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A phylogeny and revised classification of Squamata, including 4161 species oflizards and snakes
BMC Evolutionary Biology 2013, 13:93 doi:10.1186/1471-2148-13-93
Robert Alexander Pyron ([email protected])Frank T Burbrink ([email protected])
John J Wiens ([email protected])
ISSN 1471-2148
Article type Research article
Submission date 30 January 2013
Acceptance date 19 March 2013
Publication date 29 April 2013
Article URL http://www.biomedcentral.com/1471-2148/13/93
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A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes
Robert Alexander Pyron1* * Corresponding author Email: [email protected]
Frank T Burbrink2,3 Email: [email protected]
John J Wiens4 Email: [email protected]
1 Department of Biological Sciences, The George Washington University, 2023 G St. NW, Washington, DC 20052, USA
2 Department of Biology, The Graduate School and University Center, The City University of New York, 365 5th Ave., New York, NY 10016, USA
3 Department of Biology, The College of Staten Island, The City University of New York, 2800 Victory Blvd., Staten Island, NY 10314, USA
4 Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ 85721-0088, USA
Abstract
Background
The extant squamates (>9400 known species of lizards and snakes) are one of the most diverse and conspicuous radiations of terrestrial vertebrates, but no studies have attempted to reconstruct a phylogeny for the group with large-scale taxon sampling. Such an estimate is invaluable for comparative evolutionary studies, and to address their classification. Here, we present the first large-scale phylogenetic estimate for Squamata.
Results
The estimated phylogeny contains 4161 species, representing all currently recognized families and subfamilies. The analysis is based on up to 12896 base pairs of sequence data per species (average = 2497 bp) from 12 genes, including seven nuclear loci (BDNF, c-mos, NT3, PDC, R35, RAG-1, and RAG-2), and five mitochondrial genes (12S, 16S, cytochrome b, ND2, and ND4). The tree provides important confirmation for recent estimates of higher-level squamate phylogeny based on molecular data (but with more limited taxon sampling), estimates that are very different from previous morphology-based hypotheses. The tree also includes many relationships that differ from previous molecular estimates and many that differ from traditional taxonomy.
Conclusions
We present a new large-scale phylogeny of squamate reptiles that should be a valuable resource for future comparative studies. We also present a revised classification of squamates at the family and subfamily level to bring the taxonomy more in line with the new phylogenetic hypothesis. This classification includes new, resurrected, and modified subfamilies within gymnophthalmid and scincid lizards and boid, colubrid, and lamprophiid snakes.
Keywords
Amphisbaenia, Lacertilia, Likelihood support measures, Missing data, Serpentes, Squamata, Phylogenetics, Reptiles, Supermatrices, Systematics
Background
Squamate reptiles (lizards, snakes, and amphisbaenians ["worm lizards"]) are among the most diverse radiations of terrestrial vertebrates. Squamata includes more than 9400 species as of December 2012 [1]. The rate of new species descriptions shows no signs of slowing, with a record 168 new species described in 2012 [1], greater than the highest yearly rates of the 18th and 19th centuries (e.g. 1758, 118 species; 1854, 144 species [1]). Squamates are presently found on every continent except Antarctica, and in the Indian and Pacific Oceans, and span many diverse ecologies and body forms, from limbless burrowers to arboreal gliders (summarized in [2-4]).
Squamates are key study organisms in numerous fields, from evolution, ecology, and behavior [3] to medicine [5,6] and applied physics [7]. They have also been the focus of many pioneering studies using phylogenies to address questions about trait evolution (e.g. [8,9]). Phylogenies are now recognized as being integral to all comparative studies of squamate biology (e.g. [10,11]). However, hypotheses about squamate phylogeny have changed radically in recent years [12], especially when comparing trees generated from morphological [13-15] and molecular data [16-20]. Furthermore, despite extensive work on squamate phylogeny at all taxonomic levels, a large-scale phylogeny (i.e. including thousands of species and multiple genes) has never been attempted using morphological or molecular data.
Squamate phylogenetics has changed radically in the last 10 years, revealing major conflicts between the results of morphological and molecular analyses [12]. Early estimates of squamate phylogeny [21] and recent studies based on morphological data [13-15,22] consistently supported a basal division between Iguania (including chameleons, agamids, and iguanids, sensu lato), and Scleroglossa, which comprises all other squamates (including skinks, geckos, snakes, and amphisbaenians). Within Scleroglossa, many phylogenetic analyses of morphological data have also supported a clade containing limb-reduced taxa, including various combinations of snakes, dibamids, amphisbaenians, and (in some analyses) limb-reduced skinks and anguids [13-15,19,22], though some of these authors also acknowledged that this clade was likely erroneous.
In contrast, recent molecular analyses have estimated very different relationships. Novel arrangements include placement of dibamids and gekkotans near the root of the squamate tree, a sister-group relationship between amphisbaenians and lacertids, and a clade (Toxicofera) uniting Iguania with snakes and anguimorphs within Scleroglossa [16-20,23,24]. These molecular results (and the results of combined morphological and molecular analyses) suggest that some estimates of squamate phylogeny based on morphology may have been misled, especially by convergence associated with adaptations to burrowing [19]. However, there have also been disagreements among molecular studies, such as placement of dibamids relative to gekkotans and other squamates and relationships among snakes, iguanians, and anguimorphs (e.g. [17,20]).
Analyses of higher-level squamate relationships based on molecular data have so far included relatively few (less than 200) species, and none have included representatives from all described families and subfamilies [17-20,23,24]. This limited taxon sampling makes existing molecular phylogenies difficult to use for broad-scale comparative studies, with some exceptions based on supertrees [10,11]. In addition, limited taxon sampling is potentially a serious issue for phylogenetic accuracy [25-28]. Thus, an analysis with extensive taxon sampling is critically important to test hypotheses based on molecular datasets with more limited sampling and to provide a framework for comparative analyses.
Despite the lack of a large-scale phylogeny across squamates, recent molecular studies have produced phylogenetic estimates for many of the major groups of squamates, including iguanian lizards [29-34], higher-level snake groups [35-37], typhlopoid snakes [38,39], colubroid snakes [40-46], booid snakes [47,48], scincid lizards [49-52], gekkotan lizards [53-60], teiioid lizards [61-64], lacertid lizards [65-69], and amphisbaenians [70,71]. These studies have done an outstanding job of clarifying the phylogeny and taxonomy of these groups, but many were limited in some ways by the number of characters and taxa that they sampled (and which were available at the time for sequencing).
Here, we present a phylogenetic estimate for Squamata based on combining much of the existing sequence data for squamate reptiles, using the increasingly well-established supermatrix approach [41,72-77]. We present a new phylogenetic estimate including 4161 squamate species. The dataset includes up to 12896 bp per species from 12 loci (7 nuclear, 5 mitochondrial). We include species from all currently described families and subfamilies. In terms of species sampled, this is 5 times larger than any previous phylogeny for any one squamate group [30,41], 3 times larger than the largest supertree estimate [11], and 25 times larger than the largest molecular study of higher-level squamate relationships [20]. While we did not sequence any new taxa specifically for this project, much of the data in the combined matrix were generated in our labs or from our previous collaborative projects [16,19,20,34,36,37,41,44,78-82], including thousands of gene sequences from hundreds of species (>550 species; ~13% of the total).
The supermatrix approach can provide a relatively comprehensive phylogeny, and uncover novel relationships not seen in any of the separate analyses in which the data were generated. Such novel relationships can be revealed via three primary mechanisms. First, different studies may have each sampled different species from a given group for the same genes, and combining these data may reveal novel relationships not apparent in the separate analyses. Second, different studies may have used different genetic markers for the same taxa, and combining these markers can dramatically increase character sampling, potentially revealing new relationships and providing stronger support for previous hypotheses. Third, even for
clades that were previously studied using complete taxon sampling and multiple loci, novel relationships may be revealed by including these lineages with other related groups in a large-scale phylogeny.
The estimated tree and branch-lengths should be useful for comparative studies of squamate biology. However, this phylogeny is based on a supermatrix with extensive missing data (mean = 81% per species). Some authors have suggested that matrices with missing cells may yield misleading estimates of topology, support, and branch lengths [83]. Nevertheless, most empirical and simulation studies have not found this to be the case, at least for topology and support [41,73,84,85]. Though fewer studies have examined the effects of missing data on branch lengths [44,86,87], these also suggest that missing data do not strongly impact estimates. Here, we test whether branch lengths for terminal taxa are related to their completeness.
In general, our results corroborate those of many recent molecular studies with regard to higher-level relationships, species-level relationships, and the monophyly, composition, and relationships of most families, subfamilies, and genera. However, our results differ from previous estimates for some groups, and reveal (or corroborate) numerous problems in the existing classification of squamates. We therefore provide a conservative, updated classification of extant squamates at the family and subfamily level based on the new phylogeny, while highlighting problematic taxonomy at the genus level, without making changes. The generic composition of all families and subfamilies under our revised taxonomy are provided in Appendix I.
We note dozens of problems in the genus-level taxonomy suggested by our tree, but we acknowledge in advance that we do not provide a comprehensive review of the previous literature dealing with all these taxonomic issues (this would require a monographic treatment). Similarly, we do not attempt to fix these genus-level problems here, as most will require more extensive taxon (and potentially character) sampling to adequately resolve.
Throughout the paper, we address only extant squamates. Squamata also includes numerous extinct species classified in both extant and extinct families, subfamilies, and genera. Relationships and classification of extinct squamates based on morphological data from fossils have been addressed by numerous authors (e.g. [14,15,19,22,88-93]). A classification based only on living taxa may create some problems for classifying fossil taxa, but these can be addressed in future studies that integrate molecular and fossil data [19,86].
Results
Supermatrix phylogeny
We generated the final tree (lnL = −2609551.07) using Maximum Likelihood (ML) in RAxMLv7.2.8. Support was assessed using the non-parametric Shimodaira-Hasegawa-Like (SHL) implementation of the approximate likelihood-ratio test (aLRT; see [94]). The tree and data matrix are available in NEXUS format in DataDryad repository 10.5061/dryad.82h0m and as Additional file 1: Data File S1. A skeletal representation of the tree (excluding several species which are incertae sedis) is shown in Figure 1. The full species-level phylogeny (minus the outgroup Sphenodon) is shown in Figures 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20, 21, 22, 23, 24, 25, 26, 27, 28. The analysis yields a generally well-
supported phylogenetic estimate for squamates (i.e. 70% of nodes have SHL values >85, indicating they are strongly supported). There is no relationship between proportional completeness (bp of non-missing data in species / 12896 bp of complete data) and branch length (r = −0.29, P = 0.14) for terminal taxa, strongly suggesting that the estimated branch lengths are not consistently biased by missing data.
Figure 1 Higher-level squamate phylogeny. Skeletal representation of the 4161-species tree from maximum-likelihood analysis of 12 genes, with tips representing families and subfamilies (following our taxonomic revision; species considered incertae sedis are not shown). Numbers at nodes are SHL values greater than 50%. The full tree is presented in Figures 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20, 21, 22, 23, 24, 25, 26, 27, 28, with panels indicated by bold italic letters.
Figure 2 Species-level squamate phylogeny. Large-scale maximum likelihood estimate of squamate phylogeny, containing 4161 species. Numbers at nodes are SHL values greater than 50%. A skeletal version of this tree is presented in Figure 1. Bold italic letters indicate figure panels (A). Within panels, branch lengths are proportional to expected substitutions per site, but the relative scale differs between panels.
Figure 3 Species-level squamate phylogeny continued (B).
Figure 4 Figure 5 Species-level squamate phylogeny continued (D) Species-level squamate phylogeny continued (D).
Figure 6 Species-level squamate phylogeny continued (E).
Figure 7 Species-level squamate phylogeny continued (F).
Figure 8 Species-level squamate phylogeny continued (G).
Figure 9 Species-level squamate phylogeny continued (H).
Figure 10 Species-level squamate phylogeny continued (I).
Figure 11 Species-level squamate phylogeny continued (J).
Figure 12 Species-level squamate phylogeny continued (K) Species-level squamate phylogeny continued (K).
Figure 13 Species-level squamate phylogeny continued (L).
Figure 14 Species-level squamate phylogeny continued (M).
Figure 15 Species-level squamate phylogeny continued (N).
Figure 16 Species-level squamate phylogeny continued (O).
Figure 17 Species-level squamate phylogeny continued (P).
Figure 18 Species-level squamate phylogeny continued (Q).
Figure 19 Species-level squamate phylogeny continued (R).
Figure 20 Species-level squamate phylogeny continued (S).
Figure 21 Species-level squamate phylogeny continued (T).
Figure 22 Species-level squamate phylogeny continued (U).
Figure 23 Species-level squamate phylogeny continued (V).
Figure 24 Species-level squamate phylogeny continued (W).
Figure 25 Species-level squamate phylogeny continued (X).
Figure 26 Species-level squamate phylogeny continued (Y).
Figure 27 Species-level squamate phylogeny continued (Z).
Figure 28 Species-level squamate phylogeny continued (AA).
Higher-level relationships
Our tree (Figure 1) is broadly congruent with most previous molecular studies of higher-level squamate phylogeny using both nuclear data and combined nuclear and mitochondrial data (e.g. [16-20]), providing important confirmation of previous molecular studies based on more limited taxon sampling. Specifically we support (Figure 1): (i) the placement of dibamids and gekkotans near the base of the tree (Figure 1A); (ii) a sister-group relationship between Scincoidea (scincids, cordylids, gerrhosaurids, and xantusiids; Figure 1B) and a clade (Episquamata; Figure 1C) containing the rest of the squamates excluding dibamids and gekkotans; (iii) Lacertoidea (lacertids, amphisbaenians, teiids, and gymnophthalmids; Figure 1D), and (iv) a clade (Toxicofera; Figure 1E) containing anguimorphs (Figure 1F), iguanians (Figure 1G), and snakes (Figure 1H) as the sister taxon to Lacertoidea.
These relationships are strongly supported in general (Figure 1), but differ sharply from most trees based on morphological data [13-15,19,22,95]. Nevertheless, many clades found in previous morphological taxonomies and phylogenies are also present in this tree in some form, including Amphisbaenia, Anguimorpha, Gekkota, Iguania, Lacertoidea (but including amphisbaenians), Scincoidea, Serpentes, and many families and subfamilies. In contrast, the relationships among these groups differ strongly between molecular analyses [17-20] and morphological analyses [14,15]. Our results demonstrate that this incongruence is not explained by limited taxon sampling in the molecular data sets. In fact, our species-level sampling is far more extensive than in any morphological analyses (e.g. [14,15]), by an order of magnitude.
We find that the basal squamate relationships are strongly supported in our tree. The family Dibamidae is the sister group to all other squamates, and Gekkota is the sister group to all squamates excluding Dibamidae (Figure 1), as in some previous studies (e.g. [16,18]). Other recent molecular analyses have also placed Dibamidae near the squamate root, but differed in placing it as either the sister taxon to all squamates excluding Gekkota [17], or the sister-group of Gekkota [19,20]. Our results also corroborate that the New World genus
Anelytropsis is nested within the Old World genus Dibamus [96], but the associated branches are weakly supported (Figure 2).
Gekkota
Within Gekkota, we corroborate both earlier morphological [97] and recent molecular estimates [55,56,59,98] in supporting a clade containing the Australian radiation of "diplodactylid" geckos (Carphodactylidae and Diplodactylidae) and the snakelike pygopodids (Figures 1, 2). As in previous studies [55], Carphodactylidae is the weakly supported sister group to Pygopodidae, and this strongly supported clade is the sister group to Diplodactylidae (Figures 1, 2). We recover clades within the former Gekkonidae that correspond to the strongly supported families Eublepharidae, Sphaerodactylidae, Phyllodactylidae, and Gekkonidae as in previous studies, and similar relationships among these groups [55-57,59,60,98-100].
Within Gekkota, we find evidence for non-monophyly of many genera. Many relationships among the New Caledonian diplodactylids are weakly supported (Figure 2), and there is apparent non-monophyly of the genera Rhacodactylus, Bavayia, and Eurydactylodes with respect to each other and Oedodera, Dierogekko, Paniegekko, Correlophus, and Mniarogekko [101]. In the Australian diplodactylids, Strophurus taenicauda is strongly supported as belonging to a clade that is only distantly related to the other sampled Strophurus species (Figure 2). The two species of the North African sphaerodactylid genus Saurodactylus are divided between the two major sphaerodactylid clades (Figure 3), but the associated branches are weakly supported. The South American phyllodactylid genus Homonota is strongly supported as being paraphyletic with respect to Phyllodactylus (Figure 3).
A number of gekkonid genera (Figure 4) also appear to be non-monophyletic, including the Asian genera Cnemaspis (sampled species divided into two non-sister clades), Lepidodactylus (with respect to Pseudogekko and some Luperosaurus), Gekko (with respect to Ptychozoon and Lu. iskandari), Luperosaurus (with respect to Lepidodactylus and Gekko), Mediodactylus (with respect to Pseudoceramodactylus, Tropiocolotes, Stenodactylus, Cyrtopodion, Bunopus, Crossobamon, and Agamura), and Bunopus (with respect to Crossobamon), and the African Afrogecko (with respect to Afroedura, Christinus, Cryptactites, and Matoatoa), Afroedura (with respect to Afrogecko, Blaesodactylus, Christinus, Geckolepis, Pachydactylus, Rhoptropus, and numerous other genera), Chondrodactylus (with respect to Pachydactylus laevigatus), and Pachydactylus (with respect to Chondrodactylus and Colopus). Many of these taxonomic problems in gekkotan families have been identified in previous studies (e.g. [59,99,102]), and extensive changes will likely be required to fix them.
Scincoidea
We strongly support (SHL = 100; Figures 1, 5, 6, 7, 8, 9, 10) the monophyly of Scincoidea (Scincidae, Xantusiidae, Gerrhosauridae, and Cordylidae), as in other recent studies [16-20]. All four families are strongly supported (Figures 5, 6, 7, 8, 9, 10). A similar clade is also recognized in morphological phylogenies [14], though without Xantusiidae in some [13].
Within the New World family Xantusiidae, we corroborate previous analyses [103,104] that found strong support for a sister-group relationship between Xantusia and Lepidophyma, excluding Cricosaura (Figure 5). These relationships support the subfamily Cricosaurinae for
Cricosaura [105]. We also recognize Xantusiinae for the North American genus Xantusia and Lepidophyminae for the Central American genus Lepidophyma [106,107].
Within the African and Madagascan family Gerrhosauridae (Figure 5), the genus Gerrhosaurus is weakly supported as being paraphyletic with respect to the clade comprising Tetradactylus + Cordylosaurus, with G. major placed as the sister group to all other gerrhosaurids. Within Cordylidae (Figure 5), we use the generic taxonomy from a recent phylogenetic analysis and re-classification based on multiple nuclear and mitochondrial genes [108]. This classification broke up the non-monophyletic Cordylus [109] into several smaller genera, and we corroborate the non-monophyly of the former Cordylus and support the monophyly of the newly recognized genera (Figure 5). We find support the distinctiveness of Platysaurus (Figure 5) and recognition of the subfamily Platysaurinae [108].
We find strong support (SHL = 100) for the monophyly of Scincidae (Figure 6) as in previous studies (e.g. [20,50,51]). We find strong support for the basal placement of the monophyletic subfamily Acontiinae (Figure 6), as found in some previous studies (e.g. [20,51]) but not others (e.g. [50]). Similar to earlier studies, we find that the subfamily Scincinae sensu [110] is non-monophyletic, as Feylininae is nested within Scincinae (also found in [20,50,51,111]). Based on these results, synonymizing Feylininae with Scincinae produces a monophyletic Scincinae (SHL = 97), which is then sister to a monophyletic Lygosominae (SHL = 100 excluding Ateuchosaurus; see below) with 94% SHL support (Figures 6, 7, 8, 9, 10). This yields a new classification in which all three subfamilies (Acontiinae, Lygosominae, Scincinae) are strongly supported. Importantly, these definitions approximate the traditional content of the three subfamilies [50,110], except for recognition of Feylininae.
We note that a recent revision of the New World genus Mabuya introduced a nontraditional family-level classification for Scincidae [112]. These authors divided Scincidae into seven families: Acontiidae, Egerniidae, Eugongylidae, Lygosomidae, Mabuyidae, Scincidae and Sphenomorphidae. However, there was no phylogenetic need for considering these clades as families (or another rank below family), since the family Scincidae is clearly monophyletic, based on our results and others (see above). Thus, their new taxonomy changes the long-standing definition of Scincidae unnecessarily (see [113]). Furthermore, these changes were done without defining the full content (beyond a type genus) of any of these families other than Scincidae (the former Scincinae + Feylininae), and Acontiidae (the former Acontiinae).
Most importantly, the new taxonomy proposed by these authors [112] is at odds with the phylogeny estimated here, with respect to the familial and subfamilial classification of >1000 skink species (Figures 6, 7, 8, 9, 10). For instance, Sphenomorphus stellatus is found in a strongly supported clade containing Lygosoma (presumably Lygosomidae; Figure 10), which is separate from the other clade (presumably Sphenomorphidae) containing the other sampled Sphenomorphus species (Figure 7; but note that these Sphenomorphus species are divided among several subclades within this latter clade). An additional problem is that Egernia, Lygosoma, and Sphenomorphus are the type genera of Egerniidae, Lygosomidae, and Sphenomorphidae, but are paraphyletic as currently defined (Figures 7, 8, 9, 10), leading to further uncertainty in the content and definition of these putative families.
Furthermore, Mabuyidae apparently refers to the clade (Figure 9) containing Chioninia, Dasia, Mabuya, Trachylepis, with each of these genera placed in its own subfamily (Chioniniinae, Dasiinae, Mabuyinae, and Trachylepidinae). However, several other genera are strongly placed in this group, such as Eumecia, Eutropis, Lankaskincus, and Ristella
(Figure 9). These other genera cannot be readily fit into these subfamilial groups (i.e. they are not the sister group of any genera in those subfamilies), and Trachylepis is paraphyletic with respect to Eumecia, Chioninia, and Mabuya (Figure 9). Also, we find that Emoia is divided between clades containing Lygosoma (Lygosomidae) and Eugongylus (Eugongylidae), and many of these relationships have strong support (Figure 10). Finally, Ateuchosaurus is apparently not accounted for in their classification, and here is weakly placed as the sister-group to a clade comprising their Sphenomorphidae, Egerniidae, Mabuyidae, and Lygosomidae (i.e. Lygosominae as recognized here; Figures 7, 8, 9, 10).
These authors [112] argued that a more heavily subdivided classification for skinks may be desirable for facilitating future taxonomic revisions and species descriptions. However, this classification seems likely to only exacerbate existing taxonomic problems (e.g. placing congeneric species in different families without revising the genus-level taxonomy). Here, we retain the previous definition of Mabuya (restricted to the New World clade; sensu [114]), and we support the traditional definitions of Scincidae, Acontiinae, Scincinae (but including Feylininae), and Lygosominae (Figures 6, 7, 8, 9, 10; note that we leave Ateuchosaurus as incertae sedis). The other taxonomic issues in Scincidae identified here and elsewhere should be resolved in future studies. Our phylogeny provides a framework in which these analyses can take place (i.e. identifying major subclades within skinks), which we think may be more useful than a classification lacking clear taxon definitions.
Of the 133 scincid genera [1], we can assign the 113 sampled in our tree to one of the three subfamilies in our classification (Acontiinae, Lygosominae, and Scincinae; Appendix I). We place 19 of the remaining genera into one of the three subfamilies based on previous classifications (e.g. [110]), with Ateuchosaurus as incertae sedis in Scincidae. Below, we review the non-monophyletic genera in our tree. Many of these problems have been reported by previous authors [51,111,115-118], and for brevity we do not distinguish between cases reported in previous studies, and potentially new instances found here.
Within Acontiinae, we find that the two genera are both strongly supported as monophyletic (Figure 6). Within Scincinae, many genera are now strongly monophyletic (thanks in part to the dismantling of Eumeces; [49,50,119]), but some problems remain (Figure 6). The genera Scincus and Scincopus are strongly supported as being nested inside of the remaining Eumeces. Among Malagasy scincines (see [49,120]), Pseudacontias is nested inside Madascincus, and the genera Androngo, Pygomeles, and Voeltzkowia are all nested in Amphiglossus (Figure 6).
We also find numerous taxonomic problems with lygosomines (Figures 7, 8, 9, 10). Species of Sphenomorphus are widely dispersed among other lygosomine genera. The genus Tropidophorus is paraphyletic with respect to a clade containing many other genera (Figure 7). The sampled species of Asymblepharus are only distantly related to each other, including one species (A. sikimmensis) nested inside of Scincella (Figure 7). The genus Lipinia is polyphyletic, with one species (L. vittigera) strongly placed as the sister taxon to Isopachys, and with two other species (L. pulchella and L. noctua) forming a well-supported clade that also includes Papuascincus (Figure 7).
Among Australian skinks, the genus Eulamprus is polyphyletic with respect to Nangura, Calyptotis, Gnypetoscincus, Coggeria, Coeranoscincus, Ophioscincus, Saiphos, Anomalopus, Eremiascincus, Hemiergis, Glaphyromorphus, Notoscincus, Ctenotus, and Lerista, and most of the relevant nodes are strongly supported (Figure 8). The genera Coeranoscincus and
Ophioscincus are polyphyletic with respect to each other and to Saiphos and Coggeria (Figure 8). The genus Glaphyromorphus is paraphyletic with respect to a clade of Eulamprus (Figure 8). The genus Egernia is paraphyletic with respect to Bellatorias (which is paraphyletic with respect to Egernia and Lissolepis) and Lissolepis, although many of the relevant nodes are not strongly supported (Figure 9). The genera Cyclodomorphus and Tiliqua are paraphyletic with respect to each other (Figure 9).
Among other lygosomines, Trachylepis is non-monophyletic [121], with two species (T. aurata and T. vittata) that fall outside the strongly supported clade containing the other species (Figure 9). The latter clade is weakly supported as the sister group to a clade containing Chioninia, Eumecia, and Mabuya. In Mabuya, a few species (M. altamazonica, M. bistriata, and M. nigropuncata) have unorthodox placements within a monophyletic Mabuya, potentially due to uncertain taxonomic assignment of specimens by previous authors [51,122]. The genus Lygosoma is paraphyletic with respect to Lepidothyris and Mochlus, and many of the relevant nodes are strongly supported (Figure 10). Among New Caledonian skinks, the genus Lioscincus is polyphyletic with respect to Marmorosphax, Celatiscincus, and Tropidoscincus, and both Lioscincus and Tropidoscincus are paraphyletic with respect to, Kanakysaurus, Lacertoides, Phoboscincus, Sigaloseps, Tropidoscincus, Graciliscincus, Simiscincus, and Caledoniscincus, with strong support for most relevant nodes (Figure 10). The genera Emoia and Bassiana are massively polyphyletic and divided across multiple lygosomine clades (Figure 10). The genus Lygisaurus appears to be nested inside of Carlia, although many of the relevant branches are only weakly supported (Figure 10).
Lacertoidea
Within Lacertoidea (Figure 1), we corroborate recent molecular analyses (e.g. [16,17,19,20]) and morphology-based phylogenies and classifications (e.g. [13,15]) in supporting the clade including the New World families Gymnophthalmidae and Teiidae (Figure 11). Within a weakly supported Teiidae (Figure 11), the subfamilies Tupinambinae and Teiinae are each strongly supported as monophyletic, as in previous studies [61]. In Tupinambinae, Callopistes is the sister group to a clade containing Tupinambis, Dracaena, and Crocodilurus. The clade Dracaena + Crocodilurus is nested within Tupinambis, and the associated clades have strong support (Figure 11). We find that the teiine genera Ameiva and Cnemidophorus are non-monophyletic (Figure 11), interdigitating with each other and the monophyletic genera Aspidoscelis, Dicrodon (monotypic), and Kentropyx, as in previous phylogenies [62].
