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Scincomorpha
Scincomorpha
Scincomorpha
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Scincomorphs
Temporal range: Middle Jurassic—present, 170–0 Ma
Eastern blue-tongued lizard
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Squamata
Infraorder: Scincomorpha
Camp, 1923
Subgroups

Scincomorpha is an infraorder and clade of lizards including skinks (Scincidae) and their close relatives. These include the living families Cordylidae (girdled lizards), Gerrhosauridae (plated lizards), and Xantusiidae (night lizards), as well as many extinct taxa. Other roughly equivalent terms include the suborder Scinciformata,[2] or the superfamily Scincoidea, though different authors use these terms in a broader or more restricted usage relative to true skinks. They first appear in the fossil record about 170 million years ago, during the Jurassic period.[3]

Systematics

[edit]

Alifanov (2016) found the following phylogeny with morphological data:[4][5] Note that Dibamidae was included, unlike in analyses with molecular data.[6]

Scincomorpha

Alternatively, Zheng & Wiens (2016) found the following phylogeny of extant groups using molecular data; this phylogeny has largely been supported through other studies using molecular evidence:[6][7]

Scincoidea

There are many characteristics that are shared upon all skinks. All skinks have very cone shaped heads with large, symmetrical, and shield-like scales. Their scales are smooth, glossy, and circular all throughout their body. Once you get to their back and belly areas, they have more round scales that overlap like roof shingles. Their bodies are cylindrical in their cross section and have body scales that have bony plates underneath them called "osteoderms". Osteoderms are dermal bone structures that support the upper layer of skin and serve as protection against the elements in a large variety of extinct and extant organisms, especially reptiles. This structure is commonly called "dermal armor" and serves to protect the organism, while also helping with temperature regulation. The roofs of their mouths are made up of two bony plates instead of one. One of their bony plates is called a palate and the other bony palate in the roof of their mouth separates the respiratory and digestive passages. They also have very long tapering tails with small legs and five toes. 

For their distribution, there are around 1,275 species of skinks all around the world. They are very popular and can be mostly found in Southeast Asia, most areas of Australia, and temperate regions of North America. There are also desert species skinks that are called "sand swimmers" that are found in Florida. Five-lined skinks are very popular throughout Georgia and North Carolina, they are found in very wooded areas and like to hide in fallen trees. Skinks are more abundant and endangered in Africa and Indo Australia because of predators and loss of habitat. Another very popular area is New Zealand, the pale-flecked garden sunskink (Lampropholis guichenoti) is very common. They are also found in the suburban gardens in Auckland. Some skink species are more terrestrial and fossorial, some arboreal meaning tree-dwelling, and others are semiaquatic.

References

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Scincomorpha

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from Grokipedia
Scincomorpha is an infraorder of squamate reptiles within the order , encompassing a diverse of that includes skinks and their close relatives, such as girdled lizards, plated lizards, and night lizards. This group is characterized by its , supported by molecular phylogenetic analyses, and comprises four primary families: Scincidae (skinks), (girdled and flat lizards), Gerrhosauridae (plated lizards and grubsnakes), and Xantusiidae (night lizards). With approximately 1,900 described species, Scincomorpha represents about one-quarter of all lizard diversity and exhibits a global distribution, excluding polar regions, across a wide range of habitats from deserts and forests to islands. The Scincidae family, the largest within Scincomorpha, includes over 1,700 species and displays remarkable morphological diversity, including limb-reduced burrowing forms, limbless species, and arboreal types with elongated bodies and smooth, overlapping scales. In contrast, and Gerrhosauridae are predominantly African, featuring armored scales and osteoderms for protection, with species like the (Ouroborus cataphractus) known for their spiny, defensive postures. Xantusiidae, limited to the including , includes small, nocturnal adapted to rocky or subterranean environments, with viviparous reproduction unique among many Scincomorpha. Phylogenetically, Scincomorpha forms part of the superfamily Scincoidea and is the sister group to the more derived Episquamata clade (including lacertids, teiids, anguimorphs, and snakes), with divergence estimated around 180-200 million years ago in the Early Jurassic. Fossil records of Scincomorpha extend back to the Late Jurassic, with early forms like Ardeosaurus showing primitive traits, and the group has since radiated extensively, contributing to ecological roles as predators, prey, and ecosystem engineers in various biomes. Notable evolutionary adaptations include high speciation rates in island systems, such as the Caribbean and Indo-Australian archipelagos, driven by vicariance and adaptive radiation. Conservation concerns affect several species due to habitat loss and invasive predators, highlighting the importance of ongoing taxonomic and phylogenetic research for biodiversity management.

