of bone loss that can be safely tolerated. These results have potentially important implications for the evolution of vertebral morphology in birds and their extinct relatives.

What can lycophytes teach us about plant evolution and development? Modern perspectives on an ancient lineage.

All Evo-Devo studies rely on representative sampling across the tree of interest to elucidate evolutionary trajectories through time. In land plants, genetic resources are well established in model species representing lineages including bryophytes (mosses, liverworts, and hornworts), monilophytes (ferns and allies), and seed plants (gymnosperms and flowering plants), but few resources are available for lycophytes (club mosses, spike mosses, and quillworts). Living lycophytes are a sister group to the euphyllophytes (the fern and seed plant clade), and have retained several ancestral morphological traits despite divergence from a common ancestor of vascular plants around 420 million years ago. This sister relationship offers a unique opportunity to study the conservation of traits such as sporophyte branching, vasculature, and indeterminacy, as well as the convergent evolution of traits such as leaves and roots which have evolved independently in each vascular plant lineage. To elucidate the evolution of vascular development and leaf formation, molecular studies using RNA Seq, quantitative reverse transcription polymerase chain reaction, in situ hybridisation and phylogenetics have revealed the diversification and expression patterns of KNOX, ARP, HD-ZIP, KANADI, and WOX gene families in lycophytes. However, the molecular basis of further trait evolution is not known. Here we describe morphological traits of living lycophytes and their extinct relatives, consider the molecular underpinnings of trait evolution and discuss future research required in lycophytes to understand the key evolutionary innovations enabling the growth and development of all vascular plants.

Intra- and interspecific variation in tooth morphology of procyon cancrivorus and p. Lotor (carnivora, procyonidae), and its bearing on the Taxonomy of fossil south American procyonids

The family Procyonidae (raccoons, coatis, olingos, ringtails, kinkajous, and their extinct relatives) consists of six extant genera and is restricted to North and South America. Currently recognized fossil species suggest that procyonid diversity was previously much greater, including six extinct genera throughout South America. However, it is unusual that so many confamilial taxa are represented in a relatively brief span of time and restricted geographic region, and, considering that six of ten are based on badly preserved specimens, often fragments of bone with worn teeth, the validity of many of these taxa is suspect. As a step towards reevaluating past procyonid diversity in South America, we sought to identify the degree of intra- and interspecific variation in six molariform teeth of extant Procyon , particularly to identify which teeth are potentially most useful for identifying fossil procyonids. The six molariform cheek teeth analyzed consistently yielded smaller intra- than interspecific variation, permitting high accuracy of taxonomic classification. However, this accuracy varied by tooth, and the upper and lower first molars proved to be the most reliable. Thus, these particular teeth should be preferred, if available, as bases for recognizing extinct species of procyonids or reevaluating currently recognized extinct species, as a means to prevent nomina dubia .

The craniomandibular anatomy of the early archosauriform Euparkeria capensis and the dawn of the archosaur skull.

Archosauria (birds, crocodilians and their extinct relatives) form a major part of terrestrial ecosystems today, with over 10 000 living species, and came to dominate the land for most of the Mesozoic (over 150 Myr) after radiating following the Permian–Triassic extinction. The archosaur skull has been essential to this diversification, itself diversified into myriad forms. The archosauriform Euparkeria capensis from the Middle Triassic (Anisian) of South Africa has been of great interest since its initial description in 1913, because its anatomy shed light on the origins and early evolution of crown Archosauria and potentially approached that of the archosaur common ancestor. Euparkeria has been widely used as an outgroup in phylogenetic analyses and when investigating patterns of trait evolution among archosaurs. Although described monographically in 1965, subsequent years have seen great advances in the understanding of early archosaurs and in imaging techniques. Here, the cranium and mandible of Euparkeria are fully redescribed and documented using all fossil material and computed tomographic data. Details previously unclear are fully described, including vomerine dentition, the epiptergoid, number of premaxillary teeth and palatal arrangement. A new diagnosis and cranial and braincase reconstruction is provided, and an anatomical network analysis is performed on the skull of Euparkeria and compared with other amniotes. The modular composition of the cranium suggests a flexible skull well adapted to feeding on agile food, but with a clear tendency towards more carnivorous behaviour, placing the taxon at the interface between ancestral diapsid and crown archosaur ecomorphology, corresponding to increases in brain size, visual sensitivity, upright locomotion and metabolism around this point in archosauriform evolution. The skull of Euparkeria epitomizes a major evolutionary transition, and places crown archosaur morphology in an evolutionary context.

