Frequency-dependent and correlational selection pressures have conflicting consequences for assortative mating in a color-polymorphic lizard, Uta stansburiana

Acknowledgments We would like to thank the numerous undergraduate researchers involved with this project for their invaluable assistance in lizard rearing and data collection. We also thank D. Haisten, A. Runemark, Y. Takahashi, and M. Verzijden for insightful comments on the manuscript. This project was funded by National Science Foundation DEBOS-15973 to A.G.M. and B.R.S.Peer reviewedPublisher PD

Beyond fossil calibrations: realities of molecular clock practices in evolutionary biology

Molecular-based divergence dating methods, or molecular clocks, are the primary neontological tool for estimating the temporal origins of clades. While the appropriate use of vertebrate fossils as external clock calibrations has stimulated heated discussions in the paleontological community, less attention has been given to the quality and implementation of other calibration types. In lieu of appropriate fossils, many studies rely on alternative sources of age constraints based on geological events, substitution rates and heterochronous sampling, as well as dates secondarily derived from previous analyses. To illustrate the breadth and frequency of calibration types currently employed, we conducted a literature survey of over 600 articles published from 2007 to 2013. Over half of all analyses implemented one or more fossil dates as constraints, followed by geological events and secondary calibrations (15% each). Vertebrate taxa were subjects in nearly half of all studies, while invertebrates and plants together accounted for 43%, followed by viruses, protists and fungi (3% each). Current patterns in calibration practices were disproportionate to the number of discussions on their proper use, particularly regarding plants and secondarily derived dates, which are both relatively neglected in methodological evaluations. Based on our survey, we provide a comprehensive overview of the latest approaches in clock calibration, and outline strengths and weaknesses associated with each. This critique should serve as a call to action for researchers across multiple communities, particularly those working on clades for which fossil records are poor, to develop their own guidelines regarding selection and implementation of alternative calibration types. This issue is particularly relevant now, as time-calibrated phylogenies are used for more than dating evolutionary origins, but often serve as the backbone of investigations into biogeography, diversity dynamics and rates of phenotypic evolution

Integration of Bayesian molecular clock methods and fossil-based soft bounds reveals early Cenozoic origin of African lacertid lizards

Background Although current molecular clock methods offer greater flexibility in modelling historical evolutionary events, calibration of the clock with dates from the fossil record is still problematic for many groups. Here we implement several new approaches in molecular dating to estimate evolutionary ages of Lacertidae, an Old World family of lizards with a poor fossil record and uncertain phylogeny. Four different models of rate variation are tested in a new program for Bayesian phylogenetic analysis called TreeTime, based on a combination of mitochondrial and nuclear gene sequences. We incorporate paleontological uncertainty into divergence estimates by expressing multiple calibration dates as a range of probabilistic distributions. We also test the reliability of our proposed calibrations by exploring effects of individual priors on posterior estimates. Results According to the most reliable model, as indicated by Bayes factor comparison, modern lacertids arose shortly after the K/T transition and entered Africa about 45 million years ago, with the majority of their African radiation occurring in the Eocene and Oligocene. Our findings indicate much earlier origins for these clades than previously reported, and we discuss our results in light of paleogeographic trends during the Cenozoic. Conclusions This study represents the first attempt to estimate evolutionary ages of a specific group of reptiles exhibiting uncertain phylogenetic relationships, molecular rate variation and a poor fossil record. Our results emphasize the sensitivity of molecular divergence dates to fossil calibrations, and support the use of combined molecular data sets and multiple, well-spaced dates from the fossil record as minimum node constraints. The bioinformatics program used here, TreeTime, is publicly available, and we recommend its use for molecular dating of taxa faced with similar challenges

Ontogenetic allometry underlies trophic diversity in sea turtles (Chelonioidea)

Despite only comprising seven species, extant sea turtles (Cheloniidae and Dermochelyidae) display great ecological diversity, with most species inhabiting a unique dietary niche as adults. This adult diversity is remarkable given that all species share the same dietary niche as juveniles. These ontogenetic shifts in diet, as well as a dramatic increase in body size, make sea turtles an excellent group to examine how morphological diversity arises by allometric processes and life habit specialisation. Using three-dimensional geometric morphometrics, we characterise ontogenetic allometry in the skulls of all seven species and evaluate variation in the context of phylogenetic history and diet. Among the sample, the olive ridley (Lepidochelys olivacea) has a seemingly average sea turtle skull shape and generalised diet, whereas the green (Chelonia mydas) and hawksbill (Eretmochelys imbricata) show different extremes of snout shape associated with their modes of food gathering (grazing vs. grasping, respectively). Our ontogenetic findings corroborate previous suggestions that the skull of the leatherback (Dermochelys coriacea) is paedomorphic, having similar skull proportions to hatchlings of other sea turtle species and retaining a hatchling-like diet of relatively soft bodied organisms. The flatback sea turtle (Natator depressus) shows a similar but less extreme pattern. By contrast, the loggerhead sea turtle (Caretta caretta) shows a peramorphic signal associated with increased jaw muscle volumes that allow predation on hard shelled prey. The Kemp’s ridley (Lepidochelys kempii) has a peramorphic skull shape compared to its sister species the olive ridley, and a diet that includes harder prey items such as crabs. We suggest that diet may be a significant factor in driving skull shape differences among species. Although the small number of species limits statistical power, differences among skull shape, size, and diet are consistent with the hypothesis that shifts in allometric trajectory facilitated diversification in skull shape as observed in an increasing number of vertebrate groups

Integration of molecules and new fossils supports a Triassic origin for Lepidosauria (lizards, snakes, and tuatara)