Etymology and definition

Name origin

The name Scincomorpha is derived from the Ancient Greek σκίγκος (skínkos), denoting a type of lizard akin to the modern skink, combined with the suffix -morphē (μορφή), meaning "form" or "shape," reflecting a grouping of lizards sharing similar body forms. This taxonomic term was first proposed by German anatomist Max Fürbringer in 1900 as part of his systematic classification of reptiles, where he used it to categorize lizards exhibiting skink-like characteristics based on comparative anatomy. Fürbringer's work emphasized structural similarities in the pectoral girdle and musculature to define higher-level groupings within Squamata. In its initial application during the early , Scincomorpha served as a morphological section within classifications, primarily denoting taxa with shared features such as smooth, overlapping scales arranged in regular rows and specific osteological traits like the configuration of the shoulder apparatus and limb elements. American herpetologist Charles Lewis Camp further elaborated on this in 1923, adopting the term to encompass a diverse array of "scincoid" unified by these external and skeletal resemblances, distinguishing them from other saurian groups like . This usage highlighted phenetic similarities in scale patterns and bone morphology rather than strict evolutionary relationships. With the advent of cladistic methods in the late , the term Scincomorpha evolved from a subordinal or sectional rank to an infraorder in modern phylogenetic systems, accommodating monophyletic clades supported by both morphological and molecular data while retaining its focus on skink-related forms. This rank adjustment reflects broader taxonomic refinements within , as seen in comprehensive revisions that integrate fossil and extant evidence.

Clade scope

Scincomorpha constitutes an infraorder and monophyletic clade within the order , comprising approximately 1,900 extant (as of 2023) that primarily consist of and their close relatives, accounting for roughly 24% of global diversity. This diversity underscores Scincomorpha's prominence among squamate reptiles, with the vast majority of species belonging to the skink families, supplemented by smaller numbers in allied families. The clade's core taxonomic structure encompasses two primary superfamilies: Scincoidea, which includes the families and Xantusiidae, and Cordyloidea, which includes the families Gerrhosauridae and . These groups represent all living scincomorphs, with the nine families harboring approximately 1,743 species combined (as of 2023), while Xantusiidae, Gerrhosauridae, and contribute fewer than 200 species combined. Scincomorpha is delineated from distantly related squamate groups, such as the infraorders Iguania and , through its , which is affirmed by morphological phylogenies emphasizing shared derived cranial features, including a well-developed alar process on the prootic and the presence of two pairs of supraocular scales (II and III).

Description

Morphological traits

Scincomorpha encompasses a diverse array of exhibiting a general characterized by cylindrical to slightly depressed bodies, with forms ranging from elongate species with reduced limbs, such as certain members of Scincidae, to more robust taxa with well-developed limbs, as seen in Gerrhosauridae. Most species attain snout-vent lengths (SVL) of 5–30 cm, though extremes exist within the . A common integumentary feature across many scincomorphs is the presence of smooth, overlapping scales, often underlain by compound osteoderms that provide dermal armor, particularly prominent in scincids. Cranially, scincomorphs share several osteological synapomorphies, including the fusion of frontal bones in adults and parietals, which contribute to a rigid roof. The postorbitofrontal complex features an anteroposteriorly elongate postfrontal component, and the squamosal lacks a dorsal process, while a palpebral is present. The shows fusion of the articular, prearticular, and surangular into a single unit, with the dentary exhibiting a straight long axis; additionally, the sphenoid possesses posterolateral ventral flanges that overlie the basioccipital laterally. A rudimentary secondary is formed by a ventromedial fold on the that partly obscures the choanal groove, and pterygoid teeth are often arranged in multiple rows or patches. The is typically absent, though reversals occur in some lineages, and the jaw adductor musculature originates ventrally on the parietal. Postcranially, scincomorphs are distinguished by compound osteoderms present both dorsally and ventrally, enhancing structural support beneath the scales. In the pelvic girdle, the symphysial portion of the pubis exceeds the tubercular portion by more than half its length, a feature unique to the . varies but commonly includes transverse or subpleurodont implantation in many members; subpleurodont attachment—where teeth are partially fused to the margin—is prevalent in scincids and other groups. The nasal process of the is reduced or absent in several lineages, contributing to a streamlined rostral morphology.

