Tetrapod Clades Research Articles

Cranial Musculoskeletal Description of Black-Throated Finch (Aves: Passeriformes: Estrildidae) with DiceCT.

Synopsis Dietary requirements and acquisition strategies change throughout ontogeny across various clades of tetrapods, including birds. For example, birds hatch with combinations of various behavioral, physiological, and morphological factors that place them on an altricial–precocial spectrum. Passeriformes (=songbirds) in particular, a family constituting approximately more than half of known bird species, displays the most drastic difference between hatchling and adults in each of these aspects of their feeding biology. How the shift in dietary resource acquisition is managed during ontogeny alongside its relationship to the morphology of the feeding apparatus has been largely understudied within birds. Such efforts have been hampered partly due to the small size of many birds and the diminutive jaw musculature they employ. In this study, we used standard and diffusible iodine-based contrast-enhanced computed tomography in conjunction with digital dissection to quantify and describe the cranial musculature of the Black-throated Finch (Poephila cincta) at fledgling and adult stages. Our results reveal that in both the fledgling and the adult, cranial musculature shows clear and complex partitioning in the Musculus adductor mandibulae externus that is consistent with other families within Passeriformes. We quantified jaw-muscle sizes and found that the adult showed a decrease in muscle mass in comparison to the fledgling individual. We propose that this could be the result of low sample size or a physiological effect of parental care in Passeriformes. Our study shows that high-resolution visualization techniques are informative at revealing morphological discrepancies for studies that involve small specimens such as Passeriformes especially with careful specimen selection criteria.

Computed tomographic analysis of the cranium of the early Permian recumbirostran ‘microsaur’ Euryodus dalyae reveals new details of the braincase and mandible

AbstractRecumbirostran ‘microsaurs’ are a clade of Palaeozoic tetrapods that possess numerous morphological adaptations for fossorial ecologies. Re‐study of many ‘microsaurs’ using tomographic methods has provided substantial new data on poorly known anatomy that informs their debated phylogenetic position. Recent studies have identified suites of features among recumbirostrans that place the group within crown Amniota, contrary to hypothesized positions on the amniote stem or the lissamphibian stem. Herein we describe the cranial anatomy of the early Permian gymnarthrid Euryodus dalyae through tomographic analysis of the holotype from South Grandfield, Oklahoma and new specimens from karst deposits near Richards Spur. The braincase of E. dalyae is composed of well‐ossified pleurosphenoids, orbitosphenoids that brace against the skull roof, and unpaired median ossifications. The otic capsules are well‐ossified, and the occiput is unconsolidated. Analysis of the mandibles, typically obscured in articulated specimens, reveals a second tooth row on the dentary, a feature previously unknown in ‘microsaurs’ that is reminiscent of the condition of the co‐occurring captorhinid Captorhinus aguti. The Richards Spur specimens share many of these features, including the second tooth row, but the neurocranium of the scanned specimen (OMNH 53519) differs from that of the holotype of E. dalyae (e.g. absence of unpaired median ossifications), and these specimens are referred to Euryodus sp. These data add to the growing neurocranial dataset of ‘microsaurs’, which is essential for iterative reevaluation of early tetrapod phylogeny. This discovery of multiple tooth rows in ‘microsaurs’ provides further support for the hypothesized close relationships between ‘microsaurs’ and reptiles.

A redescription of the late Carboniferous trematopid Actiobates peabodyi from Garnett, Kansas.

