Assessing sources of variation in amphibian skin thickness: ecological and evolutionary implications

The skin is the largest organ of the body and provides many functions. Among tetrapod vertebrates, amphibian skin is semi-permeable and responsible for a greater proportion of water absorption and gas exchange. Myriad factors affect the physiological performance of amphibian skin. Morphological traits linked with amphibian skin physiology or ecology have remained difficult to discern because of a lack of quantitative comparative research and the discovery of sources of intraspecific variation that are mostly ignored in study designs. This thesis aims to address the effects of these sources of variation using a trait that is known to vary between sexes, among seasons, and among body regions and thought to be linked with physiology or ecology, skin thickness. The first source of variation addressed is sexual dimorphism. Specimens of the white-lipped treefrog, Litoria infrafrenata, that display sexual dimorphism in body size and skin thickness were used to test if body size was the main determinate of sexually dimorphic skin thickness. Size corrected values did not significantly differ between males and females, although the sample size was small. Seasonal variation in skin thickness has also been documented in some species, so the American bullfrog (Lithobates catesbeianus), the Northern leopard frog (L. pipiens), and the spring peeper (Pseudacris crucifer) from multiple months of the year were sampled to determine if skin thickness increased in the autumn or winter months. Seasonal skin thickening was only detected in L. catesbeianus, and skin from autumn and winter was significantly thicker than from earlier in the year. This pattern was also detectable in museum specimens collected 80 years ago, although the signal was damped, possibly due to preservation. Using a dataset of 10 species and published data, a general pattern was uncovered whereby the dorsal skin is the thickest region and the ventral thigh region is the thinnest. However, this pattern is not always true for every individual of every species (L. pipiens and P. crucifer) and in some species the dorsal skin is thinnest (Bokermannohyla alvarengai and Litoria infrafrenata). The same dataset found that skin thickness is significantly related to body size, as was found in the chapter on Litoria infrafrenata. Summer specimens of Lithobates catesbeianus were outliers below the interspecific regression line and winter specimens fell within the range of variation of other species, hinting that seasonal skin thickening could be renamed seasonal skin thinning in this species. Finally, a link between ecology and skin thickness was tested using the 10 species from previous analyses and data from the literature. At a phylogenetially broad scale, body size explained a greater amount of the variation in environmental parameters than skin thickness. At smaller taxonomic scales, skin thickness appears more closely linked with ecology. It is concluded that amphibians generally follow an allometric trend for skin thickness and when faced with suboptimal conditions over long periods of time, evolve integumentary structures like iridiophores to compensate for any physiological disadvantage of an ‘ideal’ skin thickness. In the interim, however, skin thickness may change, thus sacrificing e.g. mechanical support.Gates Cambridge Scholarship, German Academic Exchange Program (DAAD), Museum für Naturkunde Innovation Fun

Testing for a facultative locomotor mode in the acquisition of archosaur bipedality

Bipedal locomotion is a defining characteristic of humans and birds and has a profound effect on how these groups interact with their environment. Results from extensive hominin research indicate that there exists an intermediate stage in hominin evolution—facultative bipedality—between obligate quadrupedality and obligate bipedality that uses both forms of locomotion. It is assumed that archosaur locomotor evolution followed this sequence of functional and hence character-state evolution. However, this assumption has never been tested in a broad phylogenetic context. We test whether facultative bipedality is a transitionary state of locomotor mode evolution in the most recent early archosaur phylogenies using maximum-likelihood ancestral state reconstructions for the first time. Across a total of seven independent transitions from quadrupedality to a state of obligate bipedality, we find that facultative bipedality exists as an intermediary mode only once, despite being acquired a total of 14 times. We also report more independent acquisitions of obligate bipedality in archosaurs than previously hypothesized, suggesting that locomotor mode is more evolutionarily fluid than expected and more readily experimented with in these reptiles

The Function and Evolution of the Syncervical in Ceratopsian Dinosaurs with a Review of Cervical Fusion in Tetrapods

Mobility of the vertebral column is important for many ecological aspects of vertebrates, especially in the cervical series, which connects the head to the main body. Thus, fusion within the cervical series is hypothesized to have ecological and behavioural implications. Fused, anterior cervical vertebrae have evolved independently over 20 times in ecologically disparate amniotes, most commonly in pelagic, ricochetal, and fossorial taxa, suggesting an adaptive function for the ‘syncervical.’ Fusion may help increase out-force during head-lift digging or prevent anteroposteriorly shortened vertebrae from mechanically failing during locomotion, but no hypothesis for syncervical function has been tested. The syncervical of neoceratopsian dinosaurs is hypothesized to support large heads or aid in intraspecific combat. Tests of correlated character evolution within a ceratopsian phylogeny falsify these hypotheses, as the syncervical evolves before large heads and cranial weaponry. Alternative functional hypotheses may involve ancestral burrowing behaviour or unique feeding ecology in early neoceratopsians.MAS

Forearm Posture and Mobility in Quadrupedal Dinosaurs

Quadrupedality evolved four independent times in dinosaurs; however, the constraints associated with these transitions in limb anatomy and function remain poorly understood, in particular the evolution of forearm posture and rotational ability (i.e., active pronation and supination). Results of previous qualitative studies are inconsistent, likely due to an inability to quantitatively assess the likelihood of their conclusions. We attempt to quantify antebrachial posture and mobility using the radius bone because its morphology is distinct between extant sprawled taxa with a limited active pronation ability and parasagittal taxa that have an enhanced ability to actively pronate the manus. We used a sliding semi-landmark, outline-based geometric morphometric approach of the proximal radial head and a measurement of the angle of curvature of the radius in a sample of 189 mammals, 49 dinosaurs, 35 squamates, 16 birds, and 5 crocodilians. Our results of radial head morphology showed that quadrupedal ceratopsians, bipedal non-hadrosaurid ornithopods, and theropods had limited pronation/supination ability, and sauropodomorphs have unique radial head morphology that likely allowed limited rotational ability. However, the curvature of the radius showed that no dinosaurian clade had the ability to cross the radius about the ulna, suggesting parallel antebrachial elements for all quadrupedal dinosaurs. We conclude that the bipedal origins of all quadrupedal dinosaur clades could have allowed for greater disparity in forelimb posture than previously appreciated, and future studies on dinosaur posture should not limit their classifications to the overly simplistic extant dichotomy