Diversity in form

Scincomorpha exhibits remarkable morphological diversity, particularly in limb structure, where members range from fully limbed terrestrial forms to highly specialized limbless species adapted for lifestyles. In the family , limb reduction has evolved repeatedly, often associated with burrowing habits, as seen in genera like Lerista, where species display a spectrum from pentadactyl (five-toed) limbs to complete absence of external limbs, facilitating efficient sand-swimming and underground . Similarly, in Brachymeles skinks, ontogenetic development leads to externally limbless adults with concealed limb rudiments beneath the skin, enhancing streamlining for subterranean movement while retaining vestigial structures. These extremes contrast with fully limbed members in families like Gerrhosauridae and , highlighting adaptive convergence for diverse microhabitats within the . Scale morphology and coloration further underscore the in Scincomorpha, with variations reflecting ecological roles from to armor. Skinks (Scincidae) typically possess smooth, scales that are shiny and imbricate, often underlain by compound osteoderms, providing flexibility for agile movement and subtle coloration patterns in browns and greens for blending into leaf litter or . In contrast, plated lizards (Gerrhosauridae) feature heavily keeled, rectangular dorsal scales embedded with osteoderms, forming a rigid, armored plating in earthy tones that offers protection against predators in rocky or arid environments. Coloration across the varies widely, from cryptic mottling in forms to bolder patterns in diurnal species, aiding in and mate signaling without compromising the shared baseline of overlapping granular scales. Sensory adaptations in Scincomorpha are tuned to activity patterns and predation pressures, with notable nocturnal modifications in groups like Xantusiidae. Night lizards possess enlarged eyes with vertical pupils and a high of photoreceptors, enhancing low-light vision for crepuscular foraging under rocks or bark, where ambient light is minimal. Tail autotomy, a widespread defense mechanism, is particularly prevalent, allowing rapid detachment at preformed fracture planes to distract predators; in scincids, the shed tail continues wriggling via stored , buying escape time while the lizard regenerates a functional, though cartilaginous, replacement. These traits collectively enable Scincomorpha to occupy niches from deserts to forests, emphasizing form-function linkages beyond unifying morphological features like lower temporal arches.

Distribution and habitat

Geographic range

Scincomorpha, a diverse of , display a primarily and subtropical distribution across all continents except , with limited extensions into temperate zones but absence from extreme polar and boreal regions. The clade's range encompasses tropical rainforests, savannas, deserts, and Mediterranean habitats, reflecting broad ecological adaptability while avoiding high-latitude extremes. The highest diversity occurs in and , where skinks (Scincidae)—the most speciose family within Scincomorpha—dominate faunas. Australia alone hosts approximately 389 skink across 38 genera, comprising about half of the country's diversity and underscoring the region's status as a global hotspot. Similarly, , , and support hundreds of skink , with these areas accounting for over 50% of species in their respective bioregions. In , Scincomorpha are represented by endemic families such as the girdled lizards (), confined to sub-Saharan regions including southern, eastern, and central areas as far north as , and the plated lizards (Gerrhosauridae), distributed across mainland , , and the . Skinks further contribute to African diversity, particularly in southern and central zones, as well as . The Americas host a more restricted presence, primarily through the night lizards (Xantusiidae), which are limited to southwestern (including parts of the and ), extending to , and . Some species occur in the and northern , but overall diversity remains low compared to the . In and , Scincomorpha include fossorial forms such as snake-eyed skinks of the genus Ablepharus, which range from southeastern (e.g., ) through southwestern to , including countries like , , and . These distributions highlight biogeographic patterns tied to Gondwanan origins, with pronounced dominance in diversity centers like and .