A recent study [123] proposed a re-classification of the family Teiidae based on analysis of 137 morphological characters for 101 terminal species (with ~150 species in the family). Those authors erected several new genera and subfamilies in an attempt to deal with the apparent non-monophyly of currently recognized taxa in their tree. However, in our tree, some of these new taxa conflict strongly with the phylogeny or are rendered unnecessary. First, they recognize Callopistinae as a distinct subfamily for Callopistes, arguing that failure to do so would produce a taxonomy inconsistent with teiid phylogeny. However, we find strong support for Callopistes in its traditional placement as part of Tupinambinae (Figure 11), and this change is thus not needed based on our results. The genus Ameiva is paraphyletic under traditional definitions [62]. In our tree, their conception of Ameiva is also non-monophyletic, with species found in three distinct clades (Figure 11). We also find non-monophyly of many of their species groups within Ameiva, including the ameiva, bifrontata, dorsalis, and erythrocephala groups (Figure 11). Their genera Aurivela (Cnemidophorus
longicaudus) and Contomastyx (Cnemidophorus lacertoides) are strongly supported as sister taxa in our tree, and are nested within Ameiva in their erythrocephala species group, along with Dicrodon (Figure 11).
On the positive side, many of the genera they recognize are monophyletic and are not nested in other genera in our tree, including their Ameivula (Cnemidophorus ocellifer), Aspidoscelis (unchanged from previous definitions), Cnemidophorus (excluding C. ocellifer, C. lacertoides, and C. longicaudus), Holcosus (Ameiva undulata, A. festiva, and A. quadrilineatus), Kentropyx (unchanged from previous definitions), Salvator (Tupinambis rufescens, T. duseni, and T. merianae), and Teius (unchanged from previous definitions). We did not sample Ameiva edracantha (their Medopheos).
Given our results, major taxonomic rearrangements within Teiidae seem problematic at present, especially with the extensive paraphyly of many traditional and re-defined teiid genera, the lack of strong resolution of many of these relationships based on molecular and morphological data, and incomplete taxon sampling in all studies so far. Thus, we provisionally retain the traditional taxonomy of Teiidae, pending additional data and analyses. However, we note that Ameiva, Cnemidophorus, and Tupinambis are clearly non-monophyletic based on both our results and those of recent authors [123], and will require taxonomic changes in the future. We anticipate that many of these newly proposed genera [123] will be useful in such revisions.
We find strong support (SHL = 100) for monophyly of Gymnophthalmidae (Figure 11). Within Gymnophthalmidae, we find strong support for the monophyly of the previously recognized subfamilies [63,64,124], with the exception of Cercosaurinae (Figure 11). Previous researchers considered the genus Bachia a distinct tribe (Bachiini) within Cercosaurinae, based on a poorly supported sister-group relationship with the tribe Cercosaurini [63,64]. Here, we find a moderately well supported relationship (SHL = 84) between Bachia and Gymnophthalminae + Rhachisaurinae, and we find that this clade is only distantly related to other Cercosaurinae. Therefore, we restrict Cercosaurinae to the tribe Cercosaurini, and elevate the tribe Bachiini [64] to the subfamily level. The subfamily Bachiinae contains only the genus Bachia (Figure 11), identical in content to the previously recognized tribe [64]. Within Cercosaurinae, we find that the genus Petracola is nested within Proctoporus (Figure 11). In Ecpleopinae, Leposoma is divided into two clades, separated by Anotosaura, Colobosauroides, and Arthrosaura (Figure 11), and many of the relevant nodes are very strongly supported. These issues should be addressed in future studies.
Our results show strong support for a clade uniting Lacertidae and Amphisbaenia (Lacertoidea; Figure 1), as in many previous studies [16-20,23]. We also find strong support for monophyly of amphisbaenians (SHL = 99), in contrast to some molecular analyses [19,20]. Relationships among amphisbaenian families (Figure 12) are generally strongly supported and similar to those in earlier molecular studies (Figure 12), including the placement of the New World family Rhineuridae as sister group to all other amphisbaenians [70,71,125]. The family Cadeidae is placed as the sister-group to Amphisbaenidae + Trogonophiidae (Figures 1 and Figure 12) with weak support, but has been placed with Blanidae in previous studies, with strong support but less extensive taxon sampling [125,126].
We find strong support for monophyly of the Old World family Lacertidae (Figure 13). Within Lacertidae, branch support for the monophyly of most genera and for the subfamilies Gallotiinae and Lacertinae is very high (Figure 13). However, we find that relationships among many genera are poorly supported, as in previous studies [65,67,68]. Our results (Figure 13) also indicate that several lacertid genera are non-monophyletic with strong support for the associated nodes, including Algyroides (paraphyletic with respect to Dinarolacerta), Ichnotropis (paraphyletic with respect to Meroles), Meroles (paraphyletic with respect to Ichnotropis), Nucras (polyphyletic with respect to several genera, including Pedioplanis, Poromera, Latastia, Philocortus, Pseuderemias, and Heliobolus), and Pedioplanis (paraphyletic with respect to Nucras).
Higher-level phylogeny of Toxicofera
We find strong support (SHL = 96) for monophyly of Toxicofera (Anguimorpha, Iguania, and Serpentes; Figure 1), and moderate support for a sister-group relationship between Iguania and Anguimorpha (SHL = 79). Relationships among Anguimorpha, Iguania, and Serpentes were weakly supported in some Bayesian and likelihood analyses [16-19], but strongly supported in others [20]. We further corroborate previous studies in also placing Anguimorpha with Iguania [16-20]. In contrast, some other studies have placed anguimorphs with snakes as the sister group to iguanians [127,128].
Anguimorpha
Our hypothesis for family-level anguimorphan relationships (Figures 1, 14) is generally similar to that of other recent studies [17,19,20,129], and is strongly supported. Our results differ from some analyses based only on morphology, which place Anguidae near the base of Anguimorpha [130]. Here, Shinisauridae is strongly supported as the sister taxon to a well-supported clade of Varanidae + Lanthanotidae (Figures 1, 14). Varanid relationships are similar to previous estimates (e.g. [131]). Xenosauridae is here strongly supported as the sister-group to a strongly supported clade containing Helodermatidae and the strongly supported Anniellidae + Anguidae clade (Figures 1, 14). However, previous molecular analyses have placed Helodermatidae as the sister to Xenosauridae (Anniellidae + Anguidae), typically with strong support [16,17,19,20].
Within Anguidae (Figure 14), our phylogeny indicates non-monophyly of genera within every subfamily, including Diploglossinae (Diploglossus and Celestus are strongly supported as paraphyletic with respect to each other and to Ophiodes), Anguinae (Ophisaurus is strongly supported as paraphyletic with respect to Anguis, Dopasia, and Pseudopus), and Gerrhonotinae (Abronia and Mesaspis are non-monophyletic, and Coloptychon is nested inside Gerrhonotus). Some of these problems were not reported previously (e.g. Coloptychon, Abronia, and Mesaspis), due to incomplete taxon sampling in previous studies [129,132,133], but relationships within Gerrhonotinae are under detailed investigation by other researchers, so these issues are likely to be resolved in the near future.
Iguania
We find strong support (SHL = 100) for the monophyly of Iguania (Figure 1). This clade is strongly supported by nuclear data [16,17,19,20], but an apparent episode of convergent molecular evolution in several mitochondrial genes has seemingly misled some analyses of mtDNA, leading to weak support for Iguania [134], or even separation of the acrodonts and
pleurodonts [17,135] in previous studies. Within Iguania (Figure 1), we find strong support (SHL = 100) for a sister-group relationship between Chamaeleonidae and Agamidae (Acrodonta), and for a clade of mostly New World families (Pleurodonta; SHL = 100).
We find strong support for the monophyly of Chamaeleonidae and the subfamily Chamaeleoninae, and weak support for the paraphyly of Brookesiinae (Figures 1, 15). The sampled species of the Brookesia nasus group appear as the sister group to all other chamaeleonids (the latter clade weakly supported) as found by some previous authors [136], though other studies have recovered a monophyletic Brookesia [137,138]. Within Chamaeleoninae (Figure 15), we find strong support for the monophyly of most genera and species-level relationships. However, we find strong support for the non-monophyly of Calumma, with some species strongly placed with Chamaeleo, others strongly placed with Rieppeleon, and a third set weakly placed with Nadzikambia + Rhampoleon. While non-monophly of Calumma has also been found in previous studies [138], a recent study strongly supports monophyly of Brookesia and weakly supports monophyly of Calumma [139].
Monophyly of Agamidae is strongly supported (Figure 16; SHL = 100), contrary to some previous estimates [15,31]. Most relationships among agamid subfamilies and genera are strongly supported (Figure 16), and largely congruent with earlier studies [17,29,34,140]. There are some differences with earlier studies. For example, previous studies based on 29–44 loci [20,29,34] placed Hydrosaurinae as sister to Amphibolurinae + (Agaminae + Draconinae) with strong support, whereas we place Hydrosaurinae as the sister-group to Amphibolurinae with weak support. Other authors [140] placed Leiolepiedinae with Uromastycinae, but we (and most other studies) place Uromastycinae as the sister group to all other agamids.
Our phylogeny indicates several taxonomic problems within amphibolurine agamids (Figure 16). The genera Moloch and Chelosania render Hypsilurus paraphyletic, although the support for the relevant clades is weak. The species Lophognathus gilberti is placed in a strongly supported clade with Chlamydosaurus and some Amphibolurus (including the type species, A. muricatus), a clade that is not closely related to the other Lophognathus. Many of these taxonomic problems were also noted by previous authors [141].
Within agamine agamids (Figure 16), most relationships are well supported and monophyly of all sampled genera is strongly supported. In contrast, within draconine agamids (Figure 16), many intergeneric relationships are weakly supported, and some genera are non-monophyletic (Figure 16; see also [142]), including Gonocephalus (G. robinsonii is only distantly related to other Gonocephalus) and Japalura (with species distributed among three distantly related clades, including one allied with Ptyctolaemus, another with G. robinsonii, and a third with Pseudocalotes).
Recent authors suggested dividing Laudakia into three genera (Stellagama, Paralaudakia, and Laudakia) based on a non-phylogenetic analysis of morphology [143]. Here, Laudakia (as previously defined) is strongly supported as monophyletic (Figure 16), and this change is not necessitated by the phylogeny. Similarly, based on genetic and morphological data, recent authors [144] suggested resurrecting the genus Saara for the basal clade of Uromastyx (U. asmussi, U. hardwickii, and U. loricata). However, Uromastyx (as previously defined) is strongly supported as monophyletic in our results (Figure 16) and in those of the recent revision [144], and this change is not needed. We therefore retain Laudakia and Uromastyx as previously defined, to preserve taxonomic stability in these groups [113]. We note that recent
studies have also begun to revise species limits in other groups such as Trapelus [145], and taxa such as T. pallidus (Figure 16) may represent populations within other species.
Within Pleurodonta we generally confirm the monophyly and composition of the clades that were ranked as families (or subfamilies) within the group (e.g. Phrynosomatidae, Opluridae, Leiosauridae, Leiocephalidae, and Corytophanidae; Figures 1, 17, 18, 19) based on previous molecular studies [31,33,34] and earlier morphological studies [146,147].
One important exception is the previously recognized Polychrotidae. Our results confirm that Anolis and Polychrus are not sister taxa (Figures 1, 18, 19), as also found in some previous molecular studies [31,33,34], but not others [20,29]. Our results provide strong support for non-monophyly of Polychrotidae, placing Polychrus with Hoplocercidae (SHL = 99) and Anolis with Corytophanidae (SHL = 99; the latter also found by [34]). Recent analyses placing Anolis with Polychrus showed only weak support for this relationship [20,29], despite many loci (30–44). We support continued recognition of Dactyloidae for Anolis and Polychrotidae for Polychrus [34], based on a limited number of loci but extensive taxon sampling. We note that these families are still monophlyetic, even if they prove to be sister taxa.
Interestingly, our results for relationships among pleurodont families differ from most previous studies, and are surprisingly well-supported in some cases by SHL values (but see below). In previous studies, many relationships among pleurodont families were poorly supported by Bayesian posterior probabilities and by parsimony and likelihood bootstrap values, though typically sampling fewer taxa or characters [17,31,33,148-151]. Studies including 29 nuclear loci found strong concatenated Bayesian support for many relationships but weak support from ML bootstrap analyses for many of the same relationships [34]. The latter pattern (typically weak ML support) was also found in an analysis including those same 29 loci and mitochondrial data for >150 species [29]. We also find a mixture of strongly and weakly supported clades, but with many relationships that are incongruent with these previous studies. First, we find that Tropiduridae is weakly supported as the sister group to all other pleurodonts (also found by [29]), followed successively (Figures 1, 17, 18, 19) by Iguanidae, Leiocephalidae, Crotaphytidae + Phrynosomatidae, Polychrotidae + Hoplocercidae, and Corytophanidae + Dactyloidae.
The relatively strong support for the clades Crotaphytidae + Phrynosomatidae (SHL = 87), Polychrotidae + Hoplocercidae (SHL = 99), and Corytophanidae + Dactyloidae (SHL = 99) is largely unprecedented in previous studies (although Corytophanidae + Dactyloidae is strongly supported in some Bayesian analyses [34]). As in many previous analyses, deeper relationships among the families remain weakly supported. We also find a strongly supported clade containing Liolaemidae, Opluridae, and Leiosauridae (SHL = 95), with Opluridae + Leiosauridae also strongly supported (SHL = 99). Both clades have also been found in previous studies [149,151], including studies based on 29 or more nuclear loci [20,29,34].
We note that previous studies have shown strong support for some pleurodont relationships (e.g. basal placement of phrynosomatids; see [34]), only to be strongly overturned with additional data [20,29]. Therefore, the relationships found here should be taken with some caution (even if strongly supported), with the possible exception of the recurring clade of Liolaemidae + (Opluridae + Leiosauridae).
All pleurodont families are strongly supported as monophyletic (SHL > 85). Within the pleurodont families, our results generally support the current generic-level taxonomy (Figures 17, 18, 19). However, there are some exceptions. Within Tropiduridae (Figure 17), Tropidurus is paraphyletic with respect to Eurolophosaurus, Strobilurus, Uracentron, and Plica. Within Opluridae (Figure 18), the monotypic genus Chalarodon renders Oplurus paraphyletic. Two leiosaurid genera are also problematic (Figure 18). In Enyaliinae, Anisolepis is paraphyletic with respect to Urostrophus, and this clade is nested within Enyalius. In Leiosaurinae, Pristidactylus is rendered paraphyletic by Leiosaurus and Diplolaemus (Figure 18).
Within Dactyloidae, a recent study re-introduced a more subdivided classification of anoles [152], an issue that has been debated extensively in the past [153-156]. Our results support the monophyly of all the genera recognized by recent authors [152], including Anolis, Audantia, Chamaelinorops, Dactyloa, Deiroptyx, Norops, and Xiphosurus (see Figure 19). However, since Anolis is monophyletic as previously defined, we retain that definition here (including the seven listed genera) for continuity with the recent literature [113,157].
Serpentes
Relationships among the major serpent groups (Figure 1) are generally similar to other recent studies [20,35,36,38,41,44,47,158-160]. We find that the blindsnakes, Scolecophidia (Figures 1, 20) are not monophyletic, as in previous studies [19,20,36,44,159,160]. Similar to some previous studies [44,159], our data weakly place Anomalepididae as the sister taxon to all snakes, and the scolecophidian families Gerrhopilidae, Leptotyphlopidae, Typhlopidae, and Xenotyphlopidae as the sister-group to all other snakes excluding Anomalepididae (Figures 1, 20). Previous studies have also placed Anomalepididae as the sister-group to all non-scolecophidian snakes [19,20,35,36,158,160], in some cases with strong support [20]. Although it might appear that recent analyses of scolecophidian relationships [38] support monophyly of Scolecophidia (e.g. Figure 1 of [38]), the tree including non-snake outgroups from that study shows weak support for placing anomalepidids with alethinophidians, as in other studies [19,20,36,160].
We follow recent authors [38] in recognizing Xenotyphlopidae (strongly placed as the sister taxon of Typhlopidae) and Gerrhopilidae (strongly placed as the sister group of Xenotyphlopidae + Typhlopidae) as distinct families (Figure 20). Leptotyphlopidae is strongly supported as the sister group of a clade comprising Gerrhopilidae, Xenotyphlopidae, and Typhlopidae (Figure 20). As in previous studies [38,44], we find strong support for non-monophyly of several typhlopid genera (Afrotyphlops, Austrotyphlops, Ramphotyphlops, Letheobia, and Typhlops; Figure 20). There are also undescribed taxa (e.g. Typhlopidae sp. from Sri Lanka; [44]) of uncertain placement within this group (Figure 20). The systematics of typhlopoid snakes will thus require extensive revision in the future, with additional taxon and character sampling.
Within Alethinophidia (SHL = 100), Aniliidae is strongly supported (SHL = 98) as the sister taxon of Tropidophiidae (together comprising Anilioidea), and all other alethinophidians form a strongly supported sister group to this clade (SHL = 97; Figures 1, 21). The enigmatic family Xenophidiidae is weakly placed as the sister-group to all alethinophidians exclusive of Anilioidea (Figures 1, 21). The family Bolyeriidae is weakly placed as the sister-group to pythons, boas, and relatives (Booidea), which are strongly supported (SHL = 88).
Relationships in this group are generally consistent with other recent molecular studies [20,35-37,47,159].
Relationships among other alethinophidians are a mixture of strongly and weakly supported nodes (Figures 1, 21). We find strong support (SHL = 100) for a clade containing Anomochilidae + Cylindrophiidae + Uropeltidae. This clade of three families is strongly supported (SHL = 89) as the sister taxon to Xenopeltidae + (Loxocemidae + Pythonidae). Together, these six families form a strongly supported clade (SHL = 89; Figures 1, 21) that is weakly supported as the sister group to the strongly supported clade of Boidae + Calabariidae. Within the clade of Anomochilidae, Cylindrophiidae, and Uropeltidae (Figure 21), we weakly place Anomochilus as the sister group to Cylindrophiidae [44], in contrast to previous studies which placed Anomochilus within Cylindrophis [161]. However, support for monophyly of Cylindrophis excluding Anomochilus is weak (Figure 21). As in previous studies [44,162], we find several taxonomic problems within Uropeltidae (Figure 21). Specifically, Rhinophis and Uropeltis are paraphyletic with respect to each other and to Pseudotyphlops. The problematic taxa are primarily Sri Lankan [44], and forthcoming analyses will address these issues.
Within Pythonidae (Figure 21), the genus Python is the sister group to all other genera. Some species that were traditionally referred to as Python (P. reticulatus and P. timoriensis) are instead sister to an Australasian clade consisting of Antaresia, Apodora, Aspidites, Bothrochilus, Leiopython, Liasis, and Morelia (Figure 21). These taxa (P. reticulatus and P. timoriensis) have been referred to as Broghammerus, a name originating from an act of "taxonomic vandalism" (i.e. an apparently intentional attempt to disrupt stable taxonomy) in a non-peer reviewed organ without data or analyses [163,164]. However, this name was, perhaps inadvertently, subsequently used by researchers in peer-reviewed work [165] and has entered into somewhat widespread usage [1]. This name should be ignored and replaced with a suitable substitute. Within the Australasian clade (Figure 21), Morelia is paraphyletic with respect to all other genera, and Liasis is non-monophyletic with respect to Apodora, although many of the relevant relationships are weakly supported.
Within Boidae (Figure 21), our results and those of other recent studies [20,36,47,48,150,166] have converged on estimated relationships that are generally similar to each other but which differ from traditional taxonomy [167]. However, the classification has yet to be modified to reflect this, and we rectify this situation here. We find that Calabariidae is nested within Boidae [150], but this is poorly supported, and contrary to most previous studies [47,48]. While Calabaria has been classified as an erycine boid in the past, this placement is strongly rejected here and in other studies [47,48]. If the current placement of Calabaria is supported in the future, it would require recognition as the subfamily Calabariinae.
The Malagasy boine genera Acrantophis and Sanzinia are placed as the sister taxa to a weakly-supported clade containing Calabariidae and a strongly supported clade (SHL = 99) comprising the currently recognized subfamilies Erycinae, Ungaliophiinae, and other boines (Figure 21). Regardless of the position of Calabariidae, this placement of Malagasy boines renders Boinae paraphyletic. We therefore resurrect the subfamily Sanziniinae [168] for Acrantophis and Sanzinia. This subfamily could be recognized as a distinct family if future studies also support placement of this clade as distinct from other Boidae + Calabariidae.
The genera Lichanura and Charina are currently classified as erycines [1], but are strongly supported as the sister group to Ungaliophiinae, as in previous studies [20,36,47,166]. We expand Ungaliophiinae to include these two genera (Figure 21), rather than erect a new subfamily for these taxa. The subfamily Ungaliophiinae is placed as the sister group to a well-supported clade (SHL = 87) containing the rest of the traditionally recognized Erycinae and Boinae. We restrict Erycinae to the Old World genus Eryx.
The genus Candoia (Boinae) from Oceania and New Guinea [1], is placed as the sister taxon to a moderately supported clade (SHL = 83) consisting of Erycinae (Eryx) and the remaining genera of Boinae (Boa, Corallus, Epicrates, and Eunectes). To solve the non-monophyly of Boinae with respect to Erycinae (due to Candoia), we place Candoia in a new subfamily (Candoiinae, subfam. nov.; see Appendix I). Boinae then comprises the four Neotropical genera that have traditionally been classified in this group (Boa, Corallus, Epicrates, and Eunectes). We acknowledge that non-monophyly of Boinae could be resolved in other ways (e.g. expanding it to include Erycinae). However, our taxonomy maintains the traditionally used subfamilies Boinae, Erycinae, and Ungaliophiinae, modifies them to reflect the phylogeny, and recognizes the phylogenetically distinct boine clades as separate subfamilies (Candoiinae, Sanziniinae). Within Boinae, Eunectes renders Epicrates paraphyletic, but this is not strongly supported see also ([48]).
Our results for advanced snakes (Caenophidia) are generally similar to those of other recent studies [41,42,169], and will only be briefly described. However, in contrast to most recent studies [20,36,41,42,81,159,160], Acrochordidae is here strongly placed (SHL = 95) as the sister group to Xenodermatidae. This clade is then the sister group to the remaining Colubroidea, which form a strongly supported clade (SHL = 100; Figures 1, 22). This relationship has been found in some previous studies [169,170], and was hypothesized by early authors [171]. Further evidence will be required to resolve this conclusively. Analyses based on concatenation of 20–44 loci do not support this grouping [20,36], though preliminary species-tree analyses of >400 loci do (Pyron et al., in prep.). Relationships in Pareatidae are similar to recent studies [172], and the group is strongly placed as the sister taxon to colubroids excluding xenodermatids (SHL = 100; Figures 1 , 22), as in most recent analyses (e.g. [41,43,44]).
The family Viperidae is the sister group to all colubroids excluding xenodermatids and pareatids (Figure 1), as in other recent studies. The family Viperidae is strongly supported (Figure 22), as is the subfamily Viperinae, and the sister-group relationship between Azemiopinae and Crotalinae (SHL = 100). Our results generally support the existing generic-level taxonomy within Viperinae (Figure 22). However, we recover a strongly supported clade within Viperinae consisting of Daboia russelii, D. palaestinae, Macrovipera mauritanica, and M. deserti (Figure 22), as in previous studies [173]. We corroborate previous suggestions that these taxa be included in Daboia [174], though this has not been widely adopted [1]. The other Macrovipera species (including the type species) remain in that genus (Figure 22).
Within Crotalinae (Figure 22), a number of genera appear to be non-monophyletic. The species Trimeresurus gracilis is strongly supported as the sister taxon to Ovophis okinavensis and distantly related to other Trimeresurus, whereas the other Ovophis are strongly placed as the sister group to Protobothrops. A well-supported clade (SHL = 90) containing Atropoides picadoi, Cerrophidion, and Porthidium renders Atropoides paraphyletic (see also [175]). The species Bothrops pictus, considered incertae sedis in previous studies [176], is here strongly
supported as the sister taxon to a clade containing Rhinocerophis, Bothropoides, Bothriopsis, and Bothrops (Figure 22). Most of these relationships are strongly supported.
Viperidae is strongly placed (SHL = 95) as the sister taxon to a well-supported clade (SHL = 100) containing Colubridae, Elapidae, Homalopsidae, and Lamprophiidae (Figure 1). Monophyly of Homalopsidae is also strongly supported (Figure 23). Within Homalopsidae, the non-monophyly of the genus Enhydris is strongly supported (Figure 23), and should likely be split into multiple genera. Homalopsidae is weakly supported (SHL = 58) as the sister group of Elapidae + Lamprophiidae (Figure 1). This same relationship was also weakly supported by previous analyses [41,44], but other studies have found strong support for placing Homalopsidae as the sister group of a strongly supported clade including Elapidae, Lamprophiidae, and Colubridae [20,36], including data from >400 loci (Pyron et al., in prep.).
Support for the monophyly of Lamprophiidae is strong (but excluding Micrelaps; see below), and most of its subfamilies are well-supported [40,41,177,178] including Atractaspidinae, Aparallactinae, Lamprophiinae, Prosymninae (weakly placed as the sister-group to Oxyrhabdium), Pseudaspidinae, Psammophiinae, and Pseudoxyrhophiinae (Figure 23). In Lamprophiidae, most genera are monophyletic based on our sampling (Figure 23). However, within Aparallactinae, Xenocalamus is strongly placed within Amblyodipsas, and in Atractaspidinae, Homoroselaps is weakly placed in Atractaspis. In Lamprophiinae, Lamprophis is paraphyletic with respect to Lycodonomorphus but support for the relevant clades is weak.
The enigmatic genera Buhoma from Africa and Psammodynastes from Asia were both previously considered incertae sedis within Lamprophiidae [41]. Here they are weakly placed as sister taxa, and more importantly, they form a strongly supported clade with the African genus Pseudaspis (Pseudaspidinae; SHL = 95; Figure 23). Therefore, we expand Pseudaspidinae to include these two genera.
The genus Micrelaps (putatively an aparallactine; [1]) is weakly placed as the sister taxon to Lamprophiidae + Elapidae. Along with Oxyrhabdium (see above) and Montaspis [40], this genus is treated as incertae sedis in our classification (Appendix I). If future studies strongly support these relationships, they may require a new family for Micrelaps and possibly a new subfamily for Oxyrhabdium, though placement of these taxa has been highly variable in previous studies [40,41,44,81].
Monophyly of Elapidae is strongly supported (Figure 24), and Calliophis melanurus is strongly supported as the sister group to all other elapids (see also [44]). Within Elapidae (Figure 24), relationships are generally concordant with previous taxonomy, with some exceptions. The genera Toxicocalamus, Simoselaps, and Echiopsis are all divided across multiple clades, with strong support for many of the relevant branches. A recent study has provided a generic re-classification of the sea snakes (Hydrophis group) to resolve the extensive paraphyly of genera found in previous studies (e.g. [46,179,180]). Our results support this classification.