Modern human changes in regulatory regions implicated in cortical development

BackgroundRecent paleogenomic studies have highlighted a very small set of proteins carrying modern human-specific missense changes in comparison to our closest extinct relatives. Despite being frequently alluded to as highly relevant, species-specific differences in regulatory regions remain understudied. Here, we integrate data from paleogenomics, chromatin modification and physical interaction, and single-cell gene expression of neural progenitor cells to identify derived regulatory changes in the modern human lineage in comparison to Neanderthals/Denisovans. We report a set of genes whose enhancers and/or promoters harbor modern human single nucleotide changes and are active at early stages of cortical development.ResultsWe identified 212 genes controlled by regulatory regions harboring modern human changes where Neanderthals/Denisovans carry the ancestral allele. These regulatory regions significantly overlap with putative modern human positively-selected regions and schizophrenia-related genetic loci. Among the 212 genes, we identified a substantial proportion of genes related to transcriptional regulation and, specifically, an enrichment for the SETD1A histone methyltransferase complex, known to regulate WNT signaling for the generation and proliferation of intermediate progenitor cells.ConclusionsThis study complements previous research focused on protein-coding changes distinguishing our species from Neanderthals/Denisovans and highlights chromatin regulation as a functional category so far overlooked in modern human evolution studies. We present a set of candidates that will help to illuminate the investigation of modern human-specific ontogenetic trajectories.

Incorporating Trace Fossils when Exploring the Interplay of Form and Function in the Crocodyliform Feeding Apparatus

Many crocodylians, both living and extinct, fill the semi‐aquatic ambush predator niche, and their prey preferences roughly can be predicted based on a combination of snout shape, tooth morphology, and body size. Among extant crocodylians, slender‐snouted taxa such as Gavialis gangeticus and Tomistoma schlegelii generally prefer smaller‐bodied prey, whereas the broader‐snouted taxa divide into generalists like Crocodylus niloticus and macro‐generalists like Alligator mississippiensis, who will eat a diversity of prey items including animals that are as big or even bigger than themselves. Large‐bodied members of these generalist ecomorphotypes generate the highest bite forces observed in modern taxa and are often apex predators in their ecosystems. The availability of a robust phylogenetic hypothesis for crocodylians makes them an excellent clade with which to explore these types of ecomorphological hypotheses, which link form and function in the feeding apparatus within an evolutionary framework. The resulting synthesis of extant and extinct datasets has provided both exciting new insights, but also some complications. The addition of extinct taxa into studies of morphological disparity, both within the crown group and deeper in the crocodyliform evolutionary tree, introduce a diverse array of snout shapes and associated niches that are not represented in the surviving members of the clade (e.g. fully marine, fully terrestrial, durophagous, omnivorous, and even possibly herbivorous forms). Though hypotheses regarding diet in these diverse groups are based on craniodental morphology, indirect evidence, in the form of trace fossils, provides an independent means of testing those predictions. Gut contents are extremely rare in this clade because of the highly destructive digestion of crocodyliforms, and coprolites, though comparatively common, contain remains that are so degraded as to be largely unidentifiable and uninformative. Isotopic and dental microwear studies can provide broad patterns of diet. However, for detailed, direct evidence of crocodyliform feeding, bite marks are a particularly useful source of direct evidence with which to explore patterns of diet in this clade. A recent, concentrated effort to characterize modern crocodylian bite marks has resulted in the identification of novel patterns and diagnostic traces associated with feeding behavior in that clade. These trace fossils provide critical insights into diet, feeding strategy, and even intraspecific competition in crocodyliforms and their extinct relatives. By integrating these ichnological data with traditional dietary hypotheses based on morphology and phylogeny of extinct groups, a more complete, and oftentimes more complex and realistic, concept of past ecosystems and trophic structures can be explored.Support or Funding InformationFunding provided by the National Science Foundation, the Leakey Foundation, and National Geographic.