Ecological niches

Scincomorpha predominantly occupy terrestrial niches, with many species adapted to lifestyles, burrowing into , , or under bark in diverse environments ranging from arid deserts and grasslands to humid forests. For instance, skinks (Scincidae), which comprise the majority of the clade's diversity, exhibit a broad spectrum of habitat preferences, including 45% terrestrial forms in the subfamily Lygosominae and fully taxa in Acontiinae and parts of Scincinae, often thriving in microhabitats like loose sand or organic debris layers. Arboreal adaptations are less common but present in about 15% of Lygosominae skinks, which navigate tree bark and foliage, while plated lizards (Gerrhosauridae) favor semi-open rocky savannas and mounds for shelter. Girdled lizards () are typically rupicolous, inhabiting crevices in rocky outcrops of semi-arid regions, and night lizards (Xantusiidae) are crevice-dwellers in arid to mesic rocky terrains, including deserts and woodlands. Dietary habits within Scincomorpha are primarily insectivorous, reflecting their role as active foragers in ground-level or subterranean food webs, with approximately 90% of skinks consuming arthropods such as and spiders. Omnivorous tendencies appear in about 9% of skinks and some plated lizards, which incorporate vegetation, small vertebrates, and fruits, allowing exploitation of varied resources in and edges. Herbivory is rare, limited to roughly 1% of skinks, often in insular or resource-scarce settings. Defensive behaviors emphasize and evasion, including caudal —tail shedding for distraction—common across skinks and night lizards, alongside rapid hiding in burrows or crevices to avoid predators. Reproductive strategies vary, with oviparity dominant in most skinks (about 65%), where females lay clutches of 3–4 eggs in concealed nests within soil or litter, but viviparity has evolved independently at least 31 times in the clade, notably in all (e.g., girdled lizards producing live young in rocky refugia) and Xantusiidae, enhancing survival in unstable or cold-prone habitats. These modes tie directly to ecological pressures, such as predation risk in open grasslands versus thermal regulation in fossorial environments.

Evolutionary history

Fossil record

The fossil record of Scincomorpha begins in the , with the earliest definitive evidence consisting of paramacellodid lizards from the stage (approximately 167 million years ago) of the Forest Marble Formation at Kirtlington Quarry, , . These specimens, including isolated cranial elements, represent primitive scincomorphs characterized by acrodont dentition and robust skulls, marking the initial appearance of the within during a period of early squamate diversification. Subsequent paramacellodid finds from deposits, such as the in the western United States, further illustrate their persistence and modest radiation through the end of the . Recent discoveries, such as Eoscincus ornatus from the (~150 Ma), provide additional insight into early scincomorph morphology and synapomorphies like dentition. Mesozoic diversity expanded markedly during the , encompassing a range of transversely-toothed scincomorphs that dominated lizard faunas in both hemispheres. Polyglyphanodontids, a key group featuring pleurodont teeth oriented perpendicular to the jaw, are particularly well-documented from ( to ) localities across , including the Kaiparowits Formation in , , where genera like Polyglyphanodon and Dicothodon exhibit specialized herbivorous or omnivorous adaptations. In , scincomorphan fossils from the Lower Sasayama Group ( stage) in Hyogo Prefecture, , include jaw fragments tentatively referred to paramacellodids or early teiioids, highlighting an early eastward dispersal. Gondwanan records are rarer but significant, with a partial dentary of an unnamed scincomorph from the early Los Alamitos Formation , , providing evidence of the clade's presence in by the . In the , following the Cretaceous-Paleogene mass extinction that eliminated many scincomorph lineages, surviving taxa radiated into modern families such as Scincidae, , and Xantusiidae, with fossils documenting their global proliferation. deposits in , like the early Eocene of the , yield scincoid remains, while North American Eocene sites preserve diverse scincomorphs including early xantusiids. In , the record for scincomorphs is sparser, with scattered remains reflecting limited diversification. Overall, the contributes substantially to the extinct scincomorph genera documented worldwide, underscoring the clade's resilience and ongoing diversification.