Monophyly of Colubridae and most of its subfamilies (sensu [41,44]) are strongly supported (Figures 1, 25, 26, 27, 28). However, relationships among many of these subfamilies are weakly supported (Figure 1), as in most previous studies [41,43,45,181]. The subfamilies Calamariinae and Pseudoxenodontinae are strongly supported as sister taxa, and weakly
placed as the sister-group to the rest of Colubridae (Figure 25). There is a weakly supported clade (Figure 1) comprising Natricinae + (Dipsadinae + Thermophis), but the clade uniting the New World Dipsadinae with the Asian genus Thermophis is strongly supported (SHL = 100; Figure 28). Here, we place Thermophis (recently in Pseudoxenodontinae [41]) in Dipsadinae (following [182]), making it the first and only Asian member of this otherwise exclusively New World subfamily. However, despite the strong support for its placement here, placement of this taxon has been variable in previous analyses [41,182,183], and we acknowledge that future analyses may support recognition of a distinct subfamily (Thermophiinae).
The clade of Natricinae and Dipsadinae is weakly supported as the sister group (Figures 1, 25, 26, 27, 28) to a clade containing Sibynophiinae [181] + (Colubrinae + Grayiinae). The subfamily Colubrinae is weakly supported; we find that the colubrine genera Ahaetulla, Chrysopelea, and Dendrelaphis form a strongly supported clade that is weakly placed as the sister group to the rest of Colubrinae, which form a strongly supported clade (Figure 25). This clade was also placed with Grayiinae or Sibynophiinae in many preliminary analyses, rendering Colubrinae paraphyletic. This group of three genera has been strongly supported in the past, and only weakly placed with Colubrinae [41,44]. It is possible that future analyses will reveal that the clade of Ahaetulla, Chrysopelea, and Dendrelaphis is placed elsewhere in Colubridae with strong support, and thus merits recognition as a distinct subfamily (Ahaetuliinae). A notable feature of this clade is the presence of gliding flight in most species of Chrysopelea, less-developed non-flight jumping with similar locomotor origins in Dendrelaphis, and homologous glide-related traits in Ahaetulla [184].
Numerous colubroid genera are not included in our tree and are not clearly placed in subfamilies based on previous morphological evidence. In our classification, these genera are also considered incertae sedis within Colubridae, including Blythia, Cyclocorus, Elapoidis, Gongylosoma, Helophis, Myersophis, Oreocalamus, Poecilopholis, Rhabdops, and Tetralepis, as in previous classifications [1].
Our phylogeny reveals numerous taxonomic problems within Colubrinae (Figures 25, 26). The genus Boiga is paraphyletic with respect to Crotaphopeltis, Dipsadoboa, Telescopus, Toxicodryas, and Dasypeltis, with strong support (Figure 25). The genus Philothamnus is paraphyletic with respect to Hapsidophrys (Figure 25). The genus Coluber is split between Old World and New World clades (Figures 25, 26). The species Hierophis spinalis is sister to Eirenis to the exclusion of the other Hierophis species (Figure 25). The genus Dryocalamus is nested within Lycodon (Figure 26). The species Chironius carinatus and C. quadricarinatus are weakly placed in a clade of Neotropical colubrines only distantly related to the other Chironius species (Figure 26). The genus Drymobius renders Dendrophidion paraphyletic (Figure 26). The monotypic genus Rhynchophis renders the two species of Rhadinophis paraphyletic (Figure 26). The genus Coronella is rendered paraphyletic (Figure 26) by Oocatochus with weak support see also [185]. Finally, the genus Rhinechis is nested within Zamenis (Figure 26).
We find numerous non-monophyletic genera within Natricinae (Figure 27), as in previous studies [41,44,78,186]. These non-monophyletic genera include the Asian genera Amphiesma, Atretium, and Xenocrophis. Among New World genera, we find Regina to be non-monophyletic with respect to most other genera, as in previous phylogenetic studies (e.g. [41,186]). Also, as in previous studies (e.g. [41,187]), we find that Adelophis [188] is nested deep within Thamnophis [189].
Finally, within a weakly supported Dipsadinae (Figure 28), we find non-monophyly of numerous genera, as in many earlier studies (e.g. [41-43,190]). These problems of non-monophyly include Leptodeira (with respect to Imantodes), Geophis (with respect to Atractus), Atractus (with respect to Geophis), Sibynomorphus (with respect to Dipsas), Dipsas (with respect to Sibynomorphus), Taeniophallus (with respect to Echinanthera), and Echinanthera (with respect to Taeniophallus). Recent revisions have begun to tackle these problems [42,43,190], but additional taxon and character sampling will be crucial to resolve relationships and taxonomy.
Discussion
In this study, we provide a phylogenetic estimate for 4161 species of squamates based on molecular data from up to 12 genes per species, combining much of the relevant data used in previous molecular phylogenetic analyses. This tree provides a framework for future evolutionary studies, spanning from the species level to relationships among families, utilizing a common set of branch lengths. These estimated branch lengths are critically important for most phylogenetic comparative methods. To further facilitate use of this phylogeny in comparative studies, we provide the Newick version of this tree (with estimated branch lengths) in DataDryad repository 10.5061/dryad.82h0m and Additional file 1 Data File S1. Our results also suggest that the branch lengths in this tree should not generally be compromised by missing data for some genes in some taxa.
Our results also reveal many problems in squamate classification at nearly all phylogenetic levels. We make several changes to higher-level taxonomy based on this phylogeny, including changes to the traditionally recognized subfamilies of boid snakes (i.e. resurrecting Sanziniinae for the boine genera Acrantophis and Sanzinia, erecting Candoiinae for the boine genus Candoia, and moving Lichanura and Charina from Erycinae to Ungaliophiinae), lamprophiid snakes (expansion of Pseudaspidinae to include the formerly incertae sedis genera Buhoma and Psammodynastes), colubrid snakes (expansion of Dipsadinae to include the Asian pseudoxenodontine genus Thermophis), and gymnophthalmid lizards (recognition of Bachiinae for the tribe cercosaurine Bachiini, containing Bachia) and scincid lizards (synonymizing Feylininae with Scincinae to yield a total of three scincid subfamilies: Acontiinae, Lygosominae, and Scincinae). In Appendix I, we list the generic content of all families and subfamilies. We also find dozens of problems at the genus level, many of which have been identified previously, and which we defer the resolution of to future studies. Our results also highlight potential problems in recent proposals to modify the classification of scincid [112] and teiid lizards [123].
In addition to synthesizing existing molecular data for squamate phylogeny, our analyses also reveal several apparently novel findings (Figures 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20, 21, 22, 23, 24, 25, 26, 27, 28). Given space constraints, we cannot detail every deviation from previous phylogenetic hypotheses (especially pre-molecular studies). Nevertheless, we focus on three sets of examples. First, we find some relatively novel, strongly-supported relationships at the family level. These include the placement of Helodermatidae (as sister to Xenosauridae, Anguidae, and Anniellidae) and the placement of Xenodermatidae as the sister taxon to Acrochordidae (rendering Colubroidea paraphyletic), in contrast to most recent analyses of anguimorphs and snakes (see above). We also find some novel, strongly supported relationships among pleurodont families, but we acknowledge that these may be overturned in future studies.
The second example is the higher-level relationships within Scincidae, the largest family of lizards [1]. No previous studies examining higher-level relationships within the group included more than ~50 species [50,51]. In this study, we sample 683 skink species (Figures 6, 7, 8, 9, 10), and our phylogeny provides a unique resolution of higher-level skink relationships. Previous researchers [51] placed acontiines as the sister group to all other skinks, but suggested that scincines and lygosomines were paraphyletic with respect to each other (with feyliniines placed with scincines). In contrast, others [50] suggested that acontiines, feyliniines, and lygosomines were all nested inside scincines (but with each of those three subfamilies as monophyletic), although many clades were only weakly supported. Here (Figures 1, 6, 7, 8, 9, 10), we find that acontiines are the sister group to a strongly supported clade consisting of a monophyletic Scincinae and a monophyletic Lygosominae (excepting the weakly supported placement of Ateuchosaurus and placement of Feyliniinae in Scincinae).
Third, our phylogeny reveals numerous genera that appear to be non-monophyletic, with many of these cases having strong support for the associated nodes. Our examples include genera in many families, including dibamids, diplodactylids, gekkonids, phyllodactylids, gerrhosaurids, scincids, teiids, gymnophthalmids, lacertids, anguids, chamaeleonids, agamids, tropidurids, oplurids, leiosaurids, typhlopids, pythonids, uropeltids, boids, viperids, lamprophiids, elapids, and colubrids (see Results). Although many problems noted here were found in previous studies, some seem to be new, such as placement of Crocodilurus and Draceana within Tupinambis (in Teiidae; Figure 11) and Coloptychon within Gerrhonotus (in Anguidae; Figure 14).
Our study also offers an important test of higher-level squamate relationships using a very different sampling strategy than that used in most previous analyses. Squamates have traditionally been divided into two clades based on morphology, Iguania and Scleroglossa (e.g. [13,21]). Despite considerable disagreement among morphology-based hypotheses, this basic division is supported by nearly all phylogenetic analyses based on morphological data [13-15,19,22,95,191]. In contrast, our results and those of most previous molecular analyses strongly support placement of iguanians with anguimorphs and snakes [16-20,23,24]. The causes of this conflict remain unclear, but may be related to morphological traits associated with different feeding strategies of iguanian and (traditional) scleroglossan squamates [3,17].
Additionally, analyses of morphology often place dibamids, amphisbaenians, snakes, and (in some cases) some scincids and anguids in a single clade [13-15]. Our analyses do not support such a clade (Figure 1), nor do other analyses of molecular data alone [17-20], or analyses of combined molecular and morphological data [19]. Instead, these morphological results seem to be explained by convergence associated with burrowing (e.g. [19]). Overall, molecular datasets have shown overwhelmingly strong support for placement of dibamids and gekkonids at the base of the tree, amphisbaenians with lacertoids, and iguanians with snakes and anguimorphs [17-20,23]. These results have now been corroborated with up to 22 genes (15794 bp) for 45 taxa [19], 25 genes (19020 bp) for 64 taxa [16], and 44 genes (33717 bp) for 161 taxa [20]. We now support this basic topology with 4161 species sampled for up to 12 genes each (up to 12896 bp).
Nevertheless, despite the overall strong support for most of the tree (i.e. 70% of all nodes have SHL > 85), certain clades remain poorly supported (e.g. relationships among many pleurodont iguanian families; Figures 1, 17, 18, 19). A potential criticism of the supermatrix approach used here is that this weak support may occur due to missing data. However,
previous studies of 8 datasets have shown explicitly that there is typically little relationship between branch support for terminal taxa and the amount of missing data [85]. Instead, these patterns of weak support are more likely to reflect short underlying branch lengths [20,36,41], and may be difficult to resolve even with more complete taxonomic and genomic sampling. Indeed, as noted above, many of the weakly supported nodes in our phylogeny are also weakly supported in analyses with little missing data (<20%) and large numbers of genes (e.g. 44 genes as in [20]), such as the relationships of many pleurodont lizard families and colubroid snake families and subfamilies.
We acknowledge that the differences between our results and previous studies (noted above) do not necessarily mean that our results are right and those of previous studies are wrong. In some cases, we provide strong support for novel relationships when previous, conflicting studies showed only weak support (as in scincids, see above). In other cases, our results disagree with other studies for clades that were strongly supported (e.g. placement of xenodermatids). In the best-case scenario, these conflicts may be resolved because our results are correct, possibly reflecting the beneficial effects of adding taxa and the associated subdivision of long branches [25,26,28,87,192-194]. Furthermore, in many cases, we are including more genes than used in previous studies of particular clades, increasing sampling of characters and loci. This should generally reduce spurious results caused by sampling few characters and by incongruence between gene and species trees.
However, other explanations for incongruence between our results and previous studies are also possible. Adding taxa can potentially lead to incorrect results in some cases (e.g. [195]), such as when a long terminal branch is added that further subdivides a short internal branch. In other cases, conflicts with our results might reflect the impact of our sampling fewer nuclear genes and a correspondingly increased influence of mitochondrial data. Mitochondrial genes have relatively fast evolutionary rates (potentially exacerbating the impacts of long branches), and their phylogenetic resolution for a particular node may also reflect introgression or incomplete lineage sorting rather than the species phylogeny (review in [196]). Many taxa in the matrix are represented only by mitochondrial data, and highly variable mitochondrial genes might also overcome the influence of less variable nuclear genes in combined analyses (although this scenario does not seem to be common [196]). Such cases might explain some strongly supported conflicts between our results and those based on multiple nuclear loci. Another possibility is that some cases may represent failure to find the optimal tree (although we assume that these cases will likely show only weak support). We acknowledge that there are many reasons why our results may differ from previous studies, and the ultimate test of these novel findings will be corroboration in future studies that include more taxa and characters.
This analysis also corroborates several recent studies suggesting that the supermatrix approach is a powerful strategy for large-scale phylogenetic inference [41,72,73,75,76,197]. For example, even though each species had 81% missing data on average, we found that most species were placed in the families and genera expected based on previous taxonomy, often with very strong support. Furthermore, we found that incompleteness of terminal taxa is not related to branch lengths (at least not terminal branch lengths), suggesting that missing data are not significantly biasing branch-length estimates (see also [84,86,87]). Also, the CAT-based models we used have been shown to be robust to missing data in large, sparse supermatrices [84].
Even though we did find some subfamilies and genera to be non-monophyletic, similar relationships were often found in previous studies based on data matrices with only limited missing data (e.g. non-monophyly of boid snake subfamilies [47], lacertid and scincid lizard genera [67,111], and scolecophidian, dipsadine, and natricine snake genera [38,43,186]). We suggest that further resolution of the squamate tree will be greatly facilitated if researchers deliberately sample mitochondrial genes and nuclear genes that include the set of genes used here and in recent phylogenomic studies (e.g. [20]), to increase overlap between genes and taxa, and decrease missing data.
With over 5000 species remaining to be included and only 12 genes sampled, our study is far from the last word on squamate phylogeny. We note that new data can easily be added to this matrix, in terms of both new taxa and new genes. Increased sampling of other nuclear genes is likely to be advantageous as well. Next-generation sequencing strategies and phylogenomic methods should help resolve difficult nodes [16,20,36,198-200], as should application of species-tree methods [201,202]. Species-tree analyses of 44 nuclear loci support many of these same clades across squamates [20], and data from >400 nuclear loci reinforces many of the relationships found here among the colubroid snake subfamilies (Pyron et al., in prep.). In addition, it would be useful to incorporate fossil taxa in future studies [15,19,22,86], utilizing the large morphological datasets that are now available [14,15]. Despite these areas for future studies, the present tree provides a framework for researchers analyzing patterns of squamate evolution at both lower and higher taxonomic levels (e.g. [10,11,203,204]), and for building a more complete picture of squamate phylogeny.
Conclusions
In this study, we provide a phylogenetic estimate for 4161 squamate species, based on a supermatrix approach. Our results provide important confirmation for previous studies based on more limited taxon sampling, and reveal new relationships at the level of families, genera, and species. We also provide a phylogenetic framework for future comparative studies, with a large-scale tree including a common set of estimated branch lengths. Finally, we provide a revised classification for squamates based on this tree, including changes in the higher-level taxonomy of gymnophthalmid and scincid lizards and boid, colubrid, and lamprophiid snakes.
Methods
Initial classification
Our initial squamate classification is based on the June 2009 version of the Reptile Database (http://www.reptile-database.org/), accessed in September of 2009 when this research was begun. Minor modifications to this scheme were made, primarily to update changes in colubroid snake taxonomy [41-44,205]. This initial taxonomic database consists of 8650 species (169 amphisbaenians, 5270 lizards, 3209 snakes, and 2 tuataras), against which the classification of species in the molecular sequence database was fixed. While modifications and updates (i.e. new species, revisions) have been made to squamate taxonomy subsequently, these are minor and should have no impact on our phylogenetic results. This database represents ~92% of the current estimated diversity of squamates (~9400 species as of December 2012).
Throughout the paper, we refer to the updated version of squamate taxonomy from the December 2012 update of the Reptile Database, incorporating major, well-accepted changes from recent studies (summarized in [1]). However, for large, taxonomic groups that have recently been broken up for reasons other than resolving paraphyly or matters of priority (e.g. in dactyloid and scincid lizards; see Results), we generally retain the older, more inclusive name in the interest of clarity, while providing references to the recent revision. We attempt to alter existing classifications as little as possible (see also [113]). Therefore, we generally only make changes when there is strong support for non-monophyly of currently recognized taxa and our proposed changes yield strongly supported monophyletic groups. Similarly, we only erect new taxa if they are strongly supported. Finally, although numerous genera are identified as being non-monophyletic in our tree, we refrain from changing genus-level taxonomy, given that our taxon sampling within many genera is limited.
Molecular data
Preliminary literature searches were conducted to identify candidate genes for which a substantial number of squamate species were sequenced and available on GenBank (with the sampled species spread across multiple families), and which were demonstrably useful in previous phylogenetic studies of squamates (see Introduction for references). Twelve genes were identified as meeting these criteria: seven nuclear genes (brain-derived neurotrophic factor [BDNF], oocyte maturation factor [c-mos], neurotrophin-3 [NT3], phosducin [PDC], G protein-coupled receptor 35 [R35], and recombination-activating genes 1 [RAG-1] and 2 [RAG-2]), and five mitochondrial genes (12S/16S long and short subunit RNAs, cytochrome b [cyt-b], and nicotinamide adenine dehydrogenase subunits 2 [ND2] and 4 [ND4]). This sampling of genes does not include all available markers. For example, we omitted several nuclear and mitochondrial genes because they were available only for a limited subset of taxa. We also excluded tRNAs associated with the protein-coding sequences, given their short lengths and difficulty in alignment across the large time scales considered here.
To ensure maximal taxonomic coverage from the available data, searches were conducted on GenBank by family (stopping in October 2012), and the longest sequence for every species was gathered. Sequences totaling less than 250 bp for any species were not included. Only species in the taxonomic database were included in the sequence matrix, which resulted in the exclusion of numerous named taxa of ambiguous status, a few taxa described very recently, and many sequences labeled 'sp.' Some recently described phylogeographic lineages were also omitted. Species and GenBank accession numbers are available in Additional file 2 Table S1.
With respect to the December 2012 update of the Reptile Database, we sampled 52 of 183 amphisbaenian species (28%) from 11 of 19 (58%) genera; 2847 of 5799 lizard species (49%) from 448 of 499 genera (90%); and 1262 of 3434 snake species (39%) in 396 of 500 genera (80%). This yielded a total of 4161 species in 855 genera in the final matrix, 44% of the 9416 known, extant squamate species in 84% of 1018 genera [1]. The species-level classification of squamates is in constant flux, and the numbers of species and genera changed even as this paper was under review. The extant species of tuatara (Sphenodon punctatus) was included as a non-squamate outgroup taxon (see below). We acknowledge that our sampling of outgroup taxa is not extensive. However, placement of Sphenodon as the sister group to squamates is well-established by molecular analyses with extensive taxon sampling (e.g. [16,128,206]) and morphological data (e.g. [13]).
Alignment for protein-coding sequences was relatively straightforward. We converted them to amino acids, and then used the translation alignment algorithm in the program Geneious v4.8.4 (GeneMatters Corp.), with the default cost matrix (Blosum62) and gap penalties (open=12, extension=3). Alignments were relatively unambiguous after being trimmed for quality and maximum coverage (i.e. ambiguous end regions were removed, and most sequences began and ended at the same point).
For the ribosomal RNA sequences (12S and 16S sequences), alignment was more challenging. Preliminary global alignments using the algorithms MUSCLE [207] and CLUSTAL [208] under a variety of gap-cost parameters yielded low-quality results (i.e. alignments with large numbers of gaps and little overlap of potentially homologous characters). We subsequently employed a two-step strategy for these data. We first grouped sequences by higher taxa (i.e. Amphisbaenia, Anguimorpha, Gekkota, Iguania, Scincomorpha, and Serpentes, though these are not all monophyletic as previously defined), for which alignments were relatively straightforward under the default MUSCLE parameters.
These were then combined using the profile alignment feature of MUSCLE, and the global alignment was subsequently updated using the "refine alignment" option. Minor adjustments were then made by eye, and ambiguously aligned end-regions were trimmed for maximum coverage and quality. We did not include partitions for stems and loops for the ribosomal sequences, although this has been shown to improve model fit in previous squamate studies (e.g. [82]). Although it is possible to assign individual nucleotide positions to these partitions, this would have been challenging given the large number of sequences, and the potential for stems and loops to shift across the many species and large time scales involved.
Each species was represented by a single terminal taxon in the matrix. In many cases, sequences from multiple individuals of the same species were combined, to allow us to combine data from different genes for the same species. We acknowledge the possibility that in some cases this approach may cause us to combine genes from different species in the same terminal taxon (e.g. due to changing taxonomy or incorrect identifications). Additionally, many sequences are not from vouchered specimens, and it is possible that misidentified species are present on GenBank and in our matrix. However, most of our data came from lower-level phylogenetic studies, in which the identification of species by previous authors should be highly accurate. In addition, any such mistakes should be among closely related species, and lead to minimal phylogenetic distortion, as the grossest errors are easily identified.
Some species were removed after preliminary analyses, due either to obvious sequencing errors (e.g. high BLAST homology with unrelated families or non-squamates, excessive ambiguities) or a lack of overlap in genes sampled with other members of the same genus (leading to seemingly artificial paraphyly). We also excluded species with identical sequences between taxa across all genes, arbitrarily choosing the first taxon in alphabetical order to remain in the matrix. Additionally, we also removed a few apparent "rogue taxa" [75,77]. These were identified by their poor support and suspect placement (e.g. in a clearly incorrect family), and were typically represented in the matrix by short fragments of single genes (e.g. an ND4 fragment from the enigmatic colubroid snake Oreocalamus hanitchsi).
The final combined matrix contained sequence data for: 2335 species for 12S (including 56% of all 4162 taxa, 1395 bp), 2377 for 16S (57%, 1970 bp), 730 for BDNF (18%, 714 bp), 1671 for c-mos (40%, 903 bp), 1985 for cyt-b (48%, 1000 bp), 437 for NT3 (10%, 675 bp), 1860
for ND2 (45%, 960 bp), 1556 for ND4 (37%, 696 bp), 393 for PDC (9%, 395 bp), 401 for R35 (10%, 768 bp), 1379 for RAG-1 (33%, 2700 bp), and 471 for RAG2 (11%, 720 bp). The total alignment consists of 12896 bp for 4162 taxa (4161 squamates and 1 outgroup). The mean length is 2497 bp of sequence data present per species from 3.75 genes (19% of the total matrix length of 12896 bp, or 81% missing data), and ranges from 270–11153 bp (2–86% complete). The matrix and phylogeny (see below) are available in DataDryad repository 10.5061/dryad.82h0m.
Clearly, many taxa had large amounts of missing data (some >95%), and on average each species had 81% missing cells. However, several lines of evidence suggest that these missing data are not generally problematic. First, a large body of empirical and theoretical studies has shown that highly incomplete taxa can be accurately placed in model-based phylogenetic analyses (and with high levels of branch support), especially if a large number of characters have been sampled (recent reviews in [84,85]). Second, several recent empirical studies have shown that the supermatrix approach (with extensive missing data in some taxa) yields generally well-supported large-scale trees that are generally congruent with previous taxonomies and phylogenetic estimates (e.g. [41,48,72,73,75,76,197]). Third, recent studies have also shown that there is generally little relationship between the amount of missing data in individual taxa and the support for their placement on the tree [41,73,85]. Finally, we note that some highly incomplete taxa were unstable in their placement (“rogue taxa”;[ 75]), but these were removed prior to the analysis of the final matrix (see above).
Our sampling design should be especially robust to the impacts of missing data for several reasons. Most importantly, most terminal taxon (species) had substantial data present (mean of 2497 bp per species) regardless of the number of missing data cells. Simulations (see reviews in [84,85]) suggest that the amount of data present is a key parameter in determining the accuracy with which incomplete taxa are placed in phylogenies, not the amount of data absent. Additionally, several genes (e.g. 12S/16S, cyt-b, and c-mos) were shared by many (>40%) of taxa. Thus, there was typically extensive overlap among the genes present for each taxon (as also indicated by the mean bp per species being much greater than the length of most genes). Limited overlap in gene sampling among taxa could be highly problematic, irrespective of the amount of missing data per se, but this does not appear to be a problem in our dataset. Finally, several nuclear genes (e.g. BDNF, c-mos, R35, and RAG-1) were congruently sampled in previous studies to represent most (>80%) squamate families and subfamilies (e.g. [20]), providing a scaffold of well-sampled taxa spanning all major clades, as recommended by recent authors [84].
Phylogenetic analyses
We performed phylogenetic analyses of the 12-gene concatenated matrix using Maximum Likelihood (ML). We assessed node support using the non-parametric Shimodaira-Hasegawa-Like (SHL) implementation of the approximate likelihood-ratio test (aLRT; [94]). This involved a two-stage strategy. We first performed initial ML tree inference using the program RAxML-Light v1.0.7 [209], a modification of the original RAxML algorithm [210]. This program uses the GTRCAT strategy for all genes and partitions, a high-speed approximation of the GTR+Γ model (general time-reversible with gamma-distribution of rate heterogeneity among sites). The GTR model is the only substitution model implemented in RAxML [210], and all other substitution models are simply special cases of the GTR model [211]. Previous analyses suggest that GTR is generally the best-fitting model for these genes and that they should be partitioned by gene and codon position [16,17,19,20,36,81].
To generate an initial ML estimate for final optimization and support estimation, we performed 11 ML searches from 11 parsimony starting trees generated under the default parsimony model in RAxMLv7.2.8. This number is likely to be sufficient when datasets contain many characters that have strong phylogenetic signal (A. Stamatakis, pers. comm.). Additionally, the dataset was analyzed with these settings (GTRCAT search from a randomized parsimony starting tree) numerous times (>20) as the final matrix was assembled and tested, representing hundreds of independent searches from random starting points. All of the estimated trees from these various analyses showed high overall congruence with the final topology. The concordance between the preliminary and final results suggests that the tree was not strongly impacted by searches stuck on local optima, and that it should be a good approximation of the ML tree.
We then performed a final topology optimization and assessed support. We passed our best ML estimate of the phylogeny (based on GTRCAT) from RAxML-Light to RAxMLv7.2.8, which does an additional search (using the GTRGAMMA model) to produce a nearest-neighbor interchange (NNI)-optimized estimate of the ML tree. This optimization is needed to calculate the SHL version of the aLRT for estimating support values [94]. The SHL-aLRT strategy approximates a test of the null hypothesis that the branch length subtending each node equals 0 (i.e. that the node can be resolved, rather than estimated as a polytomy) with a test of the more general null hypothesis that "the branch is incorrect" relative to the four next suboptimal arrangements of that node relative to the NNI-optimal arrangement [94]. Based on initial analyses, generating sufficient ML bootstrap replicates for a tree of this size proved computationally intractable, so we rely on SHL values alone to assess support.
The SHL approach has at least two major advantages over non-parametric bootstrapping for large ML trees: (i) values are apparently robust to many potential model violations and have the same properties as bootstrap proportions for all but the shortest branches [41,94,212], and (ii) values for short branches may be more accurate than bootstrap proportions, as support is evaluated based on whole-alignment likelihoods, rather than the frequency of re-sampled characters [94,213]. Additionally, the SHL approach is orders of magnitude faster than traditional bootstrapping [94,212,213], and it appears to be similarly robust to matrices with extensive missing data [41]. As in previous studies, we take a conservative view, considering SHL values of 85 or greater (i.e. a 15% chance that a branch is "incorrect") as strong support [41,212,213].
These analyses were performed on a 360-core SGI ICE supercomputing system ("ANDY") at the High Performance Computing Center at the City University of New York (CUNY). The final analysis was completed in 8.8 days of computer time using 188 nodes of the CUNY supercomputing cluster.