Dissorophoids are a diverse clade of predominantly Permo-Carboniferous temnospondyls with a wide geographic distribution and broad ecological diversity. Each of the various dissorophoid clades first appears in the late Carboniferous, but their records are relatively sparse and fragmentary compared to those of the early Permian when dissorophoids reach their peak diversity and distribution, particularly in terrestrial environments where they are by far the most taxonomically diverse clade of non-amniote tetrapods. This provides an impetus for further study of the late Carboniferous terrestrial dissorophoids in order to contextualize the early stages in the clade's radiation into terrestrial ecosystems. Here we present a redescription of the late Carboniferous trematopid Actiobates peabodyi from Kansas, USA, which is represented by a nearly complete skeleton and which represents the earliest occurrence of trematopids in North America. Only the skull was previously described, and the taxon has been largely overlooked in the context of early terrestrial dissorophoid evolution. Here, we provide an updated cranial description, the first postcranial description, and a discussion of the position of A. peabodyi in the context of olsoniform evolution. Our most significant finding is the characterization of postcranial anatomy that is highly similar to that of later trematopid taxa. This high degree of conservatism indicates that the earliest trematopids were already well adapted for terrestrial environments, and post-Carboniferous radiations of olsoniforms may be attributed to an expansion of the dryland terrestrial environments in which these taxa already thrived, rather than to novel acquisition of adaptive features later in the clade's evolution.

Inspiring song: The role of respiratory circuitry in the evolution of vertebrate vocal behavior.

Vocalization is a common means of communication across vertebrates, but the evolutionary origins of the neural circuits controlling these behaviors are not clear. Peripheral mechanisms of sound production vary widely: fish produce sounds with a swimbladder or pectoral fins; amphibians, reptiles, and mammalians vocalize using a larynx; birds vocalize with a syrinx. Despite the diversity of vocal effectors across taxa, there are many similarities in the neural circuits underlying the control of these organs. Do similarities in vocal circuit structure and function indicate that vocal behaviors first arose in a single common ancestor, or have similar neural circuits arisen independently multiple times during evolution? In this review, we describe the hindbrain circuits that are involved in vocal production across vertebrates. Given that vocalization depends on respiration in most tetrapods, it is not surprising that vocal and respiratory hindbrain circuits across distantly related species are anatomically intermingled and functionally linked. Such vocal-respiratory circuit integration supports the hypothesis that vocal evolution involved the expansion and functional diversification of breathing circuits. Recent phylogenetic analyses, however, suggest vocal behaviors arose independently in all major tetrapod clades, indicating that similarities in vocal control circuits are the result of repeated co-options of respiratory circuits in each lineage. It is currently unknown whether vocal circuits across taxa are made up of homologous neurons, or whether vocal neurons in each lineage arose from developmentally and evolutionarily distinct progenitors. Integrative comparative studies of vocal neurons across brain regions and taxa will be required to distinguish between these two scenarios.

Divergent evolutionary morphology of the axial skeleton as a potential key innovation in modern cetaceans.

Cetaceans represent the most diverse clade of extant marine tetrapods. Although the restructuring of oceans could have contributed to their diversity, other factors might also be involved. Similar to ichthyosaurs and sharks, variation of morphological traits could have promoted the colonization of new ecological niches and supported their diversification. By combining morphological data describing the axial skeleton of 73 cetacean species with phylogenetic comparative methods, we demonstrate that the vertebral morphology of cetaceans is associated with their habitat. All riverine and coastal species possess a small body size, lengthened vertebrae and a low vertebral count compared with open ocean species. Extant cetaceans have followed two distinct evolutionary pathways relative to their ecology. Whereas most offshore species such as baleen whales evolved towards an increased body size while retaining a low vertebral count, small oceanic dolphins underwent deep modifications of their axial skeleton with an extremely high number of short vertebrae. Our comparative analyses provide evidence these vertebral modifications have potentially operated as key innovations. These novelties contributed to their explosive radiation, resulting in an efficient swimming style that provides energetic advantages to small-sized species.

Endocranial morphology of the Brazilian Permian dicynodont Rastodon procurvidens (Therapsida: Anomodontia).

Dicynodontia is a major clade of terrestrial tetrapods that greatly diversified during the Permian and Triassic periods, reaching a worldwide distribution. In this study, the endocranial cavity of the Brazilian Permian dicynodont Rastodon procurvidens is described based on a digital endocast obtained using digital imaging (X-ray computed tomography) and 3D modeling. It was possible to reconstruct the brain, olfactory bulbs, inner ear, some neurovascular canals, cranial nerves, the nasal cavity, and the maxillary recesses. The endocast of R. procurvidens preserves a typical plesiomorphic morphology of non-mammaliaform therapsids, being predominantly tubular and displaying a relatively short and robust hindbrain. Encephalization quotients (EQs) were calculated for R. procurvidens, resulting in EQs of 0.09±0.03 and 0.13±0.05 (Jerison's EQ and Manger's EQ, respectively). Finally, some biological implications of the endocast morphology were inferred for R. procurvidens. Its inner ear is especially small, and its orientation implies a slightly downturned head posture in life. Furthermore, the presence of uncompressed maxillary recesses in R. procurvidens indicates a correlation between the enlargement of the recesses and the reduction of the tusks, also seen in other dicynodonts with reduced tusks.

