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.