Finally, we assessed the potential impact of missing data on our branch-length estimates. We performed linear regression (in R) of the proportional completeness of each terminal taxa (non-missing data in bp / maximum amount of non-missing data, 12896 bp) against the length of its terminal branch. This test addresses whether incomplete taxa have branch-length estimates that are consistently biased in one direction (shorter vs. longer) relative to more complete terminals. However, it does not directly test whether branch length estimates are correct or not, nor how branch length estimates are impacted by replacing non-missing data with missing data (see [87] for results suggesting that such replacements have little effect in real data sets).
Authors' contributions
RAP and JJW conceived the study. RAP, FTB, and JJW conducted analyses and checked results. RAP, FTB, and JJW wrote the MS. All authors read and approved the final manuscript.
Acknowledgements
We thank the many researchers who made this study possible through their detailed studies of squamate phylogeny with a (mostly) shared set of molecular markers, and uploading their sequence data to GenBank. We thank E. Paradis, J. Lombardo T. Guiher, R. Walsh, and P. Muzio for computational assistance, T. Gamble, D. Frost, D. Cannatella, H. Zaher, F. Grazziotin, P. Uetz, T. Jackman, and A. Bauer for taxonomic advice, and A. Larson, M. Vences, and five anonymous reviewers for comments on this manuscript. The research was supported in part by the U.S. National Science Foundation, including a Bioinformatics Postdoctoral grant to R.A.P. (DBI-0905765), an AToL grant to J.J.W. (EF-0334923), and grants to the CUNY HPCC (CNS-0958379 and CNS-0855217).
Appendix I
Note that we only provide here an account for the one subfamily newly erected in this study. We do not provide accounts for subfamilies with changes in content (Boinae, Erycinae, Dipsadinae, Pseudaspidinae, Scincinae, Ungaliophiinae), that have been resurrrected (Sanziniinae), or that represent elevation of lower-ranked taxa (tribe Bachiini here recognized as Bachiinae).
Candoiinae subfam. nov. (family Boidae)
Type: genus and species Candoia carinata [214].
Content: one genus, 4 species; C. aspera, C. bibroni, C. carinata, C. paulsoni.
Definition: this subfamily consists of the most recent common ancestor of the extant species of Candoia, and all its descendants. These species are morphologically distinguished in part from other boid snakes by a flattened rostrum leading to an angular snout [215] and a wide premaxillary floor [167].
Distribution: these snakes are primarily restricted to the South Pacific islands of New Guinea and Melanesia, and the eastern Indonesian archipelago [150].
Remarks: the three species from this subfamily that are sampled in our tree are strongly supported as monophyletic (SHL = 100), and are well supported (SHL = 87) as the sister taxon to a moderately supported clade consisting of Erycinae + Boinae (SHL = 83).
Proposed Generic Composition of Higher Taxa
Below, we list the familial and subfamilial assignment of all squamate genera from the December, 2012 update of the Reptile Database, updated to reflect some recent changes and
the proposed subfamily level changes listed above. As this classification includes numerous taxa not sampled in our tree, we deal with this conservatively. For traditionally recognized families and subfamilies that we found to be monophyletic, we include all taxa traditionally assigned to them. Taxa are denoted incertae sedis if they are of ambiguous familial or subfamilial assignment due to uncertain placement in our tree, or due to absence from our tree and lack of assignment by previous authors. This classification includes 67 families and 56 subfamilies, and accounts for >9400 squamate species in 1018 genera [1]. Higher taxa are listed (more-or-less) phylogenetically (starting closest to the root; Figure 1), families are listed alphabetically within higher taxa, and subfamilies and genera are listed alphabetically within families.
Dibamidae (Anelytropsis, Dibamus)
Gekkota
Carphodactylidae (Carphodactylus, Nephrurus, Orraya, Phyllurus, Saltuarius, Underwoodisaurus, Uvidicolus); Diplodactylidae (Amalosia, Bavayia, Correlophus, Crenadactylus, Dactylocnemis, Dierogekko, Diplodactylus, Eurydactylodes, Hesperoedura, Hoplodactylus, Lucasium, Mniarogekko, Mokopirirakau, Naultinus, Nebulifera, Oedodera, Oedura, Paniegekko, Pseudothecadactylus, Rhacodactylus, Rhynchoedura, Strophurus, Toropuku, Tukutuku, Woodworthia); Eublepharidae (Aeluroscalabotes, Coleonyx, Eublepharis, Goniurosaurus, Hemitheconyx, Holodactylus); Gekkonidae (Afroedura, Afrogecko, Agamura, Ailuronyx, Alsophylax, Asiocolotes, Blaesodactylus, Bunopus, Calodactylodes, Chondrodactylus, Christinus, Cnemaspis, Colopus, Crossobamon, Cryptactites, Cyrtodactylus, Cyrtopodion, Dixonius, Ebenavia, Elasmodactylus, Geckolepis, Gehyra, Gekko, Goggia, Hemidactylus, Hemiphyllodactylus, Heteronotia, Homopholis, Lepidodactylus, Luperosaurus, Lygodactylus, Matoatoa, Mediodactylus, Nactus, Narudasia, Pachydactylus, Paragehyra, Paroedura, Perochirus, Phelsuma, Pseudoceramodactylus, Pseudogekko, Ptenopus, Ptychozoon, Rhinogecko, Rhoptropella, Rhoptropus, Stenodactylus, Tropiocolotes, Urocotyledon, Uroplatus); Phyllodactylidae (Asaccus, Gymnodactylus, Haemodracon, Homonota, Phyllodactylus, Phyllopezus, Ptyodactylus, Tarentola, Thecadactylus); Pygopodidae (Aprasia, Delma, Lialis, Ophidiocephalus, Paradelma, Pletholax, Pygopus); Spha (Aristelliger, Chatogekko, Coleodactylus, Euleptes, Gonatodes, Lepidoblepharis, Pristurus, Pseudogonatodes, Quedenfeldtia, Saurodactylus, Sphaerodactylus, Teratoscincus)
Scincoidea
Cordylidae, Cordylinae (Chamaesaura, Cordylus, Hemicordylus, Karusasaurus, Namazonurus, Ninurta, Ouroborus, Pseudocordylus, Smaug), Platysaurinae (Platysaurus); Gerrhosauridae, Gerrhosaurinae (Cordylosaurus, Gerrhosaurus, Tetradactylus), Zonosaurinae (Tracheloptychus, Zonosaurus); Scincidae, Acontiinae (Acontias, Typhlosaurus), Lygosominae (Ablepharus, Afroablepharus, Anomalopus, Asymblepharus, Ateuchosaurus, Bartleia, Bassiana, Bellatorias, Caledoniscincus, Calyptotis, Carlia, Cautula, Celatiscincus, Chioninia, Coeranoscincus, Coggeria, Cophoscincopus, Corucia, Cryptoblepharus, Ctenotus, Cyclodomorphus, Dasia, Egernia, Emoia, Eremiascincus, Eroticoscincus, Eugongylus, Eulamprus, Eumecia, Eutropis, Fojia, Geomyersia, Geoscincus, Glaphyromorphus, Gnypetoscincus, Graciliscincus, Haackgreerius, Hemiergis, Hemisphaeriodon, Insulasaurus, Isopachys, Kaestlea, Kanakysaurus, Lacertaspis,
Lacertoides, Lamprolepis, Lampropholis, Lankascincus, Larutia, Leiolopisma, Lepidothyris, Leptoseps, Leptosiaphos, Lerista, Liburnascincus, Liopholis, Lioscincus, Lipinia, Lissolepis, Lobulia, Lygisaurus, Lygosoma, Mabuya, Marmorosphax, Menetia, Mochlus, Morethia, Nangura, Nannoscincus, Niveoscincus, Notoscincus, Oligosoma, Ophioscincus, Otosaurus, Panaspis, Papuascincus, Parvoscincus, Phoboscincus, Pinoyscincus, Prasinohaema, Proablepharus, Pseudemoia, Ristella, Saiphos, Saproscincus, Scincella, Sigaloseps, Simiscincus, Sphenomorphus, Tachygyia, Tiliqua, Trachylepis, Tribolonotus, Tropidophorus, Tropidoscincus, Tytthoscincus, Vietnascincus), Scincinae (Amphiglossus, Androngo, Barkudia, Brachymeles, Chabanaudia, Chalcides, Chalcidoseps, Eumeces, Eurylepis, Feylinia, Gongylomorphus, Hakaria, Janetaescincus, Jarujinia, Madascincus, Melanoseps, Mesoscincus, Nessia, Ophiomorus, Pamelaescincus, Paracontias, Plestiodon, Proscelotes, Pseudoacontias, Pygomeles, Scelotes, Scincopus, Scincus, Scolecoseps, Sepsina, Sepsophis, Sirenoscincus, Typhlacontias, Voeltzkowia); Xantusiidae, Cricosaurinae (Cricosaura), Lepidophyminae (Lepidophyma), Xantusiinae (Xantusia)
Lacertoidea (including Amphisbaenia)
Amphisbaenidae (Amphisbaena, Ancylocranium, Baikia, Chirindia, Cynisca, Dalophia, Geocalamus, Loveridgea, Mesobaena, Monopeltis, Zygaspis); Bipedidae (Bipes); Blanidae (Blanus); Cadeidae (Cadea); Gymnophthalmidae, Alopoglossinae (Alopoglossus, Ptychoglossus), Bachiinae (Bachia), Cercosaurinae (Anadia, Cercosaura, Echinosaura, Euspondylus, Macropholidus, Neusticurus, Opipeuter, Petracola, Pholidobolus, Placosoma, Potamites, Proctoporus, Riama, Riolama, Teuchocercus), Ecpleopinae (Adercosaurus, Amapasaurus, Anotosaura, Arthrosaura, Colobosauroides, Dryadosaura, Ecpleopus, Kaieteurosaurus, Leposoma, Marinussaurus, Pantepuisaurus), Gymnophthalminae (Acratosaura, Alexandresaurus, Calyptommatus, Caparaonia, Colobodactylus, Colobosaura, Gymnophthalmus, Heterodactylus, Iphisa, Micrablepharus, Nothobachia, Procellosaurinus, Psilophthalmus, Scriptosaura, Stenolepis, Tretioscincus, Vanzosaura), Rhachisaurinae (Rhachisaurus); Lacertidae, Gallotiinae (Gallotia, Psammodromus), Lacertinae (Acanthodactylus, Adolfus, Algyroides, Anatololacerta, Apathya, Archaeolacerta, Atlantolacerta, Australolacerta, Congolacerta, Dalmatolacerta, Darevskia, Dinarolacerta, Eremias, Gastropholis, Heliobolus, Hellenolacerta, Holaspis, Iberolacerta, Ichnotropis, Iranolacerta, Lacerta, Latastia, Meroles, Mesalina, Nucras, Omanosaura, Ophisops, Parvilacerta, Pedioplanis, Philochortus, Phoenicolacerta, Podarcis, Poromera, Pseuderemias, Scelarcis, Takydromus, Teira, Timon, Tropidosaura, Zootoca); Rhineuridae (Rhineura); Teiidae, Teiinae (Ameiva, Aspidoscelis, Cnemidophorus, Dicrodon, Kentropyx, Teius), Tupinambinae (Callopistes, Crocodilurus, Dracaena, Tupinambis); Trogonophiidae (Agamodon, Diplometopon, Pachycalamus, Trogonophis)
Iguania
Agamidae, Agaminae (Acanthocercus, Agama, Brachysaura, Bufoniceps, Laudakia, Phrynocephalus, Pseudotrapelus, Trapelus, Xenagama), Amphibolurinae (Amphibolurus, Chelosania, Chlamydosaurus, Cryptagama, Ctenophorus, Diporiphora, Hypsilurus, Intellagama, Lophognathus, Moloch, Physignathus, Pogona, Rankinia, Tympanocryptis), Draconinae (Acanthosaura, Aphaniotis, Bronchocela, Calotes, Ceratophora, Complicitus, Cophotis, Coryphophylax, Dendragama, Draco, Gonocephalus, Harpesaurus, Hypsicalotes, Japalura, Lophocalotes, Lyriocephalus, Mantheyus, Oriocalotes, Otocryptis, Phoxophrys, Psammophilus, Pseudocalotes, Pseudocophotis, Ptyctolaemus, Salea, Sitana, Thaumatorhynchus), Hydrosaurinae (Hydrosaurus), Leiolepidinae (Leiolepis),
Uromastycinae (Uromastyx); Chamaeleonidae, Brookesiinae (Brookesia), Chamaeleoninae (Archaius, Bradypodion, Calumma, Chamaeleo, Furcifer, Kinyongia, Nadzikambia, Rhampholeon, Rieppeleon, Trioceros); Corytophanidae (Basiliscus, Corytophanes, Laemanctus); Crotaphytidae (Crotaphytus, Gambelia); Dactyloidae (Anolis); Hoplocercidae (Enyalioides, Hoplocercus, Morunasaurus); Iguanidae (Amblyrhynchus, Brachylophus, Conolophus, Ctenosaura, Cyclura, Dipsosaurus, Iguana, Sauromalus); Leiocephalidae (Leiocephalus); Leiosauridae, Enyaliinae (Anisolepis, Enyalius, Urostrophus), Leiosaurinae (Diplolaemus, Leiosaurus, Pristidactylus); Liolaemidae (Ctenoblepharys, Liolaemus, Phymaturus); Opluridae (Chalarodon, Oplurus); Phrynosomatidae (Callisaurus, Cophosaurus, Holbrookia, Petrosaurus, Phrynosoma, Sceloporus, Uma, Urosaurus, Uta); Polychrotidae (Polychrus); Tropiduridae (Eurolophosaurus, Microlophus, Plica, Stenocercus, Strobilurus, Tropidurus, Uracentron, Uranoscodon)
Anguimorpha
Anguidae, Anguinae (Anguis, Dopasia, Ophisaurus, Pseudopus), Diploglossinae (Celestus, Diploglossus, Ophiodes), Gerrhonotinae (Abronia, Barisia, Coloptychon, Elgaria, Gerrhonotus, Mesaspis); Anniellidae (Anniella); Helodermatidae (Heloderma); Lanthanotidae (Lanthanotus); Shinisauridae (Shinisaurus); Varanidae (Varanus); Xenosauridae (Xenosaurus)
Serpentes
Acrochordidae (Acrochordus); Aniliidae (Anilius); Anomalepididae (Anomalepis, Helminthophis, Liotyphlops, Typhlophis); Anomochilidae (Anomochilus); Boidae, Boinae (Boa, Corallus, Epicrates, Eunectes), Candoiinae (Candoia), Erycinae (Eryx), Sanziniinae (Acrantophis, Sanzinia), Ungaliophiinae (Charina, Exiliboa, Lichanura, Ungaliophis); Bolyeriidae (Bolyeria, Casarea); Calabariidae (Calabaria); Colubridae incertae sedis (Blythia, Cyclocorus, Elapoidis, Gongylosoma, Helophis, Myersophis, Oreocalamus, Poecilopholis, Rhabdops, Tetralepis), Calamariinae (Calamaria, Calamorhabdium, Collorhabdium, Etheridgeum, Macrocalamus, Pseudorabdion, Rabdion), Colubrinae (Aeluroglena, Ahaetulla, Aprosdoketophis, Archelaphe, Argyrogena, Arizona, Bamanophis, Bogertophis, Boiga, Cemophora, Chilomeniscus, Chionactis, Chironius, Chrysopelea, Coelognathus, Coluber, Colubroelaps, Conopsis, Coronella, Crotaphopeltis, Cyclophiops, Dasypeltis, Dendrelaphis, Dendrophidion, Dipsadoboa, Dispholidus, Dolichophis, Drymarchon, Drymobius, Drymoluber, Dryocalamus, Dryophiops, Eirenis, Elachistodon, Elaphe, Euprepiophis, Ficimia, Geagras, Gonyophis, Gonyosoma, Gyalopion, Hapsidophrys, Hemerophis, Hemorrhois, Hierophis, Lampropeltis, Leptodrymus, Leptophis, Lepturophis, Limnophis, Liopeltis, Lycodon, Lytorhynchus, Macroprotodon, Mastigodryas, Meizodon, Oligodon, Oocatochus, Opheodrys, Oreocryptophis, Orthriophis, Oxybelis, Pantherophis, Philothamnus, Phyllorhynchus, Pituophis, Platyceps, Pseudelaphe, Pseudoficimia, Pseustes, Ptyas, Rhadinophis, Rhamnophis, Rhinechis, Rhinobothryum, Rhinocheilus, Rhynchocalamus, Rhynchophis, Salvadora, Scaphiophis, Scolecophis, Senticolis, Simophis, Sonora, Spalerosophis, Spilotes, Stegonotus, Stenorrhina, Symphimus, Sympholis, Tantilla, Tantillita, Telescopus, Thelotornis, Thrasops, Toxicodryas, Trimorphodon, Xenelaphis, Xyelodontophis, Zamenis), Dipsadinae (Adelphicos, Alsophis, Amastridium, Amnesteophis, Antillophis, Apostolepis, Arrhyton, Atractus, Boiruna, Borikenophis, Caaeteboia, Calamodontophis, Caraiba, Carphophis, Cercophis, Chapinophis, Chersodromus, Clelia, Coniophanes, Conophis, Contia, Coronelaps, Crisantophis, Cryophis, Cubophis,
Darlingtonia, Diadophis, Diaphorolepis, Dipsas, Ditaxodon, Drepanoides, Echinanthera, Elapomorphus, Emmochliophis, Enuliophis, Enulius, Erythrolamprus, Farancia, Geophis, Gomesophis, Haitiophis, Helicops, Heterodon, Hydrodynastes, Hydromorphus, Hydrops, Hypsiglena, Hypsirhynchus, Ialtris, Imantodes, Leptodeira, Lioheterophis, Lygophis, Magliophis, Manolepis, Mussurana, Ninia, Nothopsis, Ocyophis, Omoadiphas, Oxyrhopus, Paraphimophis, Phalotris, Philodryas, Phimophis, Plesiodipsas, Pliocercus, Pseudalsophis, Pseudoboa, Pseudoeryx, Pseudoleptodeira, Pseudotomodon, Psomophis, Ptychophis, Rhachidelus, Rhadinaea, Rhadinella, Rhadinophanes, Rodriguesophis, Saphenophis, Schwartzophis, Sibon, Sibynomorphus, Siphlophis, Sordellina, Synophis, Tachymenis, Taeniophallus, Tantalophis, Thamnodynastes, Thermophis, Tomodon, Tretanorhinus, Trimetopon, Tropidodipsas, Tropidodryas, Uromacer, Uromacerina, Urotheca, Xenodon, Xenopholis), Grayiinae (Grayia), Natricinae (Adelophis, Afronatrix, Amphiesma, Amphiesmoides, Anoplohydrus, Aspidura, Atretium, Balanophis, Clonophis, Hologerrhum, Hydrablabes, Hydraethiops, Iguanognathus, Lycognathophis, Macropisthodon, Natriciteres, Natrix, Nerodia, Opisthotropis, Parahelicops, Pararhabdophis, Paratapinophis, Regina, Rhabdophis, Seminatrix, Sinonatrix, Storeria, Thamnophis, Trachischium, Tropidoclonion, Tropidonophis, Virginia, Xenochrophis), Pseudoxenodontinae (Plagiopholis, Pseudoxenodon), Sibynophiinae (Scaphiodontophis, Sibynophis); Cylindrophiidae (Cylindrophis); Elapidae (Acanthophis, Aipysurus, Aspidelaps, Aspidomorphus, Austrelaps, Bungarus, Cacophis, Calliophis, Cryptophis, Demansia, Dendroaspis, Denisonia, Drysdalia, Echiopsis, Elapognathus, Elapsoidea, Emydocephalus, Ephalophis, Furina, Hemachatus, Hemiaspis, Hemibungarus, Hoplocephalus, Hydrelaps, Hydrophis, Kolpophis, Laticauda, Loveridgelaps, Maticora, Micropechis, Micruroides, Micrurus, Naja, Notechis, Ogmodon, Ophiophagus, Oxyuranus, Parahydrophis, Parapistocalamus, Parasuta, Pseudechis, Pseudohaje, Pseudolaticauda, Pseudonaja, Rhinoplocephalus, Salomonelaps, Simoselaps, Sinomicrurus, Suta, Thalassophis, Toxicocalamus, Tropidechis, Vermicella, Walterinnesia); Gerrhopilidae (Gerrhopilus); Homalopsidae (Bitia, Brachyorrhos, Cantoria, Cerberus, Djokoiskandarus, Enhydris, Erpeton, Fordonia, Gerarda, Heurnia, Homalopsis, Myron, Pseudoferania); Lamprophiidae incertae sedis (Micrelaps, Montaspis, Oxyrhabdium), Aparallactinae (Amblyodipsas, Aparallactus, Brachyophis, Chilorhinophis, Elapotinus, Hypoptophis, Macrelaps, Polemon, Xenocalamus), Atractaspidinae (Atractaspis, Homoroselaps), Lamprophiinae (Boaedon, Bothrophthalmus, Chamaelycus, Dendrolycus, Gonionotophis, Hormonotus, Inyoka, Lamprophis, Lycodonomorphus, Lycophidion, Pseudoboodon), Prosymninae (Prosymna), Psammophiinae (Dipsina, Hemirhagerrhis, Malpolon, Mimophis, Psammophis, Psammophylax, Rhagerhis, Rhamphiophis), Pseudaspidinae (Buhoma, Psammodynastes, Pseudaspis, Pythonodipsas), Pseudoxyrhophiine (Alluaudina, Amplorhinus, Bothrolycus, Brygophis, Compsophis, Ditypophis, Dromicodryas, Duberria, Exallodontophis, Heteroliodon, Ithycyphus, Langaha, Leioheterodon, Liophidium, Liopholidophis, Lycodryas, Madagascarophis, Micropisthodon, Pararhadinaea, Parastenophis, Phisalixella, Pseudoxyrhopus, Thamnosophis); Leptotyphlopidae (Epacrophis, Epictia, Leptotyphlops, Mitophis, Myriopholis, Namibiana, Rena, Rhinoleptus, Siagonodon, Tetracheilostoma, Tricheilostoma, Trilepida); Loxocemidae (Loxocemus); Pareatidae (Aplopeltura, Asthenodipsas, Pareas); Pythonidae (Antaresia, Apodora, Aspidites, Bothrochilus, Broghammerus, Leiopython, Liasis, Morelia, Python); Tropidophiidae (Trachyboa, Tropidophis); Typhlopidae (Acutotyphlops, Afrotyphlops, Austrotyphlops, Cyclotyphlops, Grypotyphlops, Letheobia, Megatyphlops, Ramphotyphlops, Rhinotyphlops, Typhlops); Uropeltidae (Brachyophidium, Melanophidium, Platyplectrurus, Plectrurus, Pseudotyphlops, Rhinophis, Teretrurus, Uropeltis); Viperidae, Azemiopinae (Azemiops), Crotalinae (Agkistrodon, Atropoides, Bothriechis, Bothriopsis, Bothrocophias, Bothropoides, Bothrops, Calloselasma, Cerrophidion, Crotalus, Deinagkistrodon, Garthius,
Gloydius, Hypnale, Lachesis, Mixcoatlus, Ophryacus, Ovophis, Porthidium, Protobothrops, Rhinocerophis, Sistrurus, Trimeresurus, Tropidolaemus), Viperinae (Atheris, Bitis, Causus, Cerastes, Daboia, Echis, Eristicophis, Macrovipera, Montatheris, Montivipera, Proatheris, Pseudocerastes, Vipera); Xenodermatidae (Achalinus, Fimbrios, Stoliczkia, Xenodermus, Xylophis); Xenopeltidae (Xenopeltis); Xenophidiidae (Xenophidion); Xenotyphlopidae (Xenotyphlops)
Additional files
Additional_file_1 as TXT Additional file 1: Data File S1 The 4162-species ML phylogeny in Newick format; taxonomic changes are given in Additional file 2: Table S1.
Additional_file_2 as DOCX Additional file 2: Table S1 GenBank accession numbers for all taxa included in this analysis.
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177. Kelly CMR, Branch WR, Broadley DG, Barker NP, Villet MH: Molecular systematics of the African snake family Lamprophiidae Fitzinger, 1843 (Serpentes: Elapoidea), with particular focus on the genera Lamprophis Fitzinger 1843 and Mehelya Csiki 1903. Mol Phylogenet Evol 2011, 58:415–426.
178. Vidal N, Branch WR, Pauwels OSG, Hedges SB, Broadley DG, Wink M, Cruaud C, Joger U, Nagy ZT: Dissecting the major African snake radiation: a molecular phylogeny of the Lamprophiidae Fitzinger (Serpentes, Caenophidia). Zootaxa 2008, 1945:51–66.
179. Sanders KL, Lee MSY, Leys R, Foster R, Keogh JS: Molecular phylogeny and divergence dates for Australasian elapids and sea snakes (Hydrophiinae): evidence from seven genes for rapid evolutionary radiations. J Evol Biol 2008, 21:682–695.
180. Sanders KL, Lee MSY: Molecular evidence for a rapid late-Miocene radiation of Australasian venomous snakes (Elapidae, Colubroidea). Mol Phylogenet Evol 2008, 46:1165–1173.
181. Chen X, Huang S, Guo P, Colli GR, de Oca AN M, Vitt LJ, Pyron RA, Burbrink FT: Understanding the formation of ancient intertropical disjunct distributions using Asian and Neotropical hinged-teeth snakes (Sibynophis and Scaphiodontophis: Serpentes: Colubridae). Mol Phylogenet Evol 2012, 66:254–261.
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184. Socha JJ: Gliding flight in Chrysopelea: turning a snake into a wing. Integr Comp Biol 2011, 51:969–982.
185. Helfenberger N: Phylogenetic relationship of Old World ratsnakes based on visceral organ topography, osteology, and allozyme variation. Russ J Herp 2001, 8:1–56.
186. Alfaro ME, Arnold SJ: Molecular systematics and evolution of Regina and the Thamnophiine snakes. Mol Phylogenet Evol 2001, 21:408–423.
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188. Cope ED: Eleventh contribution to the herpetology of tropical America: Dugés, A. Proc Amer Philos Soc 1879, 18:261–277.