Phylogenetic origins

Scincomorpha occupies a pivotal position within the order as the clade Scincoidea, comprising families such as Scincidae, , Gerrhosauridae, and Xantusiidae; molecular analyses place it as the to the broader Episquamata clade (including Lacertiformes, , and ), excluding Gekkota and other basal groups. Traditional morphological phylogenies had defined a broader Scincomorpha including (, , Gymnophthalmidae) as to , but this view is largely superseded by molecular evidence reflecting ongoing refinements in squamate relationships. Scincomorpha shares a common ancestry with other squamates tracing back to the , with crown-group estimated to have originated around 206 million years ago, and the divergence of scincomorph lineages occurring approximately 180-200 million years ago during the . This early radiation aligns with the broader lepidosaur diversification, where squamates emerged alongside rhynchocephalians, setting the stage for subsequent ecological expansions. Fossil evidence, including specimens from the (~145 Ma), indicates that scincomorph ancestors were already distinct by the mid-Mesozoic, with trans-Atlantic distributions suggesting Laurasian origins prior to continental fragmentation. Key synapomorphies defining Scincomorpha, as identified from and extant material, include the presence of ventral body osteoderms, vermiculate dermal rugosities on cranial bones indicative of imbricated scale patterns, and a posteriorly broadened retroarticular process in the lower jaw. features, such as arrangements with unicuspid or tricuspid teeth observed in Jurassic fossils like Eoscincus ornatus, further support these affinities, alongside traits like the lack of nasal-prefrontal contact and enlarged cephalic scales. These characteristics, preserved in three-dimensionally ized skulls from deposits, trace the clade's evolutionary roots and distinguish it from contemporaneous squamate outgroups.

Systematics and classification

Historical taxonomy

The concept of Scincomorpha originated with early 19th-century efforts to classify based on shared morphological traits resembling those of skinks (Scincidae), such as smooth, overlapping scales and pleurodont dentition. In 1811, Martin Oppel introduced the superfamily Scincoidea to encompass these scincoid-like forms, emphasizing cranial and squamation similarities among certain lacertilian . By the late 19th and early 20th centuries, comparative anatomists refined this grouping through detailed studies of skeletal and muscular features. Max Fürbringer's 1900 monograph on the and musculature highlighted osteological parallels, such as compound osteoderms and specific throat muscle arrangements, reinforcing Scincoidea as a cohesive assemblage distinct from other groups like iguanians. A pivotal advancement came in 1923 when Charles L. Camp elevated Scincoidea to subordinal status as Scincomorpha within Lacertilia, based on an extensive review of morphology, fossils, and . Camp's system positioned Scincomorpha as part of the autarchoglossans, including families like Scincidae, Gerrhosauridae, Xantusiidae, , and , united by traits including laminate hemipenes, dilated clavicles, and successive tooth replacement. Mid-20th-century largely retained Camp's framework amid ongoing debates over group inclusions, with Scincomorpha encompassing teiids (e.g., Cnemidophorus) and lacertids (e.g., Lacerta) due to presumed shared evolutionary origins in pleurodonty and limb structure. However, accumulating evidence from the 1960s, including North American and specimens, began challenging this polyphyletic arrangement by revealing distinct lineages for teiids and lacertids. Cladistic approaches in the 1970s and 1980s further transformed the classification, emphasizing monophyly through rigorous character analysis. Richard Estes's 1983 synthesis of morphological and fossil data redefined Scincomorpha as a more restricted, monophyletic clade sister to Anguimorpha, excluding teiids and lacertids while retaining core scincoid families based on synapomorphies like ventral osteoderms and specific caudal chevron positions. This shift, informed by enhanced fossil datasets, marked a transition from broad, similarity-based groupings to phylogeny-driven taxonomy.