Inner ear morphology of diadectomorphs and seymouriamorphs (Tetrapoda) uncovered by high‐resolution x‐ray microcomputed tomography, and the origin of the amniote crown group

AbstractThe origin of amniotes was a key event in vertebrate evolution, enabling tetrapods to break their ties with water and invade terrestrial environments. Two pivotal clades of early tetrapods, the diadectomorphs and the seymouriamorphs, have played an unsurpassed role in debates about the ancestry of amniotes for over a century, but their skeletal morphology has provided conflicting evidence for their affinities. Using high‐resolution X‐ray microcomputed tomography, we reveal the three‐dimensional architecture of the well preserved endosseous labyrinth of the inner ear in representative species belonging to both groups. Data from the inner ear are coded in a new cladistic matrix of stem and primitive crown amniotes. Both maximum parsimony and Bayesian inference analyses retrieve seymouriamorphs as derived non‐crown amniotes and diadectomorphs as sister group to synapsids. If confirmed, this sister group relationship invites re‐examination of character polarity near the roots of the crown amniote radiation. Major changes in the endosseous labyrinth and adjacent braincase regions are mapped across the transition from non‐amniote to amniote tetrapods and include: a ventral shift of the cochlear recess relative to the vestibule and the semicircular canals; cochlear recess (primitively housed exclusively within the opisthotic) accommodated within both the prootic and the opisthotic; development of a distinct fossa subarcuata. The inner ear of seymouriamorphs foreshadows conditions of more derived groups, whereas that of diadectomorphs shows a mosaic of plesiomorphic and apomorphic traits, some of which are unambiguously amniote‐like, including a distinct and pyramid‐like cochlear recess.

Smooth Muscle in Cardiac Chambers is Common in Turtles and Extensive in the Emydid Turtle, Trachemys scripta.

ABSTRACTA prominent layer of smooth muscle lining the luminal side of the atria of freshwater turtles (Emydidae) was described more than a century ago. We recently demonstrated that this smooth muscle provides a previously unrecognized mechanism to change cardiac output in the emydid red‐eared slider (Trachemys scripta) that possibly contributes to their tremendous diving capacity. The purpose of the present immunohistochemical study was firstly to screen major groups of vertebrates for the presence of cardiac smooth muscle. Secondly, we investigated the phylogenetic distribution of cardiac smooth muscle within the turtle order (Testudines), including terrestrial and aquatic species. Atrial smooth muscle was not detected in a range of vertebrates, including Xenopus laevis, Alligator mississippiensis, and Caiman crocodilus, all of which have pronounced diving capacities. However, we confirmed earlier reports that traces of smooth muscle are found in human atrial tissue. Only within the turtles (eight species) was there substantial amounts of nonvascular smooth muscle in the heart. This amount was greatest in the atria, while the amount in proportion to cardiac muscle was greater in the sinus venosus than in other chambers. T. scripta had more smooth muscle in the sinus venosus and atria than the other turtles. In some specimens, there was some smooth muscle in the ventricle and the pulmonary vein. Our study demonstrates that cardiac smooth muscle likely appeared early in turtle evolution and has become extensive within the Emydidae family, possibly in association with diving. Across other tetrapod clades, cardiac smooth muscle might not associate with diving. Anat Rec, 303:1327–1336, 2020. © 2019 The Authors. The Anatomical Record published by Wiley Periodicals, Inc. on behalf of American Association for Anatomy.