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0.2 subst./site
Scincidae
Xantusiidae
Gerrhosauridae
Teiidae
Gymnophthalmidae
Lacertidae
Anguidae
Chamaeleonidae
Agamidae
Leiosauridae
Boidae
Viperidae
Colubridae
Lamprophiidae
A
B
C
D
E
F
G
A) Gekkota
B) Scincoidea
C) Episquamata
D) Lacertoidea
E) Toxicofera
F) Anguimorpha
G) Iguania
H) Serpentes
H
Dipsadinae
Gekkonidae
Cercosaurinae
Acontiinae
Lacertinae
Hydrosaurinae
Xenodermatidae
Anniellidae
Crotaphytidae
Hoplocercidae
Sphenodontidae
Leiolepidinae
Polychrotidae
Pseudoxyrhophiinae
Loxocemidae
Dibamidae
Prosymninae
Lygosominae
Natricinae
Xantusiinae
Tropiduridae
Corytophanidae
Ecpleopinae
Anomochilidae
Amphisbaenidae
Leiosaurinae
Xenophiidae
Scincinae
Draconinae
Bipedidae
Gerrhosaurinae
Rhineuridae
Anomalepididae
Trogonophiidae
Aniliidae
Azemiopinae
Alopoglossinae
Lanthanotidae
Pseudaspidinae
Tupinambinae
Xenotyphlopidae
Blanidae
Gerrhopilidae
Liolaemidae
Candoiinae
Gallotiinae
Leptotyphlopidae
Platysaurinae
Pythonidae
Boinae
Colubrinae
Opluridae
Elapidae
Typhlopidae
Pareatidae
Cadeidae
Crotalinae
Sibynophiinae
Chamaeleoninae
Rhachisaurinae
Grayiinae
Ungaliophiinae
Erycinae
Phyllodactylidae
Anguinae
Helodermatidae
Phrynosomatidae
Sanziniinae
Tropidophiidae
Viperinae
Cricosaurinae
Shinisauridae
Teiinae
Diplodactylidae
Cylindrophiidae
Brookesiinae
Lepidophyminae
Enyaliinae
Varanidae
Bachiinae
Aparallactinae
Sphaerodactylidae
Atractaspidinae
Uropeltidae
LeiocephalidaeIguanidae
Homalopsidae
Amphibolurinae
Xenopeltidae
Diploglossinae
Calamariinae
Xenosauridae
Uromasticinae
Agaminae
Acrochordidae
Pygopodidae
Calabariidae
Psammophiinae
Carphodactylidae
Cordylinae
Gymnophthalminae
Pseudoxenodontinae
Bolyeriidae
Dactyloidae
Gerrhonotinae
Eublepharidae
Lamprophiinae
Zonosaurinae
83
98
100
100
98
99
100
99
100
99
83
54
98
99
84
100
68
100
100
95
100
94
63
84
71
90
100
100
100
100
100
81
100
100
100
100
87
9969
90
100
100
88
65
97
72
89
100
90
100
100
100
100
96
100
100
96
100
94
100
95
100
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77
100
74
100
95
95
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100
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79
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87
100
100
100
Squamata
Cordylidae
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BTarentola gigas
Tarentola caboverdianaTarentola rudis
Tarentola darwiniTarentola chazaliae
Tarentola delalandiiTarentola gomerensis
Tarentola annularisTarentola ephippiata
Tarentola boettgeriTarentola mindiae
Tarentola neglectaTarentola angustimentalisTarentola mauritanica
Tarentola desertiTarentola boehmei
Tarentola americanaPhyllopezus maranjonensis
Phyllopezus lutzaePhyllopezus pollicaris
Phyllopezus periosusPhyllodactylus homolepidurus
Phyllodactylus davisiPhyllodactylus duellmani
Phyllodactylus delcampoiPhyllodactylus paucituberculatus
Phyllodactylus nocticolusPhyllodactylus bugastrolepisPhyllodactylus unctus
Phyllodactylus xantiPhyllodactylus bordai
Phyllodactylus laneiPhyllodactylus tuberculosus
Phyllodactylus reissiiPhyllodactylus wirshingi
Homonota andicolaHomonota darwinii
Homonota borelliiHomonota underwoodi
Homonota fasciataHomonota gaudichaudii
Ptyodactylus guttatusPtyodactylus hasselquistii
Ptyodactylus oudriiPtyodactylus ragazzii
Asaccus platyrhynchusHaemodracon riebeckii
Thecadactylus rapicaudaThecadactylus solimoensis
Sphaerodactylus nicholsiSphaerodactylus ocoae
Sphaerodactylus gaigeaeSphaerodactylus klauberi
Sphaerodactylus semasiopsSphaerodactylus goniorhynchus
Sphaerodactylus townsendiSphaerodactylus armstrongi
Sphaerodactylus macrolepisSphaerodactylus argus
Sphaerodactylus rooseveltiSphaerodactylus altavelensis
Sphaerodactylus notatusSphaerodactylus darlingtoni
Sphaerodactylus cryphiusSphaerodactylus shrevei
Sphaerodactylus elegansSphaerodactylus leucaster
Sphaerodactylus copeiSphaerodactylus nigropunctatus
Sphaerodactylus intermediusSphaerodactylus torrei
Sphaerodactylus cinereusSphaerodactylus thompsoni
Sphaerodactylus glaucusSphaerodactylus molei
Sphaerodactylus oliveriSphaerodactylus richardi
Sphaerodactylus cricoderusSphaerodactylus ramsdeni
Sphaerodactylus schwartziSphaerodactylus kirbyiSphaerodactylus vincenti
Sphaerodactylus microlepisSphaerodactylus parvus
Sphaerodactylus sabanusSphaerodactylus elegantulus
Sphaerodactylus sputatorSphaerodactylus fantasticus
Pseudogonatodes guianensisPseudogonatodes lunulatus
Pseudogonatodes manessiColeodactylus natalensis
Coleodactylus meridionalisColeodactylus brachystoma
Coleodactylus septentrionalisGonatodes ocellatusGonatodes ceciliaeGonatodes seigliei
Gonatodes concinnatusGonatodes antillensis
Gonatodes humeralisGonatodes falconensis
Gonatodes taniaeGonatodes purpurogularis
Gonatodes alexandermendesiGonatodes superciliaris
Gonatodes annularisGonatodes hasemani
Gonatodes infernalisGonatodes vittatus
Gonatodes petersiGonatodes albogularis
Gonatodes daudiniGonatodes caudiscutatus
Gonatodes eladioiLepidoblepharis festae
Lepidoblepharis xanthostigmaChatogekko amazonicus
Saurodactylus mauritanicusTeratoscincus roborowskii
Teratoscincus przewalskiiTeratoscincus scincusTeratoscincus microlepisEuleptes europaea
Saurodactylus fasciatusAristelliger georgeensis
Aristelliger praesignisAristelliger lar
Quedenfeldtia moerensQuedenfeldtia trachyblepharus
Pristurus somalicusPristurus crucifer
Pristurus carteriPristurus minimusPristurus rupestrisPristurus flavipunctatus
Pristurus abdelkuriPristurus guichardi
Pristurus sokotranusPristurus insignis
Pristurus celerrimusGoniurosaurus araneus
Goniurosaurus luiiGoniurosaurus catbaensis
Goniurosaurus lichtenfelderiGoniurosaurus kuroiwae
Hemitheconyx tayloriHemitheconyx caudicinctus
Holodactylus africanusEublepharis turcmenicusEublepharis macularius
Coleonyx elegansColeonyx mitratus
Coleonyx variegatusColeonyx brevis
Aeluroscalabotes felinus
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99
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10065
100
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Phyllodactylidae
Eublepharidae
Sphaerodactylidae
CFigure 3
CPhelsuma kely
Phelsuma lineataPhelsuma comorensis
Phelsuma pusillaPhelsuma quadriocellata
Phelsuma antanosyPhelsuma klemmeri
Phelsuma robertmertensiPhelsuma laticaudaPhelsuma serraticaudaPhelsuma malamakibo
Phelsuma berghofiPhelsuma hielscheriPhelsuma flavigularis
Phelsuma ravenalaPhelsuma dubiaPhelsuma modesta
Phelsuma nigristriataPhelsuma inexpectata
Phelsuma ornataPhelsuma guimbeaui
Phelsuma cepedianaPhelsuma borbonica
Phelsuma guentheriPhelsuma gigas
Phelsuma edwardnewtoniPhelsuma standingi
Phelsuma andamanensePhelsuma mutabilis
Phelsuma brevicepsPhelsuma pronkiPhelsuma barbouri
Phelsuma seippiPhelsuma parkeri
Phelsuma abbottiPhelsuma madagascariensis
Phelsuma guttataPhelsuma sundbergiPhelsuma astriata
Phelsuma vanheygeniLygodactylus keniensis
Lygodactylus kimhowelliLygodactylus luteopicturatusLygodactylus picturatus
Lygodactylus williamsiLygodactylus chobiensis
Lygodactylus gutturalisLygodactylus angularis
Lygodactylus thomensisLygodactylus conraui
Lygodactylus klugeiLygodactylus bradfieldiLygodactylus capensisLygodactylus stevensoni
Lygodactylus lawrenceiLygodactylus pictus
Lygodactylus mirabilisLygodactylus tuberosus
Lygodactylus montanusLygodactylus gravis
Lygodactylus paulianiLygodactylus arnoulti
Lygodactylus blancaeLygodactylus verticillatusLygodactylus heterurus
Lygodactylus tolampyaeLygodactylus guibei
Lygodactylus miopsLygodactylus madagascariensis
Lygodactylus rarusLygodactylus expectatus
Rhoptropella ocellataCnemaspis tropidogaster
Cnemaspis kandianaCnemaspis podihuna
Pachydactylus affinisPachydactylus vansoni
Pachydactylus capensisPachydactylus oshaughnessyi
Pachydactylus tigrinusPachydactylus montanus
Pachydactylus vanzyliPachydactylus rangei
Pachydactylus austeniPachydactylus mariquensis
Pachydactylus mclachlaniPachydactylus monicaePachydactylus weberi
Pachydactylus servalPachydactylus griffini
Pachydactylus carinatusPachydactylus fasciatus
Pachydactylus tsodiloensisPachydactylus waterbergensis
Pachydactylus purcelliPachydactylus oculatus
Pachydactylus maculatusPachydactylus geitje
Pachydactylus labialisPachydactylus barnardi
Pachydactylus formosusPachydactylus rugosus
Pachydactylus punctatusPachydactylus scherzi
Pachydactylus sansteynaePachydactylus caraculicus
Pachydactylus bicolorPachydactylus oreophilus
Pachydactylus gaiasensisPachydactylus reconditus
Pachydactylus parascutatusPachydactylus scutatus
Pachydactylus namaquensisPachydactylus kladaroderma
Pachydactylus haackeiColopus kochii
Colopus wahlbergiiPachydactylus robertsi
Chondrodactylus bibroniiChondrodactylus fitzsimonsi
Chondrodactylus turneriPachydactylus laevigatus
Chondrodactylus anguliferElasmodactylus tuberculosus
Elasmodactylus tetensisRhoptropus barnardi
Rhoptropus biporosusRhoptropus boultoni
Rhoptropus bradfieldiRhoptropus afer
Goggia lineataBlaesodactylus sakalava
Blaesodactylus antongilensisBlaesodactylus boiviniHomopholis mulleri
Homopholis walbergiiHomopholis fasciataGeckolepis maculataGeckolepis typica
Afroedura karroicaAfroedura pondolia
Afrogecko porphyreusMatoatoa brevipes
Cryptactites peringueyiAfrogecko swartbergensis
Christinus marmoratusParagehyra gabriellae
Uroplatus fimbriatusUroplatus giganteus
Uroplatus henkeliUroplatus sikorae
Uroplatus lineatusUroplatus pietschmanni
Uroplatus alluaudiUroplatus phantasticus
Uroplatus ebenauiUroplatus malama
Uroplatus malaheloUroplatus guentheri
Cnemaspis africanaCnemaspis dickersonae
Cnemaspis uzungwaeNarudasia festiva
Ptenopus carpiCalodactylodes aureusCalodactylodes illingworthorum
Ailuronyx trachygasterAiluronyx seychellensis
Ailuronyx tachyscopaeusParoedura bastardi
Paroedura tanjakaParoedura vazimba
Paroedura androyensisParoedura picta
Paroedura sanctijohannisParoedura stumpffi
Paroedura lohatsaraParoedura karstophila
Paroedura ovicepsParoedura homalorhina
Paroedura gracilisParoedura masobe
Ebenavia inunguisUrocotyledon inexpectata
Perochirus ateles
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(ii)
Hemidactylus homoeolepisHemidactylus forbesii
Hemidactylus oxyrhinusHemidactylus macropholis
Hemidactylus mindiaeHemidactylus lemurinus
Hemidactylus turcicusHemidactylus robustus
Hemidactylus yerburiiHemidactylus persicus
Hemidactylus grantiHemidactylus dracaenacolus
Hemidactylus pumilioHemidactylus foudaii
Hemidactylus citerniiHemidactylus modestus
Hemidactylus agriusHemidactylus palaichthusHemidactylus bouvieri
Hemidactylus brasilianusHemidactylus greeffiiHemidactylus platycephalus
Hemidactylus longicephalusHemidactylus mercatoriusHemidactylus mabouia
Hemidactylus haitianusHemidactylus angulatus
Hemidactylus gracilisHemidactylus reticulatus
Hemidactylus albofasciatusHemidactylus imbricatus
Hemidactylus sataraensisHemidactylus brookii
Hemidactylus frenatusHemidactylus flaviviridis
Hemidactylus leschenaultiiHemidactylus maculatus
Hemidactylus prashadiHemidactylus triedrus
Hemidactylus depressusHemidactylus giganteus
Hemidactylus aaronbaueriHemidactylus fasciatus
Hemidactylus platyurusHemidactylus karenorum
Hemidactylus garnotiiHemidactylus bowringii
Cyrtodactylus louisiadensisCyrtodactylus epiroticusCyrtodactylus tripartitus
Cyrtodactylus robustusCyrtodactylus klugei
Cyrtodactylus tuberculatusCyrtodactylus novaeguineae
Cyrtodactylus loriaeCyrtodactylus sermowaiensis
Cyrtodactylus pulchellusCyrtodactylus intermedius
Cyrtodactylus agusanensisCyrtodactylus philippinicus
Cyrtodactylus annulatusCyrtodactylus consobrinus
Cyrtodactylus irregularisCyrtodactylus marmoratus
Cyrtodactylus triedrusCyrtodactylus jarujini
Cyrtodactylus angularisCyrtodactylus ayeyarwadyensis
Cyrtodactylus oldhamiCyrtopodion gastropholeCyrtopodion agamuroides
Cyrtopodion caspiumCyrtopodion longipes
Cyrtopodion scabrumCyrtopodion sistanensis
Agamura persicaBunopus crassicauda
Crossobamon orientalisBunopus tuberculatus
Mediodactylus heteropholisMediodactylus heterocercum
Mediodactylus sagittiferumMediodactylus kotschyi
Stenodactylus sthenodactylusStenodactylus yemenensis
Stenodactylus doriaeStenodactylus leptocosymbotus
Stenodactylus petriiStenodactylus arabicus
Tropiocolotes tripolitanusPseudoceramodactylus khobarensis
Mediodactylus russowiiMediodactylus spinicaudum
Cnemaspis kendalliiCnemaspis limi
Tropiocolotes helenaeAlsophylax pipiens
Gehyra minutaGehyra montium
Gehyra pilbaraGehyra punctata
Gehyra variegataGehyra purpurascensGehyra nana
Gehyra xenopusGehyra australis
Gehyra occidentalisGehyra koira
Gehyra robustaGehyra borroloola
Gehyra pamelaGehyra catenata
Gehyra dubiaGehyra membranacruralis
Gehyra oceanicaGehyra marginata
Gehyra bareaGehyra baliola
Gehyra brevipalmataGehyra lacerata
Gehyra mutilataGehyra fehlmanni
Hemiphyllodactylus aurantiacusHemiphyllodactylus yunnanensis
Hemiphyllodactylus typusNactus pelagicusNactus multicarinatus
Nactus chevertiNactus galgajuga
Nactus vankampeniNactus eboracensisNactus acutus
Heteronotia speleaHeteronotia binoeiHeteronotia planiceps
Dixonius siamensisDixonius vietnamensis
Dixonius melanostictusGekko grossmanni
Gekko badeniiGekko petricolus
Gekko vittatusLuperosaurus iskandari
Ptychozoon lionotumPtychozoon kuhli
Ptychozoon rhacophorusGekko porosusGekko crombota
Gekko romblonGekko mindorensis
Gekko monarchusGekko athymus
Gekko swinhonisGekko auriverrucosus
Gekko hokouensisGekko japonicus
Gekko chinensisGekko gecko
Gekko smithiiLepidodactylus moestusLepidodactylus lugubris
Luperosaurus joloensisLuperosaurus cumingii
Luperosaurus macgregoriLepidodactylus orientalis
Pseudogekko compressicorpusPseudogekko smaragdinus
Lepidodactylus novaeguineae
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Gekko
nid
ae
(i)
100
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Figure 4
DCordylus cordylusCordylus tasmani
Cordylus oelofseniCordylus mclachlani
Cordylus nigerCordylus minorCordylus aridusCordylus imkeae
Cordylus macropholisCordylus rhodesianus
Cordylus jonesiiCordylus beraducciiCordylus ukingensisCordylus vittifer
Cordylus meculaeCordylus tropidosternum
Hemicordylus capensisHemicordylus nebulosusNinurta coeruleopunctatusPseudocordylus melanotusPseudocordylus spinosus
Pseudocordylus microlepidotusPseudocordylus langi
Chamaesaura anguinaChamaesaura aenea
Smaug giganteusSmaug warreni
Namazonurus peersiNamazonurus lawrenci
Namazonurus namaquensisNamazonurus pustulatus
Namazonurus campbelliKarusasaurus polyzonus
Karusasaurus jordaniOuroborus cataphractus
Platysaurus monotropisPlatysaurus minor
Platysaurus intermediusPlatysaurus capensis
Platysaurus broadleyiPlatysaurus mitchelli
Platysaurus pungweensisZonosaurus rufipes
Zonosaurus aeneusZonosaurus bemaraha
Zonosaurus subunicolorZonosaurus brygooiZonosaurus tsingy
Zonosaurus boettgeriZonosaurus laticaudatus
Zonosaurus anelanelanyZonosaurus karsteni
Zonosaurus quadrilineatusZonosaurus trilineatus
Zonosaurus ornatusZonosaurus madagascariensis
Zonosaurus haraldmeieriTracheloptychus petersi
Tracheloptychus madagascariensisGerrhosaurus flavigularis
Gerrhosaurus multilineatusGerrhosaurus nigrolineatus
Gerrhosaurus typicusGerrhosaurus skoogi
Gerrhosaurus validusTetradactylus tetradactylus
Tetradactylus africanusTetradactylus seps
Cordylosaurus subtessellatusGerrhosaurus major
Lepidophyma dontomasiLepidophyma radula
Lepidophyma loweiLepidophyma cuicatecaLepidophyma smithii
Lepidophyma lineriLepidophyma gaigeae
Lepidophyma sylvaticumLepidophyma micropholis
Lepidophyma occulorLepidophyma pajapanensis
Lepidophyma reticulatumLepidophyma flavimaculatumLepidophyma lipetziLepidophyma tuxtlae
Lepidophyma mayaeXantusia wigginsi
Xantusia bezyiXantusia vigilis
Xantusia arizonaeXantusia riversiana
Xantusia sancheziXantusia bolsonaeXantusia henshawi
Xantusia gracilisCricosaura typica
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Platysaurinae
Cordylinae
Cordylidae
Gerrhosauridae
Gerrhosaurinae
Zonosaurinae
Lepidophyminae
Xantusiinae
Cricosaurinae
Xantusiidae
Figure 5
EAmphiglossus frontoparietalisAmphiglossus punctatus
Amphiglossus macrocercusAmphiglossus anosyensis
Amphiglossus splendidusVoeltzkowia rubrocaudataVoeltzkowia lineata
Voeltzkowia fierinensisAmphiglossus astrolabi
Amphiglossus reticulatusAmphiglossus tsaratananensis
Androngo trivittatusPygomeles braconnieri
Amphiglossus tanysomaAmphiglossus mandokava
Amphiglossus ornaticepsAmphiglossus melanurus
Madascincus intermediusMadascincus polleni
Madascincus stumpffiMadascincus nanus
Madascincus mouroundavaeMadascincus igneocaudatus
Pseudoacontias menamaintyMadascincus melanopleura
Paracontias rothschildiParacontias hildebrandti
Paracontias manifyParacontias brocchii
Paracontias holomelasScelotes kasneri
Scelotes montispectusScelotes gronovii
Scelotes sexlineatusScelotes bipes
Scelotes arenicolusScelotes mirus
Scelotes anguineusScelotes caffer
Proscelotes eggeliHakaria simonyi
Melanoseps loveridgeiMelanoseps ater
Melanoseps occidentalisFeylinia currori
Feylinia polylepisFeylinia grandisquamis
Typhlacontias punctatissimusTyphlacontias brevipes
Sepsina angolensisChalcides montanus
Chalcides polylepisChalcides manueli
Chalcides mionectonChalcides sphenopsiformis
Chalcides viridanusChalcides coeruleopunctatus
Chalcides sexlineatusChalcides lanzaiChalcides parallelus
Chalcides boulengeriChalcides bedriagaiChalcides colosii
Chalcides sepsoidesChalcides ocellatus
Chalcides striatusChalcides pseudostriatusChalcides chalcides
Chalcides guentheriChalcides minutus
Chalcides mauritanicusGongylomorphus bojerii
Janetaescincus braueriJanetaescincus veseyfitzgeraldi
Pamelaescincus gardineriScincus mitranusScincus scincus
Eumeces algeriensisScincopus fasciatus
Eumeces schneideriEurylepis taeniolatus
Plestiodon dugesiiPlestiodon brevirostris
Plestiodon copeiPlestiodon parvulus
Plestiodon ochoterenaePlestiodon lynxe
Plestiodon sumichrastiPlestiodon parviauriculatus
Plestiodon skiltonianusPlestiodon lagunensis
Plestiodon gilbertiPlestiodon longirostris
Plestiodon septentrionalisPlestiodon multivirgatus
Plestiodon fasciatusPlestiodon obsoletus
Plestiodon callicephalusPlestiodon tetragrammus
Plestiodon laticepsPlestiodon inexpectatus
Plestiodon anthracinusPlestiodon egregius
Plestiodon reynoldsiPlestiodon stimpsonii
Plestiodon marginatusPlestiodon elegans
Plestiodon latiscutatusPlestiodon japonicusPlestiodon barbouri
Plestiodon capitoPlestiodon tunganus
Plestiodon quadrilineatusPlestiodon chinensis
Plestiodon kishinouyeiPlestiodon tamdaoensis
Brachymeles pathfinderiBrachymeles gracilis
Brachymeles bicolorBrachymeles boulengeri
Brachymeles schadenbergiBrachymeles eleraeBrachymeles talinis
Brachymeles samarensisBrachymeles cebuensis
Brachymeles minimusBrachymeles bonitae
Brachymeles tridactylusBrachymeles miriamae
Brachymeles apusOphiomorus latastiiOphiomorus punctatissimus
Mesoscincus managuaeMesoscincus schwartzei
Acontias poecilusAcontias plumbeus
Acontias gracilicaudaAcontias breviceps
Acontias rieppeliAcontias percivali
Acontias meleagrisAcontias kgalagadi
Acontias litoralisAcontias lineatus
Acontias gariepensisTyphlosaurus lomiaeTyphlosaurus vermisTyphlosaurus caecusTyphlosaurus meyeri
Typhlosaurus braini
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Scincidae
Acontiinae
Scincinae
F-I
Figure 6
Lygosominae
Lygosominae
cont.FPapuascincus stanleyanus
Lipinia noctua
Lipinia pulchella
Insulasaurus victoria
Insulasaurus wrighti
Insulasaurus arborens
Prasinohaema virens
Sphenomorphus buenloicus
Scincella assatus
Scincella cherriei
Scincella gemmingeri
Scincella lateralis
Asymblepharus sikimmensis
Scincella reevesii
Sphenomorphus sabanus
Sphenomorphus cyanolaemus
Sphenomorphus variegatus
Sphenomorphus multisquamatus
Sphenomorphus indicus
Sphenomorphus maculatus
Larutia seribuatensis
Parvoscincus lawtoni
Parvoscincus kitangladensis
Parvoscincus luzonense
Parvoscincus laterimaculatus
Parvoscincus beyeri
Parvoscincus sisoni
Parvoscincus tagapayo
Parvoscincus leucospilos
Parvoscincus decipiens
Parvoscincus steerei
Sphenomorphus acutus
Sphenomorphus diwata
Pinoyscincus llanosi
Pinoyscincus abdictus
Pinoyscincus coxi
Pinoyscincus jagori
Pinoyscincus mindanensis
Otosaurus cumingi
Sphenomorphus maindroni
Sphenomorphus cranei
Sphenomorphus leptofasciatus
Sphenomorphus fasciatus
Sphenomorphus solomonis
Sphenomorphus scutatus
Sphenomorphus concinnatus
Tytthoscincus atrigularis
Tytthoscincus hallieri
Tytthoscincus parvus
Tytthoscincus aesculeticola
Sphenomorphus muelleri
Sphenomorphus jobiensis
Sphenomorphus melanopogon
Isopachys anguinoides
Lipinia vittigera
Tropidophorus misaminius
Tropidophorus partelloi
Tropidophorus brookei
Tropidophorus beccarii
Tropidophorus baconi
Tropidophorus grayi
Tropidophorus microlepis
Tropidophorus cocincinensis
Tropidophorus thai
Tropidophorus robinsoni
Tropidophorus sinicus
Tropidophorus baviensis
Tropidophorus hainanus
Tropidophorus murphyi
Tropidophorus noggei
Tropidophorus latiscutatus
Tropidophorus matsuii
Tropidophorus berdmorei
Sphenomorphus simus
Sphenomorphus praesignis
Ablepharus pannonicus
Asymblepharus alaicus
Ateuchosaurus pellopleurus
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H-I GFigure 7
Lerista kennedyensisLerista onsloviana
Lerista uniduoLerista connivens
Lerista variaLerista lineopunctulata
Lerista yunaLerista kendricki
Lerista nichollsiLerista gascoynensis
Lerista petersoniLerista humphriesi
Lerista praepeditaLerista bougainvillii
Lerista viduataLerista muelleri
Lerista haroldiLerista allochira
Lerista stictopleuraLerista planiventralis
Lerista lineataLerista christinae
Lerista distinguendaLerista elegans
Lerista emmottiLerista punctatovittata
Lerista dorsalisLerista terdigitata
Lerista tridactylaLerista elongata
Lerista speciosaLerista zietzi
Lerista flammicaudaLerista picturata
Lerista baynesiLerista edwardsae
Lerista arenicolaLerista microtis
Lerista macropisthopusLerista axillaris
Lerista gerrardiiLerista eupoda
Lerista desertorumLerista puncticaudaLerista neander
Lerista zonulataLerista ingrami
Lerista orientalisLerista taeniata
Lerista aericepsLerista xanthura
Lerista chordaeLerista fragilis
Lerista frostiLerista borealis
Lerista walkeriLerista simillima
Lerista vermicularisLerista robusta
Lerista greeriLerista labialis
Lerista bipesLerista ips
Lerista griffiniLerista apoda
Lerista kalumburuLerista wilkinsiLerista cinerea
Lerista amelesLerista carpentariae
Lerista karlschmidtiLerista stylis
Ctenotus pulchellusCtenotus gagudju
Ctenotus hilliCtenotus hebetior
Ctenotus essingtoniiCtenotus piankai
Ctenotus hanloniCtenotus angusticeps
Ctenotus grandisCtenotus atlas
Ctenotus maryaniCtenotus olympicus
Ctenotus regiusCtenotus septenarius
Ctenotus astarteCtenotus mimetes
Ctenotus tanamiensisCtenotus greeri
Ctenotus serventyiCtenotus quattuordecimlineatus
Ctenotus leonhardiiCtenotus leae
Ctenotus australisCtenotus uber
Ctenotus rutilansCtenotus taeniolatus
Ctenotus robustusCtenotus spaldingi
Ctenotus inornatusCtenotus saxatilis
Ctenotus fallensCtenotus rawlinsoni
Ctenotus calurusCtenotus schomburgkiiCtenotus youngsoni
Ctenotus strauchiiCtenotus pantherinus
Ctenotus nasutusCtenotus rubicundus
Ctenotus brooksiCtenotus labillardieri
Notoscincus ornatusEulamprus tympanum
Eulamprus heatwoleiEulamprus kosciuskoi
Eulamprus leuraensisEulamprus quoyii
Glaphyromorphus darwiniensisGlaphyromorphus cracens
Glaphyromorphus pumilusGlaphyromorphus fuscicaudis
Glaphyromorphus mjobergiGlaphyromorphus punctulatus
Hemiergis decresiensisHemiergis millewae
Hemiergis gracilipesHemiergis quadrilineatum
Hemiergis peroniiHemiergis initialis
Eremiascincus isolepisEremiascincus douglasiEremiascincus richardsoniiEremiascincus pardalis
Eremiascincus fasciolatusAnomalopus leuckartii
Anomalopus verreauxiAnomalopus mackayi
Anomalopus swansoniOphioscincus truncatus
Coeranoscincus reticulatusSaiphos equalis
Ophioscincus ophioscincusCoeranoscincus frontalis
Coggeria naufragusEulamprus murrayi
Eulamprus tryoniEulamprus luteilateralis
Eulamprus brachyosomaEulamprus sokosoma
Eulamprus martiniEulamprus tigrinus
Eulamprus tenuisEulamprus amplus
Gnypetoscincus queenslandiaeCalyptotis ruficauda
Calyptotis scutirostrumCalyptotis lepidorostrum
Nangura spinosaEulamprus frerei
100
95
55
100
85
82
87
90
93
93
89
98
84
10096
100100
1008177
56
100
100
93
8996
90
92
100
90
100
61100
10099
100
100
100
98
90
100
99
10093
79
87
100
99
100100
100
96
8997
99
91100
99
9398
90 100
88
100
9997
95
81
99
95
85
67
76
95
86
66
51
85
78
9165
90
7992
97
83
8873
63
100
100
88
89
84
85
99
81
100
76
70
93
78
93 100
8287
95
99
91
100
8260
100100
10097
6288
100100
97
80
91
95
95
77
85
83
98
7896
88 100
10077
G
Lygosominae
cont.