Forensically relevant phylogeographic evaluation of mitogenome variation in the Basque Country

The Basque Country has been the focus of population (genetic) and evolutionary studies for decades, as it represents an interesting evolutionary feature: it is the only European country where a non-Indo-European language is still spoken today and, for which there are no known living or extinct relatives. Early studies that were based on anatomical and serological methods, along with subsequent molecular genetic investigations, contain controversial interpretations of their data. Additionally, the analysis of mitochondrial DNA, which is maternally inherited and thus suitable for the examination of the maternal phylogeny of the population, was the focus of some studies. Early mtDNA studies were however restricted to the information provided by the control region or its hypervariable segments only. These are known to harbour little phylogenetic information, particularly for haplogroup H that is dominant in Westeurasian populations including the Basques. Later studies analysed complete mitogenome sequences. Their information content is however limited, either because the number of samples was low, or because these studies only considered particular haplogroups. In this study we present the full mitogenome sequences of 178 autochthonous Basque individuals that were carefully selected based on their familial descent and discuss the observed phylogenetic signals in the light of earlier published findings. We confirm the presence of Basque-specific mtDNA lineages and extend the knowledge of these lineages by providing data on their distribution in comparison to other Basque and non-Basque populations. This dataset improves our understanding of the Basque mtDNA phylogeny and serves as a high-quality dataset that is provided via EMPOP for forensic genetic purposes.

Giant extinct caiman breaks constraint on the axial skeleton of extant crocodylians.

The number of precaudal vertebrae in all extant crocodylians is remarkably conservative, with nine cervicals, 15 dorsals and two sacrals, a pattern present also in their closest extinct relatives. The consistent vertebral count indicates a tight control of axial patterning by Hox genes during development. Here we report on a deviation from this pattern based on an associated skeleton of the giant caimanine Purussaurus, a member of crown Crocodylia, and several other specimens from the Neogene of the northern neotropics. P. mirandai is the first crown-crocodylian to have three sacrals, two true sacral vertebrae and one non-pathological and functional dorsosacral, to articulate with the ilium (pelvis). The giant body size of this caiman relates to locomotory and postural changes. The iliosacral configuration, a more vertically oriented pectoral girdle, and low torsion of the femoral head relative to the condyles are hypothesized specializations for more upright limb orientation or weight support.

Reconstructing the archosaur radiation using a Middle Triassic archosauriform tooth assemblage from Tanzania.

Following the Permo–Triassic mass extinction, Archosauriformes—the clade that includes crocodylians, birds, and their extinct relatives outside crown Archosauria—rapidly diversified into many distinct lineages, became distributed globally, and, by the Late Triassic, filled a wide array of resource zones. Current scenarios of archosauriform evolution are ambiguous with respect to whether their taxonomic diversification in the Early–Middle Triassic coincided with the initial evolution of dietary specializations that were present by the Late Triassic or if their ecological disparity arose sometime after lineage diversification. Late Triassic archosauriform dietary specialization is recorded by morphological divergence from the plesiomorphic archosauriform tooth condition (laterally-compressed crowns with serrated carinae and a generally triangular lateral profile). Unfortunately, the roots of this diversification are poorly documented, with few known Early­–Middle Triassic tooth assemblages, limiting characterizations of morphological diversity during this critical, early period in archosaur evolution. Recent fieldwork (2007–2017) in the Middle Triassic Manda Beds of the Ruhuhu Basin, Tanzania, recovered a tooth assemblage that provides a window into this poorly sampled interval. To investigate the taxonomic composition of that collection, we built a dataset of continuous quantitative and discrete morphological characters based on in situ teeth of known taxonomic status (e.g., Nundasuchus, Parringtonia: N = 65) and a sample of isolated teeth (N = 31). Using crown heights from known taxa to predict tooth base ratio (= base length/width), we created a quantitative morphospace for the tooth assemblage. The majority of isolated, unassigned teeth fall within a region of morphospace shared by several taxa from the Manda Beds (e.g., Nundasuchus, Parringtonia); two isolated teeth fall exclusively within a “Pallisteria” morphospace. A non-metric multidimensional scaling ordination (N = 67) of 11 binary characters reduced overlap between species. The majority of the isolated teeth from the Manda assemblage fall within the Nundasuchus morphospace. This indicates these teeth are plesiomorphic for archosauriforms as Nundasuchus exhibits the predicted plesiomorphic condition of archosauriform teeth. Our model shows that the conservative tooth morphologies of archosauriforms can be differentiated and assigned to species and/or genus, rendering the model useful for identifying isolated teeth. The large overlap in tooth shape among the species present and their overall similarity indicates that dietary specialization lagged behind species diversification in archosauriforms from the Manda Beds, a pattern predicted by Simpson’s “adaptive zones” model. Although applied to a single geographic region, our methods offer a promising means to reconstruct ecological radiations and are readily transferable across a broad range of vertebrate taxa throughout Earth history.