Modern phylogeny

Modern molecular phylogenies, based on multi-locus DNA datasets, strongly support the of Scincomorpha as a major within , with bootstrap support values exceeding 95% across analyses. A landmark study by Pyron et al. (2013) utilized up to 12 nuclear and mitochondrial genes (totaling 12,896 bp) from 4,161 squamate species, recovering Scincomorpha as a robust monophyletic group with nodal support up to 100% in maximum likelihood trees. Similarly, Burbrink et al. (2020) analyzed 394 anchored hybrid enrichment loci from 289 species, confirming with high gene and site concordance factors (>50%) and bootstrap values >95%, highlighting the 's stability under genomic-scale sampling. Internally, Scincomorpha is structured around the core subclade Scincoidea, which encompasses Scincidae and its allies (including , Gerrhosauridae, and Xantusiidae), supported by Shimodaira-Hasegawa-like tests yielding values of 100. Within Scincoidea, Xantusiidae emerges as the basal lineage, with strong bootstrap support (e.g., 98%), forming a to the remaining families, while Scincidae represents the largest and most derived component, confirmed as monophyletic with support of 100. These relationships reflect adaptations to diverse ecological niches, with Xantusiidae's and nocturnal habits marking early divergences. Regarding external affinities, Scincomorpha is positioned as in multi-locus analyses, with high nodal support (e.g., 100%), placing it within the scleroglossan radiation after Dibamidae and Gekkota. Genomic advancements in the 2020s have further refined Scincomorpha's phylogeny, incorporating thousands of loci to resolve longstanding debates on scincid . Early molecular studies occasionally suggested due to limited sampling, but recent phylogenomic datasets, such as those in Burbrink et al. (2020), unequivocally affirm Scincidae's by integrating ultraconserved elements and whole-genome alignments, achieving near-complete resolution of internal branches with support >95%. A 2024 species-tree analysis of scincine , using multi-locus data from over 100 , corroborates this , estimating the family's origin around 98 million years ago and attributing past signals to incomplete lineage sorting rather than true non-. These updates underscore ' role in stabilizing Scincomorpha's tree, enabling precise evolutionary inferences without reliance on morphology alone.

Constituent families

Scincomorpha encompasses four extant families, all belonging to the superfamily Scincoidea, which together comprise over 1,900 of characterized by their scaly and varied body forms adapted to diverse environments. The largest family is Scincidae, containing 1,793 of skinks that exhibit remarkable morphological diversity, including limbed and limbless forms, and are distributed across all continents except , with major concentrations in the and . These skinks often feature smooth scales and elongated bodies, enabling burrowing or terrestrial lifestyles in forests, deserts, and grasslands. The family Cordylidae includes 68 species of girdled lizards, primarily found in sub-Saharan Africa, where they inhabit rocky terrains and exhibit defensive behaviors such as armadillo-like rolling and spiny tails for protection against predators. Gerrhosauridae consists of 39 species of plated lizards, also restricted to Africa and Madagascar, noted for their heavily armored scales and fossorial or semi-arboreal habits in savannas and woodlands. Xantusiidae comprises 38 species of night lizards, endemic to the Americas from southwestern United States to Central America, with viviparous reproduction and nocturnal activity in arid and forested habitats. Within Scincidae, taxonomic structure varies across classifications, but Hedges (2014) delineates 9 major clades treated as families or subfamilies, including Acontinae (26 species, limbless forms in and ), Egerniidae (58 species, robust-bodied in and New Guinea), Eugongylidae (418 species, diverse in Indo-Pacific islands), Lygosomidae (52 species, slender forms in and Australasia), Mabuyidae (190 species, fast-moving in tropics worldwide), Sphenomorphidae (546 species, widespread in and ), and others like Ateuchosaurinae and Brachymeles group, emphasizing geographic foci from tropics to Australasia. Extinct families of Scincomorpha, primarily from the era, include Paramacellodidae, which appeared in the and persisted until the , featuring generalized scincomorph skulls across and . Polyglyphanodontidae, known from the Early to , comprised herbivorous or insectivorous forms with multicusped teeth, distributed in , , and . Other extinct groups such as Adamisauridae, Gilmoreteiidae, and Mongolochamopidae also contributed to the clade's diversity during this period, with approximately 20 genera documented from fossil records spanning to deposits.

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