Figure 8
Mabuya heathiMabuya caissara
Mabuya agilisMabuya guaporicola
Mabuya macrorhynchaMabuya agmosticha
Mabuya frenataMabuya bistriata
Mabuya falconensisMabuya unimarginata
Mabuya mabouyaMabuya meridensis
Mabuya dorsivittataMabuya cochabambae
Mabuya nigropunctataMabuya altamazonica
Mabuya sloaniiMabuya nigropalmata
Mabuya croizatiMabuya carvalhoiChioninia stangeri
Chioninia fogoensisChioninia spinalis
Chioninia cocteiChioninia delalandii
Chioninia vaillantiiEumecia anchietae
Trachylepis dumasiTrachylepis aureopunctata
Trachylepis vatoTrachylepis boettgeriTrachylepis madagascariensis
Trachylepis elegansTrachylepis gravenhorstiiTrachylepis acutilabris
Trachylepis homalocephalaTrachylepis sulcata
Trachylepis variegataTrachylepis hoeschi
Trachylepis striataTrachylepis spilogaster
Trachylepis occidentalisTrachylepis capensis
Trachylepis variaTrachylepis atlantica
Trachylepis margaritiferaTrachylepis quinquetaeniata
Trachylepis perrotetiiTrachylepis affinis
Trachylepis sechellensisTrachylepis wrightii
Trachylepis maculilabrisTrachylepis socotrana
Trachylepis brevicollisTrachylepis vittata
Trachylepis aurataDasia olivacea
Dasia griseaDasia vittata
Eutropis trivittataEutropis nagarjuniEutropis beddomii
Eutropis bibroniiEutropis clivicola
Eutropis multicarinataEutropis cumingi
Eutropis multifasciataEutropis macrophthalma
Eutropis rudisEutropis macularia
Eutropis longicaudataLankascincus fallax
Ristella rurkiiLiopholis montana
Liopholis guthegaLiopholis whitiiLiopholis margaretae
Liopholis modestaLiopholis pulchra
Liopholis kintoreiLiopholis multiscutata
Liopholis inornataLiopholis striata
Tiliqua scincoidesTiliqua gigas
Tiliqua nigroluteaTiliqua occipitalis
Tiliqua rugosaTiliqua adelaidensisCyclodomorphus branchialis
Cyclodomorphus casuarinaeCyclodomorphus michaeli
Egernia hosmeriEgernia stokesii
Lissolepis luctuosaEgernia napoleonis
Egernia richardiBellatorias frerei
Egernia kingiiEgernia depressa
Bellatorias majorEgernia saxatilis
Corucia zebrataTribolonotus pseudoponceleti
Tribolonotus ponceletiTribolonotus brongersmai
Tribolonotus schmidtiTribolonotus blanchardi
Tribolonotus gracilisTribolonotus novaeguineae
83
97
91
99
87
100
98
89
81
70
7780
10099
100
97
96
95
100
78
90
98
80
100
91
98
92
84
90
74
98
10097
95
10095
95
54
78
89
99100
100
100
100
99
83100
99
93
97
95
90
97
96 100
80
73
100
96
88
94
93
100
96
91
99
5652
90
73
9890
85
88
97
84
71
100
81
95
66
86
74
73
90
77
100
8697
93 96
100
Lygosominae
cont.
HFigure 9
Carlia vivaxCarlia dogare
Carlia rostralisCarlia amax
Carlia tetradactylaCarlia rubrigularis
Carlia rhomboidalisCarlia rufilatus
Carlia pectoralisCarlia munda
Carlia mysiCarlia longipes
Carlia fuscaCarlia storri
Carlia schmeltziiLygisaurus malleolus
Lygisaurus tanneriLygisaurus absconditaLygisaurus foliorum
Lygisaurus laevisLygisaurus aeratus
Lygisaurus novaeguineaeLygisaurus macfarlani
Lygisaurus parrhasiusLygisaurus sesbrauna
Carlia gracilisCarlia bicarinata
Carlia jarnoldaeCarlia johnstonei
Carlia triacanthaMenetia timlowi
Liburnascincus mundivensisLiburnascincus scirtetis
Liburnascincus coensisCautula zia
Niveoscincus metallicusNiveoscincus pretiosus
Niveoscincus ocellatusNiveoscincus greeni
Proablepharus reginaeLampropholis robertsi
Lampropholis coggeriLampropholis delicata
Lampropholis guichenotiSaproscincus spectabilis
Saproscincus roseiSaproscincus challengeri
Saproscincus mustelinusSaproscincus oriarus
Saproscincus hannahaeSaproscincus tetradactylus
Saproscincus czechuraiSaproscincus lewisi
Saproscincus basiliscusBartleia jigurru
Morethia adelaidensisMorethia butleri
Morethia ruficaudaBassiana trilineata
Emoia isolataEmoia pseudocyanura
Emoia cyanuraEmoia impar
Emoia physicaeEmoia jakati
Emoia atrocostataEmoia caeruleocauda
Emoia cyanogasterEmoia schmidti
Menetia alanaeMenetia greyii
Cryptoblepharus novocaledonicusCryptoblepharus boutonii
Cryptoblepharus nigropunctatusCaledoniscincus terma
Caledoniscincus aquiloniusCaledoniscincus chazeaui
Caledoniscincus atropunctatusCaledoniscincus haplorhinus
Caledoniscincus austrocaledonicusCaledoniscincus festivus
Caledoniscincus auratusCaledoniscincus renevieri
Caledoniscincus orestesSimiscincus aurantiacus
Graciliscincus shonaeTropidoscincus variabilis
Tropidoscincus boreus
Tropidoscincus aubrianusLioscincus tillieri
Lioscincus maruiaLioscincus novaecaledoniaeSigaloseps deplanchei
Sigaloseps ruficaudaPhoboscincus garnieri
Lacertoides pardalisLioscincus nigrofasciolatum
Kanakysaurus viviparusLioscincus steindachneri
Lioscincus vivaeCelatiscincus euryotis
Celatiscincus similisMarmorosphax tricolor
Marmorosphax montanaNannoscincus humectus
Nannoscincus hanchisteusNannoscincus greeriNannoscincus mariei
Nannoscincus gracilisNannoscincus slevini
Nannoscincus garrulusOligosoma whitakeri
Oligosoma oliveriOligosoma townsi
Oligosoma ornatumOligosoma macgregori
Oligosoma mocoOligosoma alani
Oligosoma fallaiOligosoma hardyi
Oligosoma levidensumOligosoma aeneum
Oligosoma microlepisOligosoma smithi
Oligosoma striatumOligosoma homalonotumOligosoma zelandicum
Oligosoma notosaurusOligosoma inconspicuumOligosoma maccanni
Oligosoma stenotisOligosoma grande
Oligosoma nigriplantareOligosoma longipes
Oligosoma lineoocellatumOligosoma chloronoton
Oligosoma otagenseOligosoma waimatenseOligosoma infrapunctatum
Oligosoma acrinasumOligosoma taumakaeOligosoma pikitanga
Oligosoma suteriOligosoma lichenigera
Pseudemoia entrecasteauxiiBassiana duperreyi
Pseudemoia pagenstecheriLeptosiaphos vigintiserierumLeptosiaphos kilimensis
Leptosiaphos amietiLeptosiaphos graueri
Leptosiaphos hackarsiLacertaspis lepesmei
Lacertaspis gemmiventrisLacertaspis chriswildi
Lacertaspis rohdeiLacertaspis reichenowi
Afroablepharus africanusAfroablepharus annobonensis
Afroablepharus wahlbergiPanaspis togoensis
Panaspis brevicepsLeiolopisma mauritianaLeiolopisma telfairii
Emoia loyaltiensisEugongylus albofasciolatusEugongylus rufescens
Lygosoma bowringiiMochlus brevicaudisLygosoma punctata
Lygosoma sundevalliMochlus afer
Lygosoma lineolatumLepidothyris fernandi
Lygosoma albopunctataLygosoma koratense
Lygosoma quadrupesLamprolepis smaragdina
Emoia tonganaEmoia concolor
Sphenomorphus stellatusAblepharus kitaibelii
Ablepharus chernoviAblepharus budaki
90
100
100
84
52
79
100
72
87
88
98
90
88
68
80
85
7653
62
92
100
93
98100
73
91
75
98
85
78
89100
100 100
99
79
100
96
91
76
100
92100
72
76
71100 97
100
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98
96
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93
91
100
77
98 71 98
98100
100
93
100
79
61
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96
96
98
87
97
95
83
53
8795
98
95
97
100
100 100
80
8585
92
100
80
90
9696
100100
98
8293
94
98
89
100
98
90
94
57
9893
899796
63
93
99
100
100
100
86
99
8598
100
98
99
100
99
9891
100
95
9980
75
84
85
100
8195 99
95
9396
91
90
9999
85
100
100
100
100
63
60
9593
7670
92100
98
96
I
Lygosominae
cont.
Figure 10
Gymnophthalmus cryptusGymnophthalmus underwoodi
Gymnophthalmus speciosusGymnophthalmus vanzoiGymnophthalmus leucomystax
Gymnophthalmus pleeiPsilophthalmus paeminosus
Calyptommatus nicterusCalyptommatus leiolepis
Calyptommatus confusionibusCalyptommatus sinebrachiatus
Nothobachia ablepharaProcellosaurinus tetradactylusProcellosaurinus erythrocercus
Vanzosaura rubricaudaMicrablepharus maximiliani
Micrablepharus atticolusTretioscincus oriximinensis
Tretioscincus agilisAcratosaura mentalisStenolepis ridleyi
Colobosaura modestaIphisa elegans
Colobodactylus taunayiColobodactylus dalcyanus
Heterodactylus imbricatusRhachisaurus brachylepis
Bachia heteropaBachia intermediaBachia barbouriBachia bicolorBachia peruana
Bachia panopliaBachia trisanale
Bachia scolecoidesBachia huallagana
Bachia dorbignyiBachia bresslaui
Bachia flavescensPetracola ventrimaculatus
Proctoporus bolivianusProctoporus unsaacae
Proctoporus guentheriProctoporus pachyurus
Proctoporus subsolanusProctoporus sucullucu
Cercosaura eigenmanniCercosaura schreibersii
Cercosaura ocellataCercosaura argulus
Cercosaura oshaughnessyiCercosaura quadrilineata
Potamites juruazensisPotamites ecpleopus
Pholidobolus macbrydeiPholidobolus montium
Placosoma glabellumPlacosoma cordylinum
Neusticurus bicarinatusNeusticurus rudis
Riama simoterusRiama colomaromani
Riama unicolorLeposoma guianense
Leposoma southiLeposoma parietale
Leposoma osvaldoiLeposoma percarinatum
Arthrosaura kockiiArthrosaura reticulata
Anotosaura collarisColobosauroides cearensis
Leposoma annectansLeposoma scincoides
Leposoma pukLeposoma baturitensis
Leposoma nanodactylusEcpleopus gaudichaudii
Alopoglossus angulatusAlopoglossus copii
Alopoglossus atriventrisPtychoglossus brevifrontalis
Cnemidophorus lacertoidesCnemidophorus longicaudus
Dicrodon guttulatumAmeiva fuscataAmeiva erythrocephala
Ameiva griswoldiAmeiva pluvianotata
Ameiva coraxAmeiva plei
Ameiva maynardiAmeiva lineolata
Ameiva taeniuraAmeiva chrysolaema
Ameiva leberiAmeiva wetmorei
Ameiva exsulAmeiva polops
Ameiva dorsalisAmeiva auberi
Aspidoscelis ceralbensisAspidoscelis hyperythra
Aspidoscelis tigrisAspidoscelis marmorata
Aspidoscelis lineattissimaAspidoscelis guttata
Aspidoscelis deppeiAspidoscelis velox
Aspidoscelis laredoensisAspidoscelis gularis
Aspidoscelis costataAspidoscelis communis
Aspidoscelis burtiAspidoscelis sexlineata
Aspidoscelis inornataCnemidophorus arenivagus
Cnemidophorus lemniscatusCnemidophorus gramivagus
Cnemidophorus vanzoiAmeiva festivaAmeiva quadrilineataAmeiva undulata
Kentropyx viridistrigaKentropyx vanzoi
Kentropyx paulensisKentropyx pelviceps
Kentropyx altamazonicaKentropyx striata
Kentropyx calcarataCnemidophorus ocellifer
Ameiva bifrontataAmeiva ameivaTeius teyou
Dracaena guianensisCrocodilurus amazonicus
Tupinambis teguixinTupinambis longilineus
Tupinambis quadrilineatusTupinambis duseni
Tupinambis merianaeTupinambis rufescens
Callopistes maculatusCallopistes flavipunctatus
100
98
100
84
99
100
100
50
100
10087
100
93
8893
86
72
9861
100
100
98
100
9196
100
93
10077
10088
9795 100
7874
98
100
99
80
100
97
55
98
94 99
100
98
99
82
100
92
100
98100
94
100
91
100
97
99
65
10095
96
86
100
10098
97
100100
99
54
100
56
93
98
92
99
59
100
94
8597
55
98
93
100
97
9695 100
100
86
100
99
84
98
100
98
99
93 99
8183
100
99
94
97 100
82
100
9196
71
99
100
97
9887
6691
100
97
J
Teiidae
Gymnophthalmidae
Tupinambinae
Teiinae
Alopoglossinae
Ecpleopinae
Cercosaurinae
Bachiinae
Rhachisaurinae
Gymnophthalminae
Figure 11
Amphisbaena xera
Amphisbaena bakeri
Amphisbaena caeca
Amphisbaena fenestrata
Amphisbaena manni
Amphisbaena schmidti
Amphisbaena carlgansi
Amphisbaena cubana
Amphisbaena barbouri
Amphisbaena anaemariae
Amphisbaena silvestrii
Amphisbaena leeseri
Amphisbaena angustifrons
Amphisbaena darwini
Amphisbaena munoai
Amphisbaena kingii
Amphisbaena cuiabana
Amphisbaena hastata
Amphisbaena infraorbitale
Amphisbaena microcephalum
Amphisbaena polystegum
Amphisbaena anomala
Amphisbaena bolivica
Amphisbaena camura
Amphisbaena alba
Amphisbaena vermicularis
Amphisbaena roberti
Amphisbaena kraoh
Amphisbaena saxosa
Amphisbaena ignatiana
Amphisbaena leali
Amphisbaena innocens
Amphisbaena hyporissor
Amphisbaena mertensii
Amphisbaena cunhai
Amphisbaena fuliginosa
Amphisbaena brasiliana
Monopeltis capensis
Geocalamus acutus
Chirindia swynnertoni
Cynisca leucura
Trogonophis wiegmanni
Diplometopon zarudnyi
Cadea blanoides
Blanus tingitanus
Blanus mettetali
Blanus cinereus
Blanus strauchi
Bipes canaliculatus
Bipes biporus
Bipes tridactylus
Rhineura floridana
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100
84
100
91
86
89
95
90
92
87
100
100
97
100
100
90
78100
93
99
100
100
100
81
100
98
95100
100
91
100
55
96
67
95
89
100
100
100
10097
100
73
Blanidae
Cadeidae
Trogonophiidae
Amphisbaenidae
K
Bipedidae
Rhineuridae
Figure 12
Darevskia derjuginiDarevskia caucasica
Darevskia daghestanicaDarevskia mixta
Darevskia clarkorumDarevskia armeniaca
Darevskia bendimahiensisDarevskia sapphirina
Darevskia uzzelliDarevskia raddei
Darevskia rostombekoviDarevskia saxicola
Darevskia brauneriDarevskia lindholmi
Darevskia alpinaDarevskia chlorogaster
Darevskia praticolaDarevskia valentini
Darevskia portschinskiiDarevskia rudis
Darevskia parvulaIranolacerta zagrosicaIranolacerta brandtii
Algyroides fitzingeriAlgyroides marchi
Dinarolacerta montenegrinaDinarolacerta mosorensis
Algyroides nigropunctatusAlgyroides moreoticus
Anatololacerta danfordiAnatololacerta oertzeni
Anatololacerta anatolicaParvilacerta parva
Parvilacerta fraasiiIberolacerta aranicaIberolacerta bonnali
Iberolacerta aurelioiIberolacerta horvathi
Iberolacerta monticolaIberolacerta cyreni
Iberolacerta galaniApathya cappadocica
Archaeolacerta bedriagaeHellenolacerta graecaDalmatolacerta oxycephala
Podarcis carbonelliPodarcis bocagei
Podarcis hispanicusPodarcis vaucheri
Podarcis liolepisPodarcis peloponnesiacusPodarcis erhardii
Podarcis raffoneaePodarcis lilfordi
Podarcis pityusensisPodarcis siculus
Podarcis muralisPodarcis tiliguerta
Podarcis filfolensisPodarcis gaigeae
Podarcis milensisPodarcis tauricus
Podarcis melisellensisScelarcis perspicillata
Teira dugesiiLacerta pamphylicaLacerta trilineata
Lacerta mediaLacerta agilis
Lacerta schreiberiLacerta strigata
Lacerta bilineataLacerta viridis
Timon tangitanusTimon lepidusTimon pater
Timon princepsTakydromus stejnegeriTakydromus septentrionalisTakydromus toyamaiTakydromus wolteriTakydromus formosanus
Takydromus hsuehshanensisTakydromus amurensis
Takydromus sylvaticusTakydromus dorsalis
Takydromus intermediusTakydromus sauteri
Takydromus smaragdinusTakydromus tachydromoides
Takydromus sexlineatusTakydromus kuehnei
Zootoca viviparaPhoenicolacerta cyanisparsa
Phoenicolacerta laevisPhoenicolacerta kulzeri
Acanthodactylus schreiberiAcanthodactylus boskianus
Acanthodactylus opheodurusAcanthodactylus gongrorhynchatus
Acanthodactylus masiraeAcanthodactylus cantoris
Acanthodactylus schmidtiAcanthodactylus longipes
Acanthodactylus scutellatusAcanthodactylus aureus
Acanthodactylus maculatusAcanthodactylus pardalis
Acanthodactylus beershebensisAcanthodactylus busacki
Acanthodactylus erythrurusAcanthodactylus blanci
Acanthodactylus tristramiAcanthodactylus orientalisOphisops elegans
Ophisops occidentalisMesalina rubropunctata
Mesalina brevirostrisMesalina adramitana
Mesalina balfouriMesalina guttulata
Mesalina bahaeldiniMesalina olivieri
Mesalina simoniOmanosaura jayakariOmanosaura cyanura
Congolacerta vauereselliEremias grammica
Eremias pleskeiEremias multiocellata
Eremias przewalskiiEremias arguta
Eremias nigrolateralisEremias persica
Eremias montanusEremias velox
Eremias vermiculataEremias argus
Eremias brenchleyiAdolfus jacksoni
Adolfus alleniAdolfus africanus
Gastropholis prasinaGastropholis vittata
Holaspis guentheriHolaspis laevis
Pedioplanis gaerdesiPedioplanis inornata
Pedioplanis rubensPedioplanis undata
Pedioplanis husabensisPedioplanis namaquensis
Pedioplanis brevicepsPedioplanis burchelli
Pedioplanis laticepsNucras tessellata
Pedioplanis lineoocellataMeroles cuneirostrisMeroles micropholidotus
Meroles ctenodactylusMeroles anchietaeIchnotropis squamulosa
Meroles suborbitalisMeroles knoxii
Meroles reticulatusIchnotropis capensis
Tropidosaura gularisAustralolacerta australis
Heliobolus spekiiHeliobolus lugubris
Pseuderemias smithiiPhilochortus spinalis
Latastia longicaudataNucras lalandii
Poromera fordiiAtlantolacerta andreanskyiGallotia bravoana
Gallotia simonyiGallotia intermedia
Gallotia caesarisGallotia galloti
Gallotia atlanticaGallotia stehlini
Psammodromus blanciPsammodromus algirus
Psammodromus hispanicus
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Lacertidae
Gallotiinae
Lacertinae
LFigure 13
Varanus baritjiVaranus acanthurus
Varanus storriVaranus primordius
Varanus kingorumVaranus bushiVaranus gilleni
Varanus caudolineatusVaranus eremius
Varanus brevicaudaVaranus semiremex
Varanus mitchelliVaranus timorensis
Varanus scalarisVaranus glauerti
Varanus tristisVaranus pilbarensis
Varanus glebopalmaVaranus komodoensis
Varanus variusVaranus salvadoriiVaranus gouldiiVaranus panoptes
Varanus rosenbergiVaranus giganteus
Varanus spenceriVaranus mertensi
Varanus bengalensisVaranus flavescens
Varanus dumeriliiVaranus rudicollis
Varanus salvatorVaranus marmoratus
Varanus finschiVaranus doreanusVaranus yuwonoiVaranus jobiensis
Varanus caerulivirensVaranus cerambonensisVaranus melinusVaranus indicus
Varanus rainerguentheriVaranus prasinusVaranus macraeiVaranus boehmei
Varanus beccariiVaranus keithhornei
Varanus olivaceusVaranus yemenensis
Varanus albigularisVaranus exanthematicusVaranus niloticus
Varanus griseusLanthanotus borneensis
Shinisaurus crocodilurusAbronia auritaAbronia campbelliAbronia anzuetoiAbronia matudai
Abronia fimbriataAbronia lythrochilaAbronia ornelasi
Abronia frostiAbronia graminea
Abronia oaxacaeAbronia chiszari
Mesaspis gadoviiMesaspis moreletiiBarisia rudicollis
Barisia herreraeBarisia imbricataBarisia levicollis
Abronia mixtecaGerrhonotus infernalisGerrhonotus liocephalusColoptychon rhombifer
Gerrhonotus parvusElgaria multicarinataElgaria panamintinaElgaria paucicarinataElgaria kingii
Elgaria coeruleaDopasia harti
Dopasia gracilisOphisaurus attenuatus
Anguis fragilisPseudopus apodus
Ophisaurus koellikeriOphisaurus ventralis
Celestus haetianusCelestus agasepsoides
Ophiodes striatusDiploglossus pleii
Diploglossus bilobatusCelestus enneagrammus
Anniella geronimensisAnniella pulchra
Heloderma horridumHeloderma suspectum
Xenosaurus grandisXenosaurus platyceps
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M
Xenosauridae
Helodermatidae
Anniellidae
Anguidae
Shinisauridae Lanthanotidae
Varanidae
Gerrhonotinae
Anguinae
Diploglossinae
Figure 14
Trioceros sternfeldiTrioceros rudis
Trioceros elliotiTrioceros hoehneliiTrioceros schubotzi
Trioceros narraiocaTrioceros jacksonii
Trioceros bitaeniatusTrioceros fuelleborni
Trioceros goetzeiTrioceros tempeli
Trioceros werneriTrioceros deremensis
Trioceros melleriTrioceros johnstoni
Trioceros oweniTrioceros pfefferiTrioceros quadricornisTrioceros wiedersheimi
Trioceros cristatusTrioceros montiumTrioceros feae
Trioceros balebicornutusTrioceros harennae
Trioceros affinisChamaeleo arabicus
Chamaeleo calyptratusChamaeleo zeylanicus
Chamaeleo chamaeleonChamaeleo calcaricarensChamaeleo africanus
Chamaeleo monachusChamaeleo senegalensis
Chamaeleo roperiChamaeleo dilepis
Chamaeleo quilensisChamaeleo gracilis
Chamaeleo necasiChamaeleo laevigatus
Chamaeleo namaquensisCalumma globifer
Calumma parsoniiCalumma oshaughnessyi
Calumma capuroniRhampholeon chapmanorum
Rhampholeon platycepsRhampholeon nchisiensis
Rhampholeon beraducciiRhampholeon boulengeriRhampholeon acuminatus
Rhampholeon uluguruensisRhampholeon moyeri
Rhampholeon marshalliRhampholeon viridis
Rhampholeon spinosusRhampholeon temporalis
Rhampholeon spectrumNadzikambia mlanjensis
Calumma nasutumCalumma boettgeri
Calumma fallaxCalumma gallus
Calumma maltheCalumma guibei
Calumma hilleniusiCalumma tsaratananense
Calumma brevicorneCalumma crypticum
Calumma cucullatumRieppeleon brevicaudatus
Rieppeleon brachyurusRieppeleon kerstenii
Calumma gastrotaeniaCalumma furcifer
Bradypodion ventraleBradypodion karrooicumBradypodion taeniabronchum
Bradypodion gutturaleBradypodion atromontanum
Bradypodion occidentaleBradypodion setaroi
Bradypodion nemoraleBradypodion dracomontanumBradypodion transvaalense
Bradypodion thamnobatesBradypodion melanocephalum
Bradypodion cafferBradypodion damaranumBradypodion pumilum
Kinyongia fischeriKinyongia vosseleri
Kinyongia multituberculataKinyongia matschiei
Kinyongia boehmeiKinyongia tavetana
Kinyongia uthmoelleriKinyongia carpenteri
Kinyongia xenorhinaKinyongia excubitor
Kinyongia adolfifridericiKinyongia tenue
Kinyongia oxyrhinaFurcifer antimenaFurcifer labordi
Furcifer lateralisFurcifer belalandaensis
Furcifer verrucosusFurcifer oustaleti
Furcifer angeliFurcifer pardalis
Furcifer polleniFurcifer cephalolepis
Furcifer campaniFurcifer petteri
Furcifer willsiiFurcifer minor
Furcifer bifidusFurcifer balteatus
Brookesia lineataBrookesia betschiBrookesia thieliBrookesia vadoni
Brookesia ebenauiBrookesia antakaranaBrookesia ambreensis
Brookesia stumpffiBrookesia griveaudi
Brookesia valerieaeBrookesia superciliaris
Brookesia therezieniBrookesia bonsiBrookesia brygooi
Brookesia decaryiBrookesia perarmata
Brookesia peyrierasiBrookesia karchei
Brookesia tuberculataBrookesia minima
Brookesia exarmataBrookesia dentata
Brookesia nasusBrookesia lolontany
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ChamaeleonidaeBrookesiinae
Chamaeleoninae
NFigure 15
Pseudocalotes flavigulaPseudocalotes brevipes
Pseudocalotes kakhienensisJapalura splendida
Japalura flavicepsOtocryptis wiegmanniSitana ponticeriana
Acanthosaura lepidogasterAcanthosaura capraAcanthosaura armataAcanthosaura crucigera
Salea horsfieldiiCalotes htunwiniCalotes calotes
Calotes irawadiCalotes versicolor
Calotes liolepisCalotes nigrilabris
Calotes ceylonensisCalotes liocephalusCalotes emma
Calotes chincolliumCalotes mystaceus
Ceratophora erdeleniCeratophora stoddartii
Ceratophora karuCeratophora aspera
Cophotis ceylanicaCophotis dumbara
Lyriocephalus scutatusGonocephalus kuhliiGonocephalus chamaeleontinus
Gonocephalus grandisBronchocela cristatella
Coryphophylax subcristatusAphaniotis fusca
Gonocephalus robinsoniiJapalura polygonata
Phoxophrys nigrilabrisDraco ornatusDraco palawanensisDraco spilopterus
Draco cornutusDraco guentheri
Draco quadrasiDraco cyanopterusDraco reticulatusDraco boschmai
Draco timorensisDraco volans
Draco bimaculatusDraco rhytismaDraco bourouniensis
Draco beccariiDraco biaroDraco caerulhians
Draco spilonotusDraco obscurusDraco taeniopterus
Draco blanfordiiDraco indochinensisDraco haematopogon