Background Lepidosauria (lizards, snakes, tuatara) is a globally distributed and ecologically important group of over 9,000 reptile species. The earliest fossil records are currently restricted to the Late Triassic and often dated to 227 million years ago (Mya). As these early records include taxa that are relatively derived in their morphology (e.g. Brachyrhinodon), an earlier unknown history of Lepidosauria is implied. However, molecular age estimates for Lepidosauria have been problematic; dates for the most recent common ancestor of all lepidosaurs range between approximately 226 and 289 Mya whereas estimates for crown-group Squamata (lizards and snakes) vary more dramatically: 179 to 294 Mya. This uncertainty restricts inferences regarding the patterns of diversification and evolution of Lepidosauria as a whole. Results Here we report on a rhynchocephalian fossil from the Middle Triassic of Germany (Vellberg) that represents the oldest known record of a lepidosaur from anywhere in the world. Reliably dated to 238–240 Mya, this material is about 12 million years older than previously known lepidosaur records and is older than some but not all molecular clock estimates for the origin of lepidosaurs. Using RAG1 sequence data from 76 extant taxa and the new fossil specimens two of several calibrations, we estimate that the most recent common ancestor of Lepidosauria lived at least 242 Mya (238–249.5), and crown-group Squamata originated around 193 Mya (176–213). Conclusion A Early/Middle Triassic date for the origin of Lepidosauria disagrees with previous estimates deep within the Permian and suggests the group evolved as part of the faunal recovery after the end-Permain mass extinction as the climate became more humid. Our origin time for crown-group Squamata coincides with shifts towards warmer climates and dramatic changes in fauna and flora. Most major subclades within Squamata originated in the Cretaceous postdating major continental fragmentation. The Vellberg fossil locality is expected to become an important resource for providing a more balanced picture of the Triassic and for bridging gaps in the fossil record of several other major vertebrate groups

A database of the morphology, ecology and literature of the world's limb‐reduced skinks

Aim Limb-reduced squamates are a convenient model system to investigate macroevolutionary trends in morphology. Here, we provide morphological, ecological and literature data on all known species of limb-reduced skinks (Scincidae) and their relatives, representing one of the most diverse and widely distributed groups of limb-reduced squamates. Location Global. Taxon Skinks (Reptilia, Squamata: Scincidae). Limb-reduced forms. Methods Morphological data were sourced from the primary literature, spanning a period of over 150 years. Linear body measurements were averaged across all values in the literature, preserving proportionality to body length. For digits and presacral vertebrae, we used maximum recorded counts. Ecological and biogeographical data were sourced from habitat assessments in the primary literature, online databases and field guides. Literature data were sorted according to type of study. To exemplify the applicability of the database, we used Markov-chain ordered models to estimate the evolutionary frequency of limb reduction and loss in skinks. Results We find evidence of limb reduction and loss in a total of 394 species worldwide, representing ~23% of all skink species, and ~30% of genera. The distribution of limb-reduced and limbless forms differs from that of fully limbed forms, as they are present in all biogeographic realms with the almost complete exclusion of the Americas. We estimate that limb reduction evolved more than 50 times in skinks, and that loss of at least one limb pair evolved at least 24 times. Main conclusions The dataset captures a broad spectrum of morphological and ecological variation in a large, globally distributed taxonomic group. It establishes a widely applicable definition of limb reduction based on limb proportions as a reference for future studies. Such an extensive collection of morphological and ecological data can pave the way for investigations of dramatic morphological transitions and their ecological drivers at a global and local scale

Macroecological and biogeographical patterns of limb reduction in the world's skinks

First published: 08 December 2022Aim: Limb reduction is a dramatic evolutionary transition, yet whether it is achieved in similar trajectories across clades, and its environmental drivers, remain unclear. We investigate the macroevolutionary and biogeographical patterns of limb reduction in skinks, where limb reduction occurred more often than in any other tetrapod clade, and test their associations with substrate categories using a global database. We test for habitat associations of body shapes in a group of Australian skinks using quantitative habitat data. Location: Global (Scincidae), Australia (Sphenomorphinae). Taxon: Skinks, Australian Sphenomorphinae. Materials and Methods: We use morphological data to explore the patterns of limb reduction in the world's skinks, investigating how body proportions differ across skink clades and subfamilies. We examine the relationships between body shape and substrate (coarsely classified). Further, we investigate the relationships between body shape and high-resolution soil and climate properties extracted from each species' distribution for Australian sphenomorphines. Results: Relationships between limb lengths and trunk elongation show idiosyncratic patterns across skink clades. Presacral vertebrae numbers positively correlate with trunk elongation in all taxa, except Glaphyromorphus. Skinks from sandy habitats show greater disparity between forelimb and hindlimb lengths than all other substrate categories. In sphenomorphines, shorter limbs and elongated trunks correlate with colder, more humid microhabitats and richer soils; high limb disparity correlates with hot, arid microhabitats and sandy, poor substrates. Main Conclusions: The evolutionary trajectories of limb reduction in skinks are cladespecific and sometimes unique. Selection for specific limb proportions and body sizes in limb-reduced forms changes across substrates. On poor, sandy substrates of arid environments, body shapes with longer hindlimbs may be more efficient for locomotion in a granular fluid (i.e. sand) and exploit the air–substrate interface than complete limblessness. On richer, more humid substrates, such morphology is rare, indicating that navigating cluttered substrates selects for more equal and shorter limb lengths.Marco Camaiti, Alistair R. Evans, Christy A. Hipsley, Mark N. Hutchinson, Shai Meiri, Rodolfo de Oliveira Anderson, Alex Slavenko, David G. Chappl