Osteology of the archosauromorph Teyujagua paradoxa and the early evolution of the archosauriform skull

Abstract Archosauriformes are a major group of fossil and living reptiles that include the crown group Archosauria (birds, crocodilians and their extinct relatives) and closely related taxa. Archosauriformes are characterized by a highly diagnostic skull architecture, which is linked to the predatory habits of their early representatives, and the development of extensive cranial pneumaticity associated with the nasal capsule. The evolution of the archosauriform skull from the more plesiomorphic configuration present ancestrally in the broader clade Archosauromorpha was, until recently, elusive. This began to change with the discovery and description of Teyujagua paradoxa, an early archosauromorph from the Lower Triassic Sanga do Cabral Formation of Brazil. Here, we provide a detailed osteological description of the holotype and, thus far, only known specimen of T. paradoxa. In addition to providing new details of the anatomy of T. paradoxa, our study also reveals an early development of skull pneumaticity prior to the emergence of the antorbital fenestra. We use these new data to discuss the evolution of antorbital openings within Archosauriformes. Reappraisal of the phylogenetic position of T. paradoxa supports previous hypotheses of a close relationship with Archosauriformes. The data presented here provide new insights into character evolution during the origin of the archosauriform skull.

Crocodylomorph cranial shape evolution and its relationship with body size and ecology.

Crocodylomorpha, which includes living crocodylians and their extinct relatives, has a rich fossil record, extending back for more than 200 million years. Unlike modern semi-aquatic crocodylians, extinct crocodylomorphs exhibited more varied lifestyles, ranging from marine to fully terrestrial forms. This ecological diversity was mirrored by a remarkable morphological disparity, particularly in terms of cranial morphology, which seems to be closely associated with ecological roles in the group. Here, I use geometric morphometrics to comprehensively investigate cranial shape variation and disparity in Crocodylomorpha. I quantitatively assess the relationship between cranial shape and ecology (i.e. terrestrial, aquatic, and semi-aquatic lifestyles), as well as possible allometric shape changes. I also characterize patterns of cranial shape evolution and identify regime shifts. I found a strong link between shape and size, and a significant influence of ecology on the observed shape variation. Terrestrial taxa, particularly notosuchians, have significantly higher disparity, and shifts to more longirostrine regimes are associated with large-bodied aquatic or semi-aquatic species. This demonstrates an intricate relationship between cranial shape, body size and lifestyle in crocodylomorph evolutionary history. Additionally, disparity-through-time analyses were highly sensitive to different phylogenetic hypotheses, suggesting the description of overall patterns among distinct trees. For crocodylomorphs, most results agree in an early peak during the Early Jurassic and another in the middle of the Cretaceous, followed by nearly continuous decline until today. Since only crown-group members survived through the Cenozoic, this decrease in disparity was likely the result of habitat loss, which narrowed down the range of crocodylomorph lifestyles.

Variation in appendages in early Cambrian bradoriids reveals a wide range of body plans in stem-euarthropods

Traditionally, the origin and evolution of modern arthropod body plans has been revealed through increasing levels of appendage specialisation exhibited by Cambrian euarthropods. Here we show significant variation in limb morphologies and patterns of limb-tagmosis among three early Cambrian arthropod species conventionally assigned to the Bradoriida. These arthropods are recovered as a monophyletic stem-euarthropod group (and sister taxon to crown-group euarthropods, i.e. Chelicerata, Mandibulata and their extinct relatives), thus implying a radiation of stem-euarthropods where trends towards increasing appendage specialisation were explored convergently with other euarthropod groups. The alternative solution, where bradoriids are polyphyletic, representing several independent origins of a small, bivalved body plan in lineages from diverse regions of the euarthropod and mandibulate stems, is only marginally less parsimonious. The new data reveal a previously unknown disparity of body plans in stem-euarthropods and both solutions support remarkable evolutionary convergence, either of fundamental body plans or appendage specialization patterns.