Bone histology of two pareiasaurs from Russia (Deltavjatia rossica and Scutosaurus karpinskii) with implications for pareiasaurian palaeobiology

Abstract Pareiasaurs were one of the main clades of large herbivorous tetrapods in Middle–Late Permian continental ecosystems. Despite abundant pareiasaur material, many aspects of their biology remain poorly known. This paper provides a description of ontogenetic changes in long-bone and rib microanatomy/histology of two Upper Permian pareiasaurs from Russia, Deltavjatia rossica and Scutosaurus karpinskii. Analysis of a growth series of bones of Deltavjatia and Scutosaurus revealed rapid and cyclical growth early in ontogeny (as indicated by fast-growing fibrolamellar bone with lines of arrested growth). This was followed by a change in the growth pattern (as indicated by an outer avascular layer of lamellar bone in the cortex) and a decrease in the growth rate after 50% of maximum body size was reached in Deltavjatia and 75% in Scutosaurus (larger body sizes in Scutosaurus were attained through an extended initial period of fast skeletal growth). The study confirms that the bones of all pareiasaurs are histologically and microanatomically uniform [spongious (porous) microstructure and a very thin compact cortex] and indicate a similar growth strategy (a short initial period of rapid and cyclical growth followed by a long period of slow growth). The microanatomical characteristics of pareiasaurs do not provide a clear indication of their lifestyle.

A new bystrowianid from the upper Permian of Germany: first record of a Permian chroniosuchian (Tetrapoda) outside Russia and China

ABSTRACTAn isolated dorsal osteoderm of a chroniosuchian from a late Permian fissure filling in the lower Zechstein (Z1) of central Germany represents the first Permian-age record of this enigmatic tetrapod clade outside Russia and China. Based on a number of features, the specimen is designated the holotype of a new taxon, Hassiacoscutum munki, and referred to the chroniosuchian clade Bystrowianidae. Phylogenetic analysis based on a recently published data set with addition of Hassiacoscutum and other recently described bystrowianids (Yumenerpeton, Jiyuanitectum, and Vyushkoviana) found good support for the clade Chroniosuchia and its subclades Chroniosuchidae and Bystrowianidae, but with little internal resolution for the latter. A second phylogenetic analysis of all known bystrowianid genera based solely on osteoderm characters yielded a well-supported and well-resolved clade Bystrowianidae with two subclades: (Yumenerpeton + (Synesuchus + Bystrowiella)) and (Jiyuanitectum + (Bystrowiana + (Hassiacoscutum + (Vyushkoviana + (Dromotectum + Axitectum))))). Hassiacoscutum provides a paleobiogeographic link to other middle and late Permian assemblages of Laurasian continental tetrapods. To date, chroniosuchians are known only from Laurasia, and their apparent absence in Gondwana hints at faunal provincialism across Pangaea.

A single-cell transcriptomic atlas of the developing chicken limb

BackgroundThrough precise implementation of distinct cell type specification programs, differentially regulated in both space and time, complex patterns emerge during organogenesis. Thanks to its easy experimental accessibility, the developing chicken limb has long served as a paradigm to study vertebrate pattern formation. Through decades’ worth of research, we now have a firm grasp on the molecular mechanisms driving limb formation at the tissue-level. However, to elucidate the dynamic interplay between transcriptional cell type specification programs and pattern formation at its relevant cellular scale, we lack appropriately resolved molecular data at the genome-wide level. Here, making use of droplet-based single-cell RNA-sequencing, we catalogue the developmental emergence of distinct tissue types and their transcriptome dynamics in the distal chicken limb, the so-called autopod, at cellular resolution.ResultsUsing single-cell RNA-sequencing technology, we sequenced a total of 17,628 cells coming from three key developmental stages of chicken autopod patterning. Overall, we identified 23 cell populations with distinct transcriptional profiles. Amongst them were small, albeit essential populations like the apical ectodermal ridge, demonstrating the ability to detect even rare cell types. Moreover, we uncovered the existence of molecularly distinct sub-populations within previously defined compartments of the developing limb, some of which have important signaling functions during autopod pattern formation. Finally, we inferred gene co-expression modules that coincide with distinct tissue types across developmental time, and used them to track patterning-relevant cell populations of the forming digits.ConclusionsWe provide a comprehensive functional genomics resource to study the molecular effectors of chicken limb patterning at cellular resolution. Our single-cell transcriptomic atlas captures all major cell populations of the developing autopod, and highlights the transcriptional complexity in many of its components. Finally, integrating our data-set with other single-cell transcriptomics resources will enable researchers to assess molecular similarities in orthologous cell types across the major tetrapod clades, and provide an extensive candidate gene list to functionally test cell-type-specific drivers of limb morphological diversification.