Evolution and function of anterior cervical vertebral fusion in tetrapods

The evolution of vertebral fusion is a poorly understood phenomenon that results in the loss of mobility between sequential vertebrae. Non-pathological fusion of the anterior cervical vertebrae has evolved independently in numerous extant and extinct mammals and reptiles, suggesting that the formation of a ‘syncervical’ is an adaptation that arose to confer biomechanical advantage(s) in these lineages. We review syncervical anatomy and evolution in a broad phylogenetic context for the first time and provide a comprehensive summary of proposed adaptive hypotheses. The syncervical generally consists of two vertebrae (e.g. hornbills, porcupines, dolphins) but can include fusion of seven cervical vertebrae in some cetaceans. Based on the ecologies of taxa with this trait, cervical fusion most often occurs in fossorial and pelagic taxa. In fossorial taxa, the syncervical likely increases the out-lever force during head-lift digging. In cetaceans and ricochetal rodents, the syncervical may stabilize the head and neck during locomotion, although considerable variation exists in its composition without apparent variability in locomotion. Alternatively, the highly reduced cervical vertebral centra may require fusion to prevent mechanical failure of the vertebrae. In birds, the syncervical of hornbills may have evolved in response to their unique casque-butting behaviour, or due to increased head mass. The general correlation between ecological traits and the presence of a syncervical in extant taxa allows more accurate interpretation of extinct animals that also exhibit this unique trait. For example, syncervicals evolved independently in several groups of marine reptiles and may have functioned to stabilize the head at the craniocervical joint during pelagic locomotion, as in cetaceans. Overall, the origin and function of fused cervical vertebrae is poorly understood, emphasizing the need for future comparative biomechanical studies interpreted in an evolutionary context

Table-S1-locomotor-diagnoses LG CvB DN supporting 'Testing for a facultative locomotor mode in the acquisition of archosaur bipedality'

The uploaded file consists of a simple data table to support the associated article, "Testing for a facultative locomotor mode in the acquisition of archosaur bipedality". The table outlines the results of a literature survey of all taxa included in the two independent character matrices discussed in the article. The table records the most recent interpretation of locomotor mode for each taxon. The literature survey was conducted to determine whether each taxon was classified as an obligate quadruped (OQ), facultative biped (FB), or obligate biped (OB)

Forearm Posture and Mobility in Quadrupedal Dinosaurs

<div><p>Quadrupedality evolved four independent times in dinosaurs; however, the constraints associated with these transitions in limb anatomy and function remain poorly understood, in particular the evolution of forearm posture and rotational ability (i.e., active pronation and supination). Results of previous qualitative studies are inconsistent, likely due to an inability to quantitatively assess the likelihood of their conclusions. We attempt to quantify antebrachial posture and mobility using the radius bone because its morphology is distinct between extant sprawled taxa with a limited active pronation ability and parasagittal taxa that have an enhanced ability to actively pronate the manus. We used a sliding semi-landmark, outline-based geometric morphometric approach of the proximal radial head and a measurement of the angle of curvature of the radius in a sample of 189 mammals, 49 dinosaurs, 35 squamates, 16 birds, and 5 crocodilians. Our results of radial head morphology showed that quadrupedal ceratopsians, bipedal non-hadrosaurid ornithopods, and theropods had limited pronation/supination ability, and sauropodomorphs have unique radial head morphology that likely allowed limited rotational ability. However, the curvature of the radius showed that no dinosaurian clade had the ability to cross the radius about the ulna, suggesting parallel antebrachial elements for all quadrupedal dinosaurs. We conclude that the bipedal origins of all quadrupedal dinosaur clades could have allowed for greater disparity in forelimb posture than previously appreciated, and future studies on dinosaur posture should not limit their classifications to the overly simplistic extant dichotomy.</p></div

Radii of sprawling and parasagittal extant taxa.

<p>The radial head (A) and long axis (B) of <i>Caiman crocodylus</i> (ROM R7719) shows the flattened ulnar articular surface and relatively straight long axis typical of sprawling taxa. The radial head (C) and long axis (D) of <i>Ursus americanas</i> (USNM 49664) shows the rounded ulnar articular surface and curved long axis typical of parasagittal mammals and chameleons. Scale bar = 5 cm. Radial heads not to scale. <b>rh</b>, radial head; <b>sp</b>, styloid process; <b>us</b>, ulnar articular surface.</p

Results from the Kruskal-Wallis test on extant taxa.

<p>Results from the Kruskal-Wallis test on extant taxa.</p

Terrestrial radial head PCA.

<p>Principal component scores from the analysis of terrestrial taxa. PC1 vs. PC2 (A) and PC1 vs. PC3 (B) have 95% convex hulls representing the sprawled and parasagittal taxa. The origin of each axis is represented by the shape of the consensus (C). Shape change along the principal component axes is shown with the location of the consensus shown at the origin (C). Landmarks 1 (middle of the ulnar articular surface) and 10 are labeled.</p