Draco melanopogonDraco quinquefasciatus
Draco mindanensisDraco maximus
Draco cristatellusDraco fimbriatus
Draco maculatusDraco lineatus
Draco dussumieriPtyctolaemus collicristatus
Ptyctolaemus gularisJapalura tricarinata
Japalura variegataMantheyus phuwuanensis
Agama paragamaAgama planiceps
Agama finchiAgama agama
Agama doriaeAgama sankaranicaAgama mwanzae
Agama kaimosaeAgama lionotus
Agama rueppelliAgama caudospinosa
Agama aculeataAgama armata
Agama hispidaAgama atraAgama anchietae
Agama weidholziAgama gracilimembris
Agama insularisAgama boulengeri
Agama castroviejoiAgama impalearis
Agama bouetiAgama spinosa
Trapelus savigniiTrapelus flavimaculatus
Trapelus pallidusTrapelus ruderatus
Trapelus agilisTrapelus mutabilisTrapelus sanguinolentus
Bufoniceps laungwalaensisXenagama taylori
Acanthocercus atricollisPseudotrapelus sinaitus
Phrynocephalus guttatusPhrynocephalus albolineatus
Phrynocephalus melanurusPhrynocephalus przewalskiiPhrynocephalus versicolorPhrynocephalus raddei
Phrynocephalus mystaceusPhrynocephalus helioscopus
Phrynocephalus axillarisPhrynocephalus lidskiiPhrynocephalus vlangalii
Phrynocephalus putjataiPhrynocephalus theobaldiPhrynocephalus forsythiiPhrynocephalus interscapularis
Phrynocephalus scutellatusLaudakia erythrogasterLaudakia caucasia
Laudakia microlepisLaudakia stoliczkana
Laudakia himalayanaLaudakia lehmanni
Laudakia nuptaLaudakia tuberculataLaudakia sacra
Laudakia stellio
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Diporiphora lalliae
Diporiphora arnhemica
Diporiphora albilabris
Diporiphora bennettii
Diporiphora nobbi
Diporiphora australis
Diporiphora amphiboluroides
Diporiphora valens
Diporiphora pindan
Diporiphora bilineata
Diporiphora magna
Diporiphora reginae
Diporiphora winneckei
Diporiphora linga
Diporiphora superba
Pogona vitticeps
Pogona minor
Pogona minima
Pogona henrylawsoni
Pogona nullarbor
Pogona barbata
Rankinia diemensis
Tympanocryptis lineata
Tympanocryptis pinguicolla
Tympanocryptis cephalus
Tympanocryptis tetraporophora
Tympanocryptis intima
Tympanocryptis uniformis
Amphibolurus norrisi
Amphibolurus muricatus
Lophognathus gilberti
Chlamydosaurus kingii
Lophognathus longirostris
Lophognathus temporalis
Ctenophorus femoralis
Ctenophorus fordi
Ctenophorus maculatus
Ctenophorus pictus
Ctenophorus mckenziei
Ctenophorus scutulatus
Ctenophorus isolepis
Ctenophorus ornatus
Ctenophorus caudicinctus
Ctenophorus fionni
Ctenophorus decresii
Ctenophorus vadnappa
Ctenophorus tjantjalka
Ctenophorus rufescens
Ctenophorus reticulatus
Ctenophorus nuchalis
Ctenophorus cristatus
Ctenophorus salinarum
Ctenophorus gibba
Ctenophorus clayi
Ctenophorus adelaidensis
Ctenophorus maculosus
Intellagama lesueurii
Hypsilurus bruijnii
Hypsilurus nigrigularis
Hypsilurus papuensis
Hypsilurus modestus
Chelosania brunnea
Moloch horridus
Hypsilurus dilophus
Hypsilurus boydii
Hypsilurus spinipes
Physignathus cocincinus
Hydrosaurus amboinensis
Leiolepis guentherpetersi
Leiolepis guttata
Leiolepis reevesii
Leiolepis belliana
Uromastyx yemenensis
Uromastyx benti
Uromastyx ocellata
Uromastyx ornata
Uromastyx leptieni
Uromastyx aegyptia
Uromastyx thomasi
Uromastyx dispar
Uromastyx acanthinura
Uromastyx geyri
Uromastyx macfadyeni
Uromastyx princeps
Uromastyx asmussi
Uromastyx loricata
Uromastyx hardwickii
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Ag
am
idae
Uromasticinae
Leiolepidinae
Am
phib
olu
rinae
Hydrosaurinae Agam
inae
Draconinae
O
(i)
(ii)
(i)
(ii)
Figure 16
Ctenosaura oaxacanaCtenosaura quinquecarinataCtenosaura flavidorsalisCtenosaura palearis
Ctenosaura bakeriCtenosaura oedirhinaCtenosaura melanosterna
Ctenosaura pectinataCtenosaura acanthuraCtenosaura hemilophaCtenosaura similis
Conolophus pallidusConolophus subcristatusAmblyrhynchus cristatus
Cyclura cychluraCyclura nubila
Cyclura rileyiCyclura colleiCyclura carinataCyclura ricordiCyclura cornuta
Cyclura pinguisSauromalus variusSauromalus hispidusSauromalus klauberiSauromalus aterIguana delicatissima
Iguana iguanaBrachylophus vitiensisBrachylophus fasciatus
Dipsosaurus dorsalisTropidurus torquatusTropidurus hispidus
Tropidurus oreadicusTropidurus insulanus
Tropidurus erythrocephalusTropidurus mucujensis
Tropidurus etheridgeiTropidurus cocorobensis
Tropidurus itambereTropidurus psammonastes
Tropidurus montanusTropidurus hygomi
Eurolophosaurus nanuzaeEurolophosaurus amathites
Eurolophosaurus divaricatusTropidurus spinulosus
Tropidurus callathelysPlica plica
Plica lumariaPlica umbra
Strobilurus torquatusTropidurus bogerti
Uracentron flavicepsUranoscodon superciliosus
Microlophus duncanensisMicrolophus albemarlensisMicrolophus pacificusMicrolophus grayii
Microlophus delanonisMicrolophus stolzmanni
Microlophus habeliiMicrolophus bivittatus
Microlophus occipitalisMicrolophus koepckeorum
Microlophus yaneziMicrolophus theresioidesMicrolophus atacamensis
Microlophus quadrivittatusMicrolophus peruvianus
Microlophus tigrisMicrolophus heterolepis
Microlophus theresiaeMicrolophus thoracicus
Stenocercus stigmosusStenocercus melanopygus
Stenocercus latebrosusStenocercus orientalis
Stenocercus ornatissimusStenocercus chrysopygus
Stenocercus cupreusStenocercus boettgeriStenocercus empetrusStenocercus eunetopsisStenocercus imitator
Stenocercus variusStenocercus torquatus
Stenocercus crassicaudatusStenocercus marmoratus
Stenocercus humeralisStenocercus angel
Stenocercus guentheriStenocercus festae
Stenocercus chotaStenocercus rhodomelasStenocercus anguliferStenocercus iridescens
Stenocercus puyangoStenocercus percultusStenocercus ornatus
Stenocercus caducusStenocercus limitaris
Stenocercus doellojuradoiStenocercus azureusStenocercus roseiventris
Stenocercus ochoaiStenocercus formosus
Stenocercus apurimacusStenocercus scapularis
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Iguanidae
Tropiduridae
PQ-RFigure 17
Sceloporus adleriSceloporus stejnegeriSceloporus formosusSceloporus subpictusSceloporus cryptus
Sceloporus lundelliSceloporus malachiticusSceloporus smaragdinus
Sceloporus taeniocnemisSceloporus spinosusSceloporus edwardtaylori
Sceloporus horridusSceloporus cautusSceloporus undulatus
Sceloporus consobrinusSceloporus woodiSceloporus virgatus
Sceloporus occidentalisSceloporus olivaceus
Sceloporus clarkiiSceloporus cyanogenysSceloporus serriferSceloporus ornatus
Sceloporus minorSceloporus dugesii
Sceloporus poinsettiiSceloporus macdougalli
Sceloporus mucronatusSceloporus bulleri
Sceloporus torquatusSceloporus jarrovii
Sceloporus insignisSceloporus megalepidurus
Sceloporus goldmaniSceloporus samcolemani
Sceloporus chaneyiSceloporus slevini
Sceloporus scalarisSceloporus bicanthalis
Sceloporus aeneusSceloporus heterolepis
Sceloporus palaciosiSceloporus grammicus
Sceloporus melanorhinusSceloporus hunsakeriSceloporus orcuttiSceloporus lickiSceloporus lineatulus
Sceloporus zosteromusSceloporus magister
Sceloporus arenicolusSceloporus graciosusSceloporus vandenburgianus
Sceloporus ochoterenaeSceloporus jalapae
Sceloporus maculosusSceloporus gadoviae
Sceloporus nelsoniSceloporus pyrocephalus
Sceloporus merriamiSceloporus siniferus
Sceloporus squamosusSceloporus carinatus
Sceloporus utiformisSceloporus angustus
Sceloporus grandaevusSceloporus cozumelae
Sceloporus smithiSceloporus variabilis
Sceloporus teapensisSceloporus chrysostictus
Sceloporus couchiiSceloporus parvus
Urosaurus auriculatusUrosaurus clarionensisUrosaurus ornatus
Urosaurus graciosusUrosaurus nigricaudusUrosaurus lahtelaiUrosaurus bicarinatus
Urosaurus gadoviUta stejnegeri
Uta stansburianaUta palmeriUta squamataPetrosaurus thalassinusPetrosaurus repens
Petrosaurus mearnsiPhrynosoma hernandesi
Phrynosoma douglassiiPhrynosoma ditmarsiPhrynosoma orbiculare
Phrynosoma modestumPhrynosoma taurusPhrynosoma braconnieri
Phrynosoma asioPhrynosoma wigginsiPhrynosoma cerroense
Phrynosoma blainvilliiPhrynosoma coronatum
Phrynosoma platyrhinosPhrynosoma mcalliiPhrynosoma solare
Phrynosoma cornutumHolbrookia maculataHolbrookia propinquaHolbrookia lacerata
Cophosaurus texanusCallisaurus draconoides
Uma notataUma inornataUma scoparia
Uma paraphygasUma exsul
Crotaphytus collarisCrotaphytus nebriusCrotaphytus bicinctores
Crotaphytus grismeriCrotaphytus insularis
Crotaphytus vestigiumCrotaphytus reticulatus
Crotaphytus antiquusGambelia copeiiGambelia sila
Gambelia wislizeniiLeiocephalus barahonensisLeiocephalus schreibersii
Leiocephalus personatusLeiocephalus psammodromus
Leiocephalus ravicepsLeiocephalus carinatus
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Leio
cep
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ae
Crotaphytidae
Ph
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dae
Liolaemus lavillaiLiolaemus ornatusLiolaemus calchaquiLiolaemus albicepsLiolaemus irregularis
Liolaemus crepuscularisLiolaemus abaucanLiolaemus espinozai
Liolaemus quilmesLiolaemus koslowskyi
Liolaemus laurentiLiolaemus darwiniiLiolaemus grosseorum
Liolaemus olongastaLiolaemus chacoensis
Liolaemus uspallatensisLiolaemus hermannuneziLiolaemus xanthoviridisLiolaemus fitzingerii
Liolaemus chehuachekenkLiolaemus morenoiLiolaemus canqueli
Liolaemus melanopsLiolaemus donosobarrosiLiolaemus cuyanus
Liolaemus telsenLiolaemus inacayali
Liolaemus boulengeriLiolaemus rothi
Liolaemus wiegmanniiLiolaemus azarai
Liolaemus scapularisLiolaemus multimaculatusLiolaemus riojanusLiolaemus salinicola
Liolaemus lutzaeLiolaemus occipitalis
Liolaemus pseudoanomalusLiolaemus andinus
Liolaemus multicolorLiolaemus molinaiLiolaemus dorbignyiLiolaemus huacahuasicusLiolaemus fabianiLiolaemus audituvelatus
Liolaemus ruibaliLiolaemus vallecurensisLiolaemus famatinae
Liolaemus orientalisLiolaemus stolzmanni
Liolaemus archeforusLiolaemus tristisLiolaemus zullyaeLiolaemus scolaroi
Liolaemus sarmientoiLiolaemus gallardoi
Liolaemus kingiiLiolaemus escarchadosiLiolaemus tari
Liolaemus bagualiLiolaemus somuncurae
Liolaemus uptoniLiolaemus magellanicus
Liolaemus kolenghLiolaemus silvanaeLiolaemus lineomaculatus
Liolaemus hatcheriLiolaemus saxatilis
Liolaemus gracilisLiolaemus robertmertensiLiolaemus ramirezae
Liolaemus yanalcuLiolaemus walkeriLiolaemus puna
Liolaemus chaltinLiolaemus bitaeniatus
Liolaemus pagaburoiLiolaemus bibronii
Liolaemus hernaniLiolaemus pictusLiolaemus cyanogaster
Liolaemus chiliensisLiolaemus schroederi
Liolaemus gravenhorstiiLiolaemus belliiLiolaemus coeruleusLiolaemus elongatus
Liolaemus kriegiLiolaemus leopardinus
Liolaemus ceiiLiolaemus buergeriLiolaemus petrophilusLiolaemus umbriferLiolaemus capillitasLiolaemus heliodermis
Liolaemus dicktracyiLiolaemus thermarum
Liolaemus austromendocinusLiolaemus paulinaeLiolaemus plateiLiolaemus nigromaculatus
Liolaemus pseudolemniscatusLiolaemus zapallarensisLiolaemus atacamensis
Liolaemus fuscusLiolaemus nigroviridisLiolaemus nitidus
Liolaemus monticolaLiolaemus lemniscatus
Liolaemus tenuisPhymaturus antofagastensisPhymaturus mallimacciiPhymaturus punaePhymaturus palluma
Phymaturus dorsimaculatusPhymaturus patagonicusPhymaturus somuncurensis
Phymaturus indistinctusCtenoblepharys adspersa
Leiosaurus catamarcensisLeiosaurus paronae
Leiosaurus belliiPristidactylus torquatus
Diplolaemus darwiniiPristidactylus scapulatus
Urostrophus vautieriAnisolepis longicauda
Urostrophus gallardoiEnyalius leechii
Enyalius bilineatusOplurus fierinensis
Oplurus grandidieriOplurus saxicola
Oplurus quadrimaculatusOplurus cuvieri
Oplurus cyclurusChalarodon madagascariensis
Enyalioides praestabilisEnyalioides microlepisEnyalioides palpebralis
Enyalioides oshaughnessyiEnyalioides heterolepis
Enyalioides laticepsMorunasaurus annularis
Hoplocercus spinosusPolychrus marmoratus
Polychrus femoralisPolychrus acutirostris
Polychrus gutturosus
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Po
lych
roti
dae
Ho
plo
cerc
idae
Opluridae
En
yaliid
ae
Lio
laem
idae
Leiosaurinae
Enyaliinae
(i)
(ii)
(i)(ii)
QRFigure 18
Anolis smaragdinusAnolis allisoniAnolis maynardi
Anolis brunneusAnolis longiceps
Anolis porcatusAnolis carolinensisAnolis isolepisAnolis oporinus
Anolis argillaceusAnolis centralis
Anolis pumilusAnolis loysiana
Anolis guazumaAnolis garridoi
Anolis angusticepsAnolis paternusAnolis alayoniAnolis sheplaniAnolis placidus
Anolis alutaceusAnolis inexpectatus
Anolis vanidicusAnolis alfaroi
Anolis macilentusAnolis cupeyalensisAnolis cyanopleurusAnolis rejectus
Anolis clivicolaAnolis lucius
Anolis whitemaniAnolis cybotes
Anolis armouriAnolis shreveiAnolis haetianusAnolis longitibialis
Anolis strahmiAnolis marcanoi
Anolis barahonaeAnolis baleatus
Anolis ricordiAnolis eugenegrahami
Anolis christopheiAnolis cuvieri
Anolis guamuhayaAnolis chamaeleonides
Anolis porcusAnolis barbatus
Anolis argenteolusAnolis alumina
Anolis semilineatusAnolis olssoni
Anolis barbouriAnolis insolitus
Anolis fowleriAnolis etheridgei
Anolis equestrisAnolis luteogularis
Anolis baracoaeAnolis nobleiAnolis smallwoodi
Anolis alinigerAnolis singularisAnolis chlorocyanusAnolis coelestinus
Anolis hendersoniAnolis dolichocephalus
Anolis bahorucoensisAnolis darlingtoni
Anolis monticolaAnolis occultus
Anolis vermiculatusAnolis bartschi
Anolis maculigulaAnolis casildae
Anolis frenatusAnolis princeps
Anolis chocorumAnolis fraseri
Anolis danieliAnolis insignisAnolis microtus
Anolis agassiziAnolis neblininusAnolis calimae
Anolis anatolorosAnolis jacare
Anolis tigrinusAnolis transversalisAnolis punctatus
Anolis inderenaeAnolis vanzoliniiAnolis heterodermusAnolis nicefori
Anolis euskalerriariAnolis gemmosusAnolis aequatorialisAnolis ventrimaculatusAnolis chlorisAnolis festae
Anolis peraccaeAnolis huilaeAnolis fitchi
Anolis extremusAnolis roquet
Anolis aeneusAnolis richardii
Anolis trinitatisAnolis griseus
Anolis bonairensisAnolis luciae
Anolis boettgeriBasiliscus basiliscus
Basiliscus plumifronsBasiliscus vittatus
Basiliscus galeritusCorytophanes cristatus
Corytophanes percarinatusLaemanctus longipes
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Anolis trachyderma
Anolis poecilopus
Anolis tropidogaster
Anolis lionotus
Anolis oxylophus
Anolis zeus
Anolis limifrons
Anolis lemurinus
Anolis bicaorum
Anolis carpenteri
Anolis ocelloscapularis
Anolis tropidonotus
Anolis capito
Anolis polylepis
Anolis cupreus
Anolis fuscoauratus
Anolis kemptoni
Anolis altae
Anolis humilis
Anolis pachypus
Anolis sericeus
Anolis isthmicus
Anolis intermedius
Anolis laeviventris
Anolis ortonii
Anolis quercorum
Anolis aquaticus
Anolis woodi
Anolis biporcatus
Anolis bitectus
Anolis uniformis
Anolis polyrhachis
Anolis sminthus
Anolis crassulus
Anolis utilensis
Anolis purpurgularis
Anolis loveridgei
Anolis chrysolepis
Anolis bombiceps
Anolis nitens
Anolis meridionalis
Anolis lineatus
Anolis onca
Anolis annectens
Anolis auratus
Anolis grahami
Anolis conspersus
Anolis garmani
Anolis opalinus
Anolis valencienni
Anolis reconditus
Anolis lineatopus
Anolis bremeri
Anolis quadriocellifer
Anolis sagrei
Anolis ophiolepis
Anolis mestrei
Anolis homolechis
Anolis jubar
Anolis confusus
Anolis guafe
Anolis ahli
Anolis allogus
Anolis rubribarbaris
Anolis imias
Anolis cristatellus
Anolis desechensis
Anolis ernestwilliamsi
Anolis scriptus
Anolis cooki
Anolis monensis
Anolis krugi
Anolis pulchellus
Anolis gundlachi
Anolis poncensis
Anolis evermanni
Anolis stratulus
Anolis acutus
Anolis caudalis
Anolis marron
Anolis brevirostris
Anolis websteri
Anolis distichus
Anolis marmoratus
Anolis sabanus
Anolis nubilus
Anolis lividus
Anolis ferreus
Anolis terraealtae
Anolis oculatus
Anolis gingivinus
Anolis bimaculatus
Anolis leachii
Anolis pogus
Anolis wattsi
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Corytophanidae
Dactyloidae
(i)
(ii)
(i)
(ii)
RFigure 19
Austrotyphlops pilbarensis
Austrotyphlops hamatus
Austrotyphlops endoterus
Austrotyphlops australis
Austrotyphlops waitii
Austrotyphlops splendidus
Austrotyphlops pinguis
Ramphotyphlops bicolor
Austrotyphlops ligatus
Austrotyphlops ganei
Austrotyphlops troglodytes
Austrotyphlops kimberleyensis
Austrotyphlops ammodytes
Austrotyphlops unguirostris
Austrotyphlops leptosomus
Austrotyphlops longissimus
Austrotyphlops grypus
Austrotyphlops bituberculatus
Austrotyphlops guentheri
Austrotyphlops howi
Austrotyphlops diversus
Ramphotyphlops polygrammicus
Acutotyphlops kunuaensis
Acutotyphlops subocularis
Ramphotyphlops acuticaudus
Ramphotyphlops lineatus
Typhlopidae sp. (Sri Lanka)
Ramphotyphlops braminus
Typhlops pammeces
Ramphotyphlops albiceps
Typhlops ruber
Typhlops luzonensis
Typhlops arenarius
Typhlops vermicularis
Afrotyphlops punctatus
Afrotyphlops congestus
Afrotyphlops lineolatus
Afrotyphlops bibronii
Afrotyphlops fornasinii
Megatyphlops schlegelii
Afrotyphlops angolensis
Typhlops elegans
Letheobia obtusa
Letheobia newtoni
Letheobia feae
Rhinotyphlops lalandei
Typhlops catapontus
Typhlops richardi
Typhlops platycephalus
Typhlops hypomethes
Typhlops granti
Typhlops dominicanus
Typhlops monastus
Typhlops notorachius
Typhlops anousius
Typhlops anchaurus
Typhlops contorhinus
Typhlops arator
Typhlops caymanensis
Typhlops agoralionis
Typhlops sylleptor
Typhlops jamaicensis
Typhlops capitulatus
Typhlops rostellatus
Typhlops sulcatus
Typhlops titanops
Typhlops schwartzi
Typhlops lumbricalis
Typhlops eperopeus
Typhlops syntherus
Typhlops hectus
Typhlops pusillus
Typhlops biminiensis
Typhlops reticulatus
Typhlops brongersmianus
Xenotyphlops grandidieri
Gerrhopilus mirus
Gerrhopilus hedraeus
Leptotyphlops scutifrons
Leptotyphlops distanti
Leptotyphlops conjunctus
Leptotyphlops sylvicolus
Leptotyphlops nigricans
Leptotyphlops nigroterminus
Namibiana occidentalis
Myriopholis adleri
Myriopholis macrorhyncha
Myriopholis blanfordi
Myriopholis boueti
Myriopholis rouxestevae
Myriopholis algeriensis
Myriopholis longicauda
Mitophis leptipileptus
Mitophis pyrites
Mitophis asbolepis
Tetracheilostoma breuili
Tetracheilostoma carlae
Rena humilis
Rena dulcis
Trilepida macrolepis
Epictia columbi
Epictia goudotii
Epictia albifrons
Siagonodon septemstriatus
Tricheilostoma bicolor
Typhlophis squamosus
Liotyphlops albirostris
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83
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87
76
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72
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988174
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93
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100 Anomalepididae
Leptotyphlopidae
Gerrhopilidae
Xenotyphlopidae
Typ
hlo
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ae
ST-AAFigure 20
Antaresia childreniAntaresia stimsoniAntaresia perthensisAntaresia maculosaMorelia bredliMorelia spilota
Morelia carinataMorelia viridisBothrochilus boaLeiopython albertisii
Liasis fuscusLiasis mackloti
Apodora papuanaLiasis olivaceus
Aspidites melanocephalusAspidites ramsayi
Morelia amethistinaMorelia boeleniMorelia oenpelliensisBroghammerus timoriensisBroghammerus reticulatus
Python molurusPython sebaePython curtusPython regius
Python brongersmaiLoxocemus bicolor
Xenopeltis unicolorRhinophis blythiiRhinophis homolepisPseudotyphlops philippinusRhinophis oxyrhynchus
Rhinophis philippinusRhinophis dorsimaculatus
Rhinophis drummondhayiUropeltis phillipsiUropeltis melanogaster
Rhinophis travancoricusUropeltis ceylanicus
Uropeltis liuraBrachyophidium rhodogaster
Melanophidium punctatumCylindrophis ruffus
Cylindrophis maculatusAnomochilus leonardi
Epicrates striatusEpicrates exsulEpicrates chrysogasterEpicrates subflavus
Epicrates fordiEpicrates monensisEpicrates inornatus
Epicrates anguliferEunectes notaeusEunectes murinus
Epicrates cenchriaCorallus annulatus
Corallus caninusCorallus hortulanusBoa constrictorEryx tataricusEryx miliaris
Eryx elegansEryx jaculusEryx johnii
Eryx conicusEryx colubrinus
Eryx jayakariCandoia bibroni
Candoia carinataCandoia aspera
Lichanura trivirgataCharina bottae
Ungaliophis continentalisExiliboa placata
Calabaria reinhardtiiAcrantophis dumeriliAcrantophis madagascariensisSanzinia madagascariensisCasarea dussumieriXenophidion schaeferiTropidophis melanurusTropidophis feicki
Tropidophis haetianusTropidophis greenwayi
Tropidophis wrightiTropidophis pardalis
Trachyboa boulengeriTrachyboa gularis
Anilius scytale
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5398
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100 Aniliidae
Tropidophiidae
Xenophidiidae
Bolyeriidae
Boidae
Anomochilidae
Cylindrophiidae
Uropeltidae
XenopeltidaeLoxocemidae
Pythonidae
Calabariidae
Sanziniinae
Ungaliophiinae
Candoiinae
Erycinae
Boinae
TU-AAFigure 21
Bitis rubida
Bitis cornuta
Bitis atropos
Bitis xeropaga
Bitis caudalis
Bitis peringueyi
Bitis gabonica
Bitis nasicornis
Bitis arietans
Bitis worthingtoni
Atheris desaixi
Atheris nitschei
Atheris squamigera
Atheris hispida
Atheris chlorechis
Atheris barbouri
Atheris ceratophora
Echis pyramidum
Echis leucogaster
Echis coloratus
Echis omanensis
Echis jogeri
Echis ocellatus
Echis carinatus
Causus defilippii
Causus rhombeatus
Causus resimus
Cerastes gasperettii
Cerastes cerastes
Cerastes vipera
Proatheris superciliaris
Vipera lotievi
Vipera renardi
Vipera dinniki
Vipera ursinii
Vipera eriwanensis
Vipera kaznakovi
Vipera barani
Vipera berus
Vipera nikolskii
Vipera seoanei
Vipera aspis
Vipera latastei
Vipera ammodytes
Macrovipera mauritanica
Macrovipera deserti
Daboia russelii
Daboia palaestinae
Montivipera wagneri
Montivipera albizona
Montivipera bornmuelleri
Montivipera raddei
Montivipera xanthina
Macrovipera schweizeri
Macrovipera lebetina
Pseudocerastes fieldi
Pseudocerastes persicus
Eristicophis macmahoni
Pareas nuchalis
Pareas carinatus
Pareas hamptoni
Pareas macularius
Pareas margaritophorus
Pareas formosensis
Pareas boulengeri
Pareas monticola
Aplopeltura boa
Asthenodipsas vertebralis