The transcriptome of the veiled chameleon (Chamaeleo calyptratus): A resource for studying the evolution and development of vertebrates.

The veiled chameleon (Chamaeleo calyptratus) is an emerging model system for studying functional morphology and evolutionary developmental biology (evo-devo). Chameleons possess body plans that are highly adapted to an arboreal life style, featuring laterally compressed bodies, split hands/ft for grasping, a projectile tongue, turreted independently moving eyes, and a prehensile tail. Despite being one of the most phenotypically divergent clades of tetrapods, genomic resources for chameleons are severely lacking. To address this lack of resources, we used RNAseq to generate 288 million raw Illumina sequence reads from four adult tissues (male and female eyes and gonads) and whole embryos at three distinct developmental stages. We used these data to assemble a largely complete de novo transcriptome consisting of only 82 952 transcripts. In addition, a majority of assembled transcripts (67%) were successfully annotated. We then demonstrated the utility of these data in the context of studying visual system evolution by examining the content of veiled chameleon opsin genes to show that chameleons possess all five ancestral tetrapod opsins. We present this de novo, annotated, multi-tissue transcriptome assembly for the Veiled Chameleon, Chamaeleo calyptratus, as a resource to address a range of evolutionary and developmental questions. The associated raw reads and final annotated transcriptome assembly are freely available for use on NCBI and Figshare, respectively.

New age constraints for the Middle Triassic archosaur Lotosaurus: Implications for basal archosaurian appearance and radiation in South China

Archosaurians were the dominant terrestrial tetrapods during the Mesozoic and became one of the most successful tetrapod clades on Earth following the end-Permian mass extinction. The lower Badong Formation in northwestern Hunan Province, South China hosts fossils representing the basal archosaurian Lotosaurus. The Badong Formation has been previously interpreted as Anisian-Ladinian (247–237 Ma) in age based on biostratigraphic correlations. This study presents new detrital zircon UPb age constraints for sandstones hosting Lotosaurus fossils in Sangzhi County of northwestern Hunan. Our data give a weighted mean age of 238.0 ± 1.4 Ma (Ladinian Stage) as the maximum depositional age for the fossil-bearing layer in the lower Badong Formation. This age is consistent with younger estimates from previous biostratigraphic studies. It is also more precise and stratigraphically constrained than previously published radiometric dating results for the units. This age estimate provides critical geochronologic constraint on the radiation of early archosaurians from China into other areas of Pangea beginning in the Early Triassic.

Laosuchus naga gen. et sp. nov., a new chroniosuchian from South-East Asia (Laos) with internal structures revealed by micro-CT scan and discussion of its palaeobiology

Chroniosuchians were a clade of non-amniotic tetrapods known from the Guadalupian (middle Permian) to Late Triassic, mainly from Russia and China. The rarity of complete or articulated remains means that relatively little is known about this group in terms of its anatomy, palaeobiology or evolutionary history. Based on a nearly complete skull with a left hemimandible, we describe the first chroniosuchian from Laos from rocks preserving the Permian–Triassic boundary in the Luang Prabang Basin, which is located on the tectonic Indochina Block. This specimen is referred to a new genus and species, Laosuchus naga, based on numerous diagnostic features, including an extremely reduced pineal foramen; absence of palatal dentition; well-developed transverse flange of the pterygoid that contacts the maxilla; internal crest on and above the dorsal side the palate; otic notch closed by the tabular horn and the posterior part of the squamosal, forming a continuous wall; thin and high ventromedial ridge on parasphenoid. A phylogenetic analysis of 51 characters and 25 taxa reveals a basal position for Laosuchus naga among Chroniosuchia. In addition, CT scan data reveal internal structures and provide new insights about the anatomy and palaeobiology of chroniosuchians. Laosuchus naga was likely amphibious and spent most of its time in water, rather than in terrestrial environments. As chroniosuchians are non-marine tetrapods previously known on the North China Block, South China Block and Laurussia, the occurrence of Laosuchus naga on the Indochina Block supports the hypothesis of physical connections between all these tectonic plates by the time of the Permo–Triassic boundary.http://zoobank.org/urn:lsid:zoobank.org:pub:11D07FA3-0F4C-4EF9-A416-E8E6BE76C970

