Elongated snake-like bodies associated with limb reduction have evolved multiple times throughout vertebrate history. Limb-reduced squamates (lizards and snakes) account for the vast majority of these morphological transformations, and thus have great potential for revealing macroevolutionary transitions and modes of body-shape transformation. Here we present a comprehensive review on limb reduction, in which we examine and discuss research on these dramatic morphological transitions. Historically, there have been several approaches to the study of squamate limb reduction: (i) definitions of general anatomical principles of snake-like body shapes, expressed as varying relationships between body parts and morphometric measurements; (ii) framing of limb reduction from an evolutionary perspective using morphological comparisons; (iii) defining developmental mechanisms involved in the ontogeny of limb-reduced forms, and their genetic basis; (iv) reconstructions of the evolutionary history of limb-reduced lineages using phylogenetic comparative methods; (v) studies of functional and biomechanical aspects of limb-reduced body shapes; and (vi) studies of ecological and biogeographical correlates of limb reduction. For each of these approaches, we highlight their importance in advancing our understanding, as well as their weaknesses and limitations. Lastly, we provide suggestions to stimulate further studies, in which we underscore the necessity of widening the scope of analyses, and of bringing together different perspectives in order to understand better these morphological transitions and their evolution. In particular, we emphasise the importance of investigating and comparing the internal morphology of limb-reduced lizards in contrast to external morphology, which will be the first step in gaining a deeper insight into body-shape variation.

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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.

The limbless skink Ophiomorus punctatissimus is a cryptozoic species found in the Peloponnese region of Greece and on the Greek island Kythira. To provide the first thorough description of the cranial and postcranial osteology of this species, both disarticulated specimens and X‐ray computed tomographies of wet‐preserved specimens were examined in detail. Resulting from this, an anatomical atlas of this species is provided. Two separate considerations, an evolutionary and an ecomorphological one, are made based on the observed adaptations related to limb loss in this skink. The structure of the girdles shows a particular pattern of reduction: whereas the pelvic girdle is mostly vestigial, the pectoral girdle is instead well developed, with all the elements typical of limbed lizards except for the actual limbs. This led us to hypothesize an asynchronous pattern of limb reduction during the evolution of this species, in which the hindlimbs regressed earlier than the forelimbs. Furthermore, considerations based on overall body morphology, osteology and the structure of the inner ear led to the recognition of this species as a burrowing ecomorph. In contrast to the morphology normally displayed in this ecomorph, O. punctatissimus is characterized by the retention of autotomic vertebrae in its tail. This is consistent with the habitats in which it lives, where active burrowing would be difficult because of the hard, rocky terrain. Instead, this skink hides among rocks on the surface and is, therefore, subject to greater predation risk.

Many animals have strict diel activity patterns, with unique adaptations for either diurnal or nocturnal activity. Diel activity is phylogenetically conserved, yet evolutionary shifts in diel activity occur and lead to important changes in an organism's morphology, physiology, and behavior. We use phylogenetic comparative methods to examine the evolutionary history of diel activity in skinks, one of the largest families of terrestrial vertebrates. We examine how diel patterns are associated with microhabitat, ambient temperatures, and morphology. We found support for a nondiurnal ancestral skink. Strict diurnality in crown group skinks only evolved during the Paleogene. Nocturnal habits are associated with fossorial activity, limb reduction and loss, and warm temperatures. Our results shed light on the evolution of diel activity patterns in a large radiation of terrestrial ectotherms and reveal how both intrinsic biotic and extrinsic abiotic factors can shape the evolution of animal activity patterns.

The skull osteology of Hierophis viridiflavus is here described and figured in detail on the basis of 18 specimens. The sample includes specimens from the ranges of both Hierophis viridiflavus viridiflavus and Hierophis viridiflavus carbonarius as well as specimens not identified at This article is protected by copyright. All rights reserved. This is the author manuscript accepted for publication and has undergone full peer review but has not been through the copyediting, typesetting, pagination and proofreading process, which may lead to differences between this version and the Version of Record. Please cite this article as

Aim: 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 | 429 CAMAITI et al.

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