Achalinus rufescens
Achalinus meiguensis
Xenodermus javanicus
Stoliczkia borneensis
Acrochordus arafurae
Acrochordus granulatus
Acrochordus javanicus
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Crotalus adamanteusCrotalus tigris
Crotalus mitchelliiCrotalus viridis
Crotalus oreganusCrotalus scutulatusCrotalus tortugensis
Crotalus atroxCrotalus catalinensis
Crotalus ruberCrotalus totonacusCrotalus molossus
Crotalus basiliscusCrotalus simusCrotalus durissus
Crotalus horridusCrotalus willardi
Crotalus ravusCrotalus cerastesCrotalus polystictus
Crotalus enyoCrotalus aquilus
Crotalus lepidusCrotalus pusillus
Crotalus triseriatusCrotalus tancitarensis
Crotalus transversusCrotalus intermedius
Crotalus priceiSistrurus miliarius
Sistrurus catenatusAgkistrodon bilineatus
Agkistrodon tayloriAgkistrodon piscivorus
Agkistrodon contortrixBothriechis rowleyi
Bothriechis auriferBothriechis bicolor
Bothriechis thalassinusBothriechis marchi
Bothriechis lateralisBothriechis nigroviridis
Bothriechis schlegeliiLachesis mutaLachesis stenophrys
Mixcoatlus melanurusMixcoatlus barbouri
Ophryacus undulatusBothrops leucurus
Bothrops moojeniBothrops atrox
Bothrops colombiensisBothrops marajoensis
Bothrops asperBothrops lanceolatus
Bothrops caribbaeusBothrops punctatusBothrops jararacussu
Bothrops braziliBothriopsis bilineata
Bothriopsis taeniataBothriopsis chloromelasBothriopsis pulchra
Bothropoides alcatrazBothropoides insularis
Bothropoides jararacaBothropoides neuwiedi
Bothropoides diporusBothropoides erythromelas
Rhinocerophis itapetiningaeRhinocerophis alternatus
Rhinocerophis fonsecaiRhinocerophis cotiara
Rhinocerophis ammodytoidesBothrops pictus
Bothrocophias hyoproraBothrocophias microphthalmus
Bothrocophias campbelliPorthidium lansbergii
Porthidium porrasiPorthidium nasutum
Porthidium yucatanicumPorthidium dunni
Porthidium ophryomegasCerrophidion petlalcalensis
Cerrophidion tzotzilorumCerrophidion godmani
Atropoides picadoiAtropoides olmec
Atropoides nummiferAtropoides occiduus
Gloydius saxatilisGloydius intermedius
Gloydius shedaoensisGloydius halys
Gloydius strauchiGloydius blomhoffiiGloydius ussuriensis
Gloydius brevicaudusGloydius tsushimaensis
Trimeresurus gracilisOvophis okinavensis
Protobothrops tokarensisProtobothrops flavoviridis
Protobothrops mucrosquamatusProtobothrops elegans
Protobothrops jerdoniiProtobothrops xiangchengensis
Protobothrops cornutusProtobothrops mangshanensis
Protobothrops sieversorumProtobothrops kaulbacki
Ovophis tonkinensisOvophis zayuensisOvophis monticola
Trimeresurus purpureomaculatusTrimeresurus erythrurus
Trimeresurus cantoriTrimeresurus albolabris
Trimeresurus andersoniiTrimeresurus septentrionalis
Trimeresurus insularisTrimeresurus fasciatus
Trimeresurus macropsTrimeresurus venustus
Trimeresurus kanburiensisTrimeresurus vogeli
Trimeresurus stejnegeriTrimeresurus gumprechti
Trimeresurus yunnanensisTrimeresurus medoensis
Trimeresurus tibetanusTrimeresurus popeiorum
Trimeresurus sumatranusTrimeresurus schultzei
Trimeresurus flavomaculatusTrimeresurus malcolmiTrimeresurus hageni
Trimeresurus trigonocephalusTrimeresurus gramineus
Trimeresurus malabaricusTrimeresurus borneensis
Trimeresurus puniceusHypnale zara
Hypnale hypnaleHypnale nepa
Calloselasma rhodostomaDeinagkistrodon acutus
Tropidolaemus wagleriGarthius chaseni
Azemiops feae
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Vip
eri
dae
Acrochordidae
Xen
od
erm
ati
dae
Vip
erinae
Azemiopinae
Cro
talin
ae
UV-AA
(i)
(ii)
(i)
(ii)
Pare
ati
dae
Figure 22
Liophidium chabaudiLiophidium mayottensis
Liophidium torquatumLiophidium therezieni
Liophidium vaillantiLiophidium rhodogaster
Liopholidophis sexlineatusLiopholidophis dolicocercus
Liopholidophis dimorphusHeteroliodon occipitalis
Pseudoxyrhopus ambreensisThamnosophis lateralis
Thamnosophis stumpffiThamnosophis epistibes
Thamnosophis martaeThamnosophis infrasignatus
Dromicodryas bernieriDromicodryas quadrilineatus
Lycodryas granulicepsLycodryas pseudogranuliceps
Lycodryas inopinaeLycodryas sanctijohannis
Lycodryas citrinusLycodryas inornatus
Madagascarophis meridionalisMadagascarophis colubrinus
Ithycyphus oursiIthycyphus miniatus
Micropisthodon ochraceusLangaha madagascariensis
Leioheterodon modestusLeioheterodon madagascariensis
Leioheterodon geayiParastenophis betsileanus
Alluaudina bellyiCompsophis infralineatus
Compsophis laphystiusCompsophis albiventris
Compsophis boulengeriDitypophis vivaxDuberria lutrix
Duberria variegataAmplorhinus multimaculatus
Lycodonomorphus whytiiLycodonomorphus laevissimus
Lycodonomorphus rufulusLycodonomorphus inornatus
Lamprophis fiskiiLamprophis aurora
Lamprophis fuscusLamprophis guttatus
Boaedon olivaceusBoaedon fuliginosus
Boaedon lineatusBoaedon virgatus
Bothrophthalmus lineatusBothrophthalmus brunneus
Bothrolycus aterPseudoboodon lemniscatus
Gonionotophis capensisGonionotophis poensis
Gonionotophis brussauxiGonionotophis nyassae
Gonionotophis stenophthalmusInyoka swazicus
Hormonotus modestusLycophidion ornatum
Lycophidion capenseLycophidion laterale
Lycophidion nigromaculatumBuhoma depressiceps
Buhoma procteraePsammodynastes pictus
Psammodynastes pulverulentusPseudaspis cana
Pythonodipsas carinataAparallactus guentheri
Aparallactus capensisAparallactus werneriAparallactus modestus
Polemon acanthiasPolemon collaris
Polemon notatusAmblyodipsas dimidiata
Xenocalamus transvaalensisAmblyodipsas polylepis
Macrelaps microlepidotusAtractaspis corpulenta
Atractaspis micropholisAtractaspis bibronii
Atractaspis boulengeriAtractaspis microlepidota
Atractaspis irregularisHomoroselaps lacteus
Psammophis phillipsiPsammophis mossambicusPsammophis leopardinus
Psammophis sibilansPsammophis rukwae
Psammophis orientalisPsammophis sudanensis
Psammophis subtaeniatusPsammophis lineatusPsammophis biseriatus
Psammophis tanganicusPsammophis praeornatus
Psammophis punctulatusPsammophis schokari
Psammophis angolensisPsammophis notostictusPsammophis leightoniPsammophis jallae
Psammophis trigrammusPsammophis condanarus
Psammophis lineolatusPsammophis crucifer
Psammophylax tritaeniatusPsammophylax variabilis
Psammophylax rhombeatusPsammophylax acutus
Hemirhagerrhis hildebrandtiiHemirhagerrhis kelleri
Hemirhagerrhis viperinaDipsina multimaculata
Mimophis mahfalensisRhagerhis moilensis
Malpolon monspessulanusRhamphiophis rubropunctatus
Rhamphiophis oxyrhynchusProsymna meleagris
Prosymna greigertiProsymna janii
Prosymna visseriProsymna ruspolii
Oxyrhabdium leporinumMicrelaps bicoloratus
Cerberus rynchopsCerberus microlepis
Cerberus australisHomalopsis buccataEnhydris punctata
Fordonia leucobaliaGerarda prevostiana
Cantoria violaceaBitia hydroides
Enhydris bocourtiErpeton tentaculatum
Pseudoferania polylepisMyron richardsonii
Enhydris innominataEnhydris longicauda
Enhydris jagoriiEnhydris enhydris
Enhydris matannensisEnhydris plumbea
Enhydris chinensis
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Homalopsidae
Prosymninae
Psammophiinae
Atractaspidinae
Aparallactinae
Pseudaspidinae
Lamprophiinae
Pseudoxyrhophiinae
Lamprophiidae
VX-AA WFigure 23
Hydrophis ornataHydrophis peronii
Hydrophis kingiiHydrophis macdowelli
Hydrophis schistosaHydrophis majorHydrophis czeblukovi
Hydrophis caerulescensHydrophis brooki
Hydrophis atricepsHydrophis stokesii
Hydrophis platuraHydrophis curtus
Hydrophis parvicepsHydrophis melanocephala
Hydrophis cyanocinctaHydrophis pacificaHydrophis semperi
Hydrophis spiralisHydrophis lapemoides
Hydrophis elegansHydrelaps darwiniensis
Ephalophis greyaeParahydrophis mertoni
Aipysurus laevisAipysurus fuscusAipysurus apraefrontalisAipysurus duboisii
Aipysurus eydouxiiEmydocephalus annulatus
Hemiaspis dameliiHemiaspis signata
Tropidechis carinatusNotechis scutatus
Echiopsis atricepsHoplocephalus bitorquatus
Austrelaps superbusAustrelaps labialis
Drysdalia coronoidesDrysdalia mastersii
Echiopsis curtaPseudonaja textilisPseudonaja modesta
Oxyuranus scutellatusOxyuranus microlepidotus
Denisonia devisiSimoselaps calonotus
Vermicella intermediaSuta spectabilis
Suta sutaSuta monachus
Suta fasciataRhinoplocephalus bicolor
Cryptophis nigrescensElapognathus coronata
Cacophis squamulosusPseudechis guttatus
Pseudechis papuanusPseudechis colletti
Pseudechis australisPseudechis butleri
Pseudechis porphyriacusAcanthophis antarcticusAcanthophis praelongus
Aspidomorphus muelleriAspidomorphus lineaticollis
Aspidomorphus schlegeliSimoselaps bertholdi
Simoselaps anomalusSimoselaps semifasciatus
Furina diademaFurina ornata
Toxicocalamus loriaeDemansia psammophis
Demansia papuensisDemansia vestigiata
Toxicocalamus preussiMicropechis ikaheka
Laticauda colubrinaLaticauda guineai
Laticauda saintgironsiLaticauda laticaudata
Bungarus candidusBungarus multicinctus
Bungarus nigerBungarus caeruleus
Bungarus ceylonicusBungarus sindanus
Bungarus fasciatusBungarus bungaroidesBungarus flavicepsElapsoidea sundevalliiElapsoidea semiannulata
Elapsoidea nigraNaja ashei
Naja nigricollisNaja mossambica
Naja katiensisNaja pallida
Naja nubiaeNaja haje
Naja annuliferaNaja nivea
Naja annulataNaja melanoleuca
Naja multifasciataNaja sumatranaNaja siamensis
Naja mandalayensisNaja naja
Naja atraNaja kaouthia
Hemachatus haemachatusAspidelaps scutatus
Walterinnesia aegyptiaDendroaspis polylepis
Dendroaspis angusticepsOphiophagus hannah
Hemibungarus calligasterMicrurus brasiliensisMicrurus frontalisMicrurus spixii
Micrurus ibibobocaMicrurus altirostris
Micrurus pyrrhocryptusMicrurus baliocoryphus
Micrurus decoratusMicrurus lemniscatusMicrurus hemprichii
Micrurus surinamensisMicrurus dissoleucus
Micrurus mipartitusMicrurus narduccii
Micrurus diastemaMicrurus fulvius
Micrurus psychesMicrurus albicinctus
Micrurus corallinusSinomicrurus macclellandi
Sinomicrurus kelloggiSinomicrurus japonicus
Micruroides euryxanthusMaticora bivirgata
Calliophis melanurus
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Elapidae
WFigure 24
Eirenis punctatolineatusEirenis barani
Eirenis collarisEirenis eiselti
Eirenis coronelloidesEirenis rothiiEirenis lineomaculatus
Eirenis thospitisEirenis medus
Eirenis levantinusEirenis decemlineatus
Eirenis persicusEirenis aurolineatus
Eirenis modestusHierophis spinalisHierophis gemonensis
Hierophis viridiflavusDolichophis caspius
Dolichophis schmidtiDolichophis jugularis
Platyceps florulentusPlatyceps collaris
Platyceps najadumPlatyceps ventromaculatus
Platyceps kareliniPlatyceps rogersi
Platyceps rhodorachisSpalerosophis diadema
Spalerosophis microlepisHemorrhois ravergieri
Hemorrhois nummiferHemorrhois algirus
Hemorrhois hippocrepisHemerophis socotrae
Macroprotodon abubakeriMacroprotodon cucullatus
Bamanophis dorriColuber zebrinus
Lytorhynchus diademaHapsidophrys principis
Hapsidophrys smaragdinaHapsidophrys lineatus
Philothamnus hoplogasterPhilothamnus natalensisPhilothamnus thomensis
Philothamnus girardiPhilothamnus angolensis
Philothamnus semivariegatusPhilothamnus nitidus
Philothamnus heterodermusPhilothamnus carinatus
Thelotornis capensisDispholidus typus
Thrasops jacksoniiCoelognathus flavolineatus
Coelognathus subradiatusCoelognathus erythrurus
Coelognathus helenaCoelognathus radiatus
Oligodon planicepsOligodon torquatus
Oligodon theobaldiOligodon cruentatus
Oligodon splendidusOligodon maculatus
Oligodon cinereusOligodon formosanus
Oligodon ocellatusOligodon chinensisOligodon taeniatusOligodon barroniOligodon cyclurus
Oligodon octolineatusOligodon taeniolatusOligodon sublineatus
Oligodon arnensisScaphiophis albopunctatus
Dasypeltis fasciataDasypeltis gansi
Dasypeltis sahelensisDasypeltis scabra
Dasypeltis atraDasypeltis confusa
Toxicodryas pulverulentaBoiga multomaculata
Boiga beddomeiBoiga ceylonensisBoiga trigonata
Boiga barnesiiBoiga cynodon
Boiga forsteniBoiga dendrophila
Boiga irregularisTelescopus fallax
Dipsadoboa unicolorCrotaphopeltis tornieri
Boiga kraepeliniDendrelaphis tristis
Dendrelaphis schokariDendrelaphis caudolineatus
Dendrelaphis caudolineolatusDendrelaphis bifrenalis
Chrysopelea paradisiChrysopelea ornata
Chrysopelea taprobanicaAhaetulla nasuta
Ahaetulla pulverulentaAhaetulla fronticincta
Grayia smithiiGrayia ornataGrayia tholloni
Sibynophis chinensisSibynophis collaris
Sibynophis triangularisSibynophis subpunctatus
Sibynophis bistrigatusScaphiodontophis annulatus
Pseudoxenodon karlschmidtiPseudoxenodon macrops
Pseudoxenodon bambusicolaPlagiopholis styani
Calamaria yunnanensisCalamaria pavimentata
Pseudorabdion oxycephalum
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Co
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Calamariinae
Pseudoxenodontinae
Sibynophiinae
Grayiinae
Colu
brinae
XY
Z-AA Colubrinae cont.Figure 25
Lampropeltis californiaeLampropeltis splendidaLampropeltis holbrooki
Lampropeltis getulaLampropeltis nigraLampropeltis alterna
Lampropeltis extenuatumLampropeltis triangulumLampropeltis ruthveniLampropeltis mexicanaLampropeltis elapsoidesLampropeltis zonataLampropeltis pyromelana
Lampropeltis webbiLampropeltis calligaster
Cemophora coccineaPseudelaphe flavirufa
Arizona elegansRhinocheilus lecontei
Bogertophis rosaliaeBogertophis subocularis
Pantherophis bairdiPantherophis obsoletusPantherophis alleghaniensis
Pantherophis spiloidesPantherophis slowinskii
Pantherophis emoryiPantherophis guttatus
Pantherophis vulpinusPituophis cateniferPituophis ruthveniPituophis melanoleucus
Pituophis lineaticollisPituophis vertebralis
Pituophis deppeiSenticolis triaspis
Oocatochus rufodorsatusCoronella austriaca
Coronella girondicaElaphe sauromates
Elaphe quatuorlineataElaphe quadrivirgata
Elaphe schrenckiiElaphe climacophora
Elaphe dioneElaphe bimaculata
Elaphe carinataElaphe davidi
Zamenis situlaRhinechis scalaris
Zamenis lineatusZamenis longissimus
Zamenis persicusZamenis hohenackeri
Orthriophis cantorisOrthriophis moellendorffi
Orthriophis hodgsoniOrthriophis taeniurus
Oreocryptophis porphyraceusEuprepiophis mandarinus
Euprepiophis conspicillataArchelaphe bella
Lycodon aulicusLycodon zawi
Lycodon osmanhilliLycodon capucinus
Lycodon carinatusDryocalamus nympha
Lycodon semicarinatumLycodon rufozonatum
Lycodon paucifasciatusLycodon fasciatus
Lycodon laoensisLycodon ruhstrati
Rhadinophis frenatusRhynchophis boulengeri
Rhadinophis prasinusGonyosoma jansenii
Gonyosoma oxycephalumSympholis lippiens
Pseudoficimia frontalisGyalopion canum
Ficimia streckeriConopsis nasusConopsis biserialis
Sonora semiannulataChilomeniscus stramineus
Chionactis occipitalisStenorrhina freminvillei
Rhinobothryum lentiginosumMastigodryas bifossatus
Mastigodryas boddaertiMastigodryas melanolomus
Drymoluber braziliDrymoluber dichrous
Chironius multiventrisChironius laurenti
Chironius exoletusChironius monticola
Chironius bicarinatusChironius flavolineatus
Chironius grandisquamisChironius laevicollisChironius scurrulus
Chironius fuscusDrymarchon corais
Pseustes sulphureusSpilotes pullatus
Phyllorhynchus decurtatusTrimorphodon biscutatus
Drymobius rhombiferDendrophidion percarinatum
Dendrophidion dendrophisLeptophis ahaetulla
Chironius carinatusChironius quadricarinatus
Coluber flagellumColuber constrictor
Coluber taeniatusTantilla melanocephala
Salvadora mexicanaOpheodrys aestivusOpheodrys vernalisOxybelis aeneus
Oxybelis fulgidusPtyas mucosa
Ptyas korrosCyclophiops major
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Colubrinae cont.
YFigure 26
Thamnophis butleri
Thamnophis radix
Thamnophis brachystoma
Thamnophis elegans
Thamnophis gigas
Thamnophis atratus
Thamnophis couchii
Thamnophis ordinoides
Thamnophis hammondii
Thamnophis marcianus
Thamnophis eques
Thamnophis cyrtopsis
Thamnophis chrysocephalus
Thamnophis fulvus
Thamnophis mendax
Thamnophis sumichrasti
Thamnophis scaliger
Thamnophis godmani
Thamnophis exsul
Thamnophis melanogaster
Thamnophis valida
Adelophis foxi
Thamnophis rufipunctatus
Thamnophis proximus
Thamnophis sauritus
Thamnophis sirtalis
Nerodia sipedon
Nerodia fasciata
Nerodia harteri
Nerodia erythrogaster
Nerodia rhombifer
Nerodia taxispilota
Nerodia floridana
Nerodia cyclopion
Tropidoclonion lineatum
Regina grahami
Regina septemvittata
Regina alleni
Regina rigida
Seminatrix pygaea
Storeria dekayi
Storeria occipitomaculata
Virginia striatula
Clonophis kirtlandii
Natrix tessellata
Natrix natrix
Natrix maura
Sinonatrix annularis
Sinonatrix percarinata
Sinonatrix aequifasciata
Opisthotropis cheni
Opisthotropis latouchii
Opisthotropis lateralis
Opisthotropis guangxiensis
Aspidura guentheri
Aspidura ceylonensis
Aspidura trachyprocta
Aspidura drummondhayi
Atretium yunnanensis
Xenochrophis asperrimus
Xenochrophis punctulatus
Atretium schistosum
Xenochrophis piscator
Xenochrophis flavipunctatus
Xenochrophis vittatus
Amphiesma stolatum
Rhabdophis nuchalis
Rhabdophis tigrinus
Rhabdophis subminiatus
Balanophis ceylonensis
Macropisthodon rudis
Lycognathophis seychellensis
Afronatrix anoscopus
Natriciteres olivacea
Amphiesma craspedogaster
Amphiesma sauteri
Trachischium monticola
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Natricinae
ZFigure 27
Sibynomorphus neuwiedi
Sibynomorphus ventrimaculatus
Dipsas albifrons
Dipsas articulata
Sibynomorphus turgidus
Sibynomorphus mikanii
Dipsas pratti
Dipsas variegata
Dipsas neivai
Dipsas catesbyi
Dipsas indica
Tropidodipsas sartorii
Sibon nebulatus
Ninia atrata
Atractus zebrinus
Atractus trihedrurus
Atractus reticulatus
Atractus badius
Atractus flammigerus
Atractus elaps
Atractus albuquerquei
Atractus schach
Atractus wagleri
Atractus zidoki
Geophis godmani
Geophis carinosus
Cryophis hallbergi
Tretanorhinus nigroluteus
Hydromorphus concolor
Adelphicos quadrivirgatum
Leptodeira rubricata
Leptodeira maculata
Leptodeira bakeri
Leptodeira annulata
Leptodeira septentrionalis
Leptodeira punctata
Leptodeira splendida
Leptodeira frenata
Leptodeira uribei
Imantodes cenchoa
Imantodes gemmistratus
Imantodes lentiferus
Imantodes inornatus
Leptodeira nigrofasciata
Tretanorhinus variabilis
Hypsiglena chlorophaea
Hypsiglena torquata
Hypsiglena ochrorhyncha
Hypsiglena affinis
Hypsiglena jani
Hypsiglena slevini
Trimetopon gracile
Pseudoleptodeira latifasciata
Rhadinaea fulvivittis
Rhadinaea flavilata
Coniophanes fissidens
Tantalophis discolor
Amastridium veliferum
Nothopsis rugosus
Farancia erytrogramma
Farancia abacura
Carphophis amoenus
Diadophis punctatus
Heterodon nasicus
Heterodon simus
Heterodon platirhinos
Contia tenuis
Thermophis baileyi
Thermophis zhaoermii
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Pseudoboa coronataPseudoboa nigra
Pseudoboa neuwiediiRhachidelus brazili
Boiruna maculataDrepanoides anomalus
Mussurana bicolorClelia clelia
Phimophis gueriniParaphimophis rusticus
Rodriguesophis iglesiasiOxyrhopus guibei
Oxyrhopus melanogenysOxyrhopus rhombifer
Oxyrhopus clathratusOxyrhopus trigeminus
Oxyrhopus formosusOxyrhopus petolarius
Siphlophis longicaudatusSiphlophis pulcher
Siphlophis compressusSiphlophis cervinus
Hydrodynastes gigasHydrodynastes bicinctus
Caaeteboia amaraliXenopholis scalarisXenopholis undulatus
Tropidodryas striaticepsTropidodryas serra
Thamnodynastes rutilusThamnodynastes strigatus
Thamnodynastes hypoconiaThamnodynastes laneiThamnodynastes pallidusTomodon dorsatus
Ptychophis flavovirgatusTachymenis peruviana
Pseudotomodon trigonatusCalamodontophis paucidens
Gomesophis brasiliensisHelicops angulatus
Helicops gomesiHelicops carinicaudus
Helicops hagmanniHelicops infrataeniatus
Pseudoeryx plicatilisHydrops triangularis
Manolepis putnamiApostolepis cearensisApostolepis sanctaeritae
Apostolepis flavotorquataApostolepis albicollaris
Apostolepis dimidiataApostolepis assimilis
Elapomorphus quinquelineatusPhalotris mertensi
Phalotris lativittatusPhalotris nasutusPhalotris lemniscatusTaeniophallus affinisEchinanthera undulataEchinanthera melanostigma
Taeniophallus nicagusTaeniophallus brevirostris
Sordellina punctataPhilodryas patagoniensis
Philodryas agassiziiPhilodryas aestiva
Philodryas psammophideaPhilodryas mattogrossensis
Philodryas georgeboulengeriPhilodryas argentea
Philodryas olfersiiPhilodryas viridissima
Philodryas nattereriPhilodryas baroni
Erythrolamprus aesculapiiErythrolamprus mimus
Eryrthrolamprus typhlusEryrthrolamprus pygmaea
Eryrthrolamprus brevicepsEryrthrolamprus reginae
Eryrthrolamprus miliarisEryrthrolamprus juliae
Eryrthrolamprus epinephelusEryrthrolamprus poecilogyrusEryrthrolamprus ceii
Eryrthrolamprus almadensisEryrthrolamprus jaegeri
Eryrthrolamprus atraventerXenodon pulcherXenodon matogrossensisXenodon semicinctusXenodon guentheri
Xenodon nattereriXenodon histricus
Xenodon dorbignyiXenodon neuwiedii
Xenodon werneriXenodon merremi
Xenodon severusLygophis meridionalis
Lygophis flavifrenatusLygophis paucidens
Lygophis lineatusLygophis elegantissimus
Lygophis anomalusUromacer frenatusUromacer oxyrhynchusUromacer catesbyi
Schwartzophis funereumSchwartzophis polylepis
Schwartzophis callilaemumAntillophis parvifrons
Hypsirhynchus feroxDarlingtonia haetiana
Ialtris dorsalisAlsophis rufiventrisAlsophis antiguaeAlsophis rijgersmaei
Alsophis antillensisBorikenophis portoricensisHaitiophis anomalusCaraiba andreae
Cubophis vudiiCubophis cantherigerus
Magliophis exiguumArrhyton dolichura
Arrhyton tanyplectumArrhyton procerum
Arrhyton landoiArrhyton vittatum
Arrhyton supernumArrhyton taeniatum
Pseudalsophis elegansPseudalsophis biserialis
Psomophis jobertiPsomophis genimaculatus
Psomophis obtususCrisantophis nevermanni
Conophis lineatus
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Figure 28
Additional files provided with this submission:
Additional file 1: 2046042875907394_add1.txt, 264Khttp://www.biomedcentral.com/imedia/1032288879981940/supp1.txtAdditional file 2: 2046042875907394_add2.docx, 681Khttp://www.biomedcentral.com/imedia/7057154239819418/supp2.docx