Euryhaline ecology of early tetrapods revealed by stable isotopes.

The fish-to-tetrapod transition-followed later by terrestrialization-represented a major step in vertebrate evolution that gave rise to a successful clade that today contains more than 30,000 tetrapod species. The early tetrapod Ichthyostega was discovered in 1929 in the Devonian Old Red Sandstone sediments of East Greenland (dated to approximately 365 million years ago). Since then, our understanding of the fish-to-tetrapod transition has increased considerably, owing to the discovery of additional Devonian taxa that represent early tetrapods or groups evolutionarily close to them. However, the aquatic environment of early tetrapods and the vertebrate fauna associated with them has remained elusive and highly debated. Here we use a multi-stable isotope approach (δ13C, δ18O and δ34S) to show that some Devonian vertebrates, including early tetrapods, were euryhaline and inhabited transitional aquatic environments subject to high-magnitude, rapid changes in salinity, such as estuaries or deltas. Euryhalinity may have predisposed the early tetrapod clade to be able to survive Late Devonian biotic crises and then successfully colonize terrestrial environments.

The indestructible insect: Velvet ants from across the United States avoid predation by representatives from all major tetrapod clades.

Velvet ants are a group of parasitic wasps that are well known for a suite of defensive adaptations including bright coloration and a formidable sting. While these adaptations are presumed to function in antipredator defense, observations between potential predators and this group are lacking. We conducted a series of experiments to determine the risk of velvet ants to a host of potential predators including amphibians, reptiles, birds, and small mammals. Velvet ants from across the United States were tested with predator's representative of the velvet ants native range. All interactions between lizards, free‐ranging birds, and a mole resulted in the velvet ants survival, and ultimate avoidance by the predator. Two shrews did injure a velvet ant, but this occurred only after multiple failed attacks. The only predator to successfully consume a velvet ant was a single American toad (Anaxyrus americanus). These results indicate that the suite of defenses possessed by velvet ants, including aposematic coloration, stridulations, a chemical alarm signal, a hard exoskeleton, and powerful sting are effective defenses against potential predators. Female velvet ants appear to be nearly impervious to predation by many species whose diet is heavily derived of invertebrate prey.

The Only Known Jawed Vertebrate with Four Eyes and the Bauplan of the Pineal Complex

The pineal and parapineal organs are dorsal outpocketings of the vertebrate diencephalon that play key roles in orientation and in circadian and annual cycles. Lampreys are four eyed in that both the pineal and parapineal form eyelike photosensory structures, but the pineal is the dominant or sole median photosensory structure in most lower vertebrate clades. The pineal complex has been thought to evolve in a single direction by losing photosensory and augmenting secretory function in the transitions from three-eyed lower vertebrates to two-eyed mammals and archosaurs [1-3]. Yet the widely accepted elaboration of the parapineal instead of the pineal as the primary median photosensory organ [4] in Lepidosauria (lizards, snakes, and tuataras) hints at a more complex evolutionary history. Here we present evidence that a fourth eye re-evolved from the pineal organ at least once within vertebrates, specifically in an extinct monitor lizard, Saniwa ensidens, in which pineal and parapineal eyes were present simultaneously. The tandem midline location of these structures confirms in a striking fashion the proposed homology of the parietal eye with the parapineal organ and refutes the classical model of pineal bilaterality. It furthermore raises questions about the evolution and functional interpretation of the median photosensory organ in other tetrapod clades.

The evolution of the tetrapod humerus: morphometrics, disparity, and evolutionary rates

ABSTRACTThe present study explores the macroevolutionary dynamics of shape changes in the humeri of all major grades and clades of early tetrapods and their fish-like forerunners. Coordinate point eigenshape analysis applied to humeral outlines in extensor view reveals that fish humeri are more disparate than those of most early tetrapod groups and significantly separate from the latter. Our findings indicate sustained changes in humeral shape in the deepest portions of the tetrapod stem group and certain portions of the crown. In the first half of sampled tetrapod history, subclades show larger than expected humeral disparity, suggesting rapid diffusion into morphospace. Later in tetrapod evolution, subclades occupy smaller and non-overlapping morphospace regions. This pattern may reflect in part increasing specialisations in later tetrapod lineages. Bayesian shifts in rates of evolutionary change are distributed discontinuously across the phylogeny, and most of them occur within rather than between major groups. Most shifts with the highest Bayesian posterior probabilities are observed in lepospondyls. Similarly, maximum likelihood analyses of shifts support marked rate accelerations in lepospondyls and in various subclades within that group. In other tetrapod groups, rates either tend to slow down or experience only small increases. Somewhat surprisingly, no shifts are concurrent with structural, functional, or ecological innovations in tetrapod evolution, including the origin of digits, the water–land transition and increasing terrestrialisation. Although counterintuitive, these results are consistent with a model of continual phenotypic innovation that, although decoupled from key evolutionary changes, is possibly triggered by niche segregation in divergent clades and grades of early tetrapods.

Miniaturization and morphological evolution in Paleozoic relatives of living amphibians: a quantitative approach

AbstractMiniaturization has been defined as the evolution of extremely small adult size in a lineage. It does not simply imply the decrease of the body size but also involves structural modifications to maintain functional efficiency at a strongly reduced size. Miniaturization has been proposed as a key factor in the origin of several major tetrapod clades. Current hypotheses propose that the living amphibians (lissamphibians) originated within a clade of Paleozoic dwarfed dissorophoid temnospondyls. Morphological traits shared by these small dissorophoids have been interpreted as resulting from constraints imposed by the extreme size reduction, but these statements were based only on qualitative observations. Herein, we assess quantitatively morphological changes in the skull previously associated with miniaturization in the lissamphibian stem lineage by comparing evolutionary and ontogenetic allometries in dissorophoids. Our results show that these features are not comparable to the morphological consequences of extreme size reduction as documented in extant miniature amphibians, but instead they resemble immature conditions of larger temnospondyls. We conclude that the truncation of the ancestral ontogeny, and not constraints related to miniaturization, might have been the factor that played a major role in the morphological evolution of small dissorophoids. Based on our results, we discuss the putative role of miniaturization in the origin of lissamphibians within Dissorophoidea.

Foramina in plesiosaur cervical centra indicate a specialized vascular system

Abstract. The sauropterygian clade Plesiosauria arose in the Late Triassic and survived to the very end of the Cretaceous. A long, flexible neck with over 35 cervicals (the highest number of cervicals in any tetrapod clade) is a synapomorphy of Pistosauroidea, the clade that contains Plesiosauria. Basal plesiosaurians retain this very long neck but greatly reduce neck flexibility. In addition, plesiosaurian cervicals have large, paired, and highly symmetrical foramina on the ventral side of the centrum, traditionally termed subcentral foramina, and on the floor of the neural canal. We found that these dorsal and the ventral foramina are connected by a canal that extends across the center of ossification of the vertebral centrum. We posit that these foramina are not for nutrient transfer to the vertebral centrum but that they are the osteological correlates of a highly paedomorphic vascular system in the neck of plesiosaurs. This is the retention of intersegmental arteries within the vertebral centrum that are usually obliterated during sclerotome re-segmentation in early embryonic development. The foramina and canals are a rare osteological correlate of the non-cranial vascular (arterial) system in fossil reptiles. The adaptive value of the retention of the intersegmental arteries may be improved oxygen transport during deep diving and thermoregulation. These features may have been important in the global dispersal of plesiosaurians.