A Palaeogene stem crotaphytid (Aciprion formosum) and the phylogenetic affinities of early fossil pleurodontan iguanians.

Pleurodonta is an ancient, diverse clade of iguanian lizard distributed primarily in the Western Hemisphere. Although the clade is a frequent subject of systematic research, phylogenetic resolution among the major pleurodontan clades is elusive. That uncertainty has complicated the interpretations of many fossil pleurodontans. I describe a fossil skull of a pleurodontan lizard from the Palaeogene of Wyoming that was previously allocated to the puzzling taxon Aciprion formosum, and provide an updated morphological matrix for iguanian lizards. Phylogenetic analyses using Bayesian inference demonstrate that the fossil skull is the oldest and first definitive stem member of Crotaphytidae (collared and leopard lizards), establishing the presence of that clade in North America during the Palaeogene. I also discuss new or revised hypotheses for the relationships of several early pleurodontans. In particular, I examine potential evidence for crown-Pleurodonta in the Cretaceous of Mongolia (Polrussia), stem Pleurodonta in the Cretaceous of North America (Magnuviator) and a stem anole in the Eocene of North America (Afairiguana). I suggest that the placement of the fossil crotaphytid is stable to the uncertain phylogeny of Pleurodonta, but recognize the dynamic nature of fossil diagnosis and the potential for updated systematic hypotheses for the other fossils analysed here.

A new perspective on the taxonomy and systematics of Arvicolinae (Gray, 1821) and a new time-calibrated phylogeny for the clade.

Background: Arvicoline rodents are one of the most speciose and rapidly evolving mammalian lineages. Fossil arvicolines are also among the most common vertebrate fossils found in sites of Pliocene and Pleistocene age in Eurasia and North America. However, there is no taxonomically robust, well-supported, time-calibrated phylogeny for the group. Methods: Here we present well-supported hypotheses of arvicoline rodent systematics using maximum likelihood and Bayesian inference of DNA sequences of two mitochondrial genes and three nuclear genes representing 146 (82% coverage) species and 100% of currently recognized arvicoline genera. We elucidate well-supported major clades, reviewed the relationships and taxonomy of many species and genera, and critically compared our resulting molecular phylogenetic hypotheses to previously published hypotheses. We also used five fossil calibrations to generate a time-calibrated phylogeny of Arvicolinae that permitted some reconciliation between paleontological and neontological data. Results: Our results are largely congruent with previous molecular phylogenies, but we increased the support in many regions of the arvicoline tree that were previously poorly-sampled. Our sampling resulted in a better understanding of relationships within Clethrionomyini, the early-diverging position and close relationship of true lemmings (Lemmus and Myopus) and bog lemmings (Synaptomys), and provided support for recent taxonomic changes within Microtini. Our results indicate an origin of ∼6.4 Ma for crown arvicoline rodents. These results have major implications (e.g., diversification rates, paleobiogeography) for our confidence in the fossil record of arvicolines and their utility as biochronological tools in Eurasia and North America during the Quaternary.

The affinities of the Late Triassic Cryptovaranoides and the age of crown squamates.

Most living reptile diversity is concentrated in Squamata (lizards, including snakes), which have poorly known origins in space and time. Recently, †Cryptovaranoides microlanius from the Late Triassic of the United Kingdom was described as the oldest crown squamate. If true, this result would push back the origin of all major lizard clades by 30-65 Myr and suggest that divergence times for reptile clades estimated using genomic and morphological data are grossly inaccurate. Here, we use computed tomography scans and expanded phylogenetic datasets to re-evaluate the phylogenetic affinities of †Cryptovaranoides and other putative early squamates. We robustly reject the crown squamate affinities of †Cryptovaranoides, and instead resolve †Cryptovaranoides as a potential member of the bird and crocodylian total clade, Archosauromorpha. Bayesian total evidence dating supports a Jurassic origin of crown squamates, not Triassic as recently suggested. We highlight how features traditionally linked to lepidosaurs are in fact widespread across Triassic reptiles. Our study reaffirms the importance of critically choosing and constructing morphological datasets and appropriate taxon sampling to test the phylogenetic affinities of problematic fossils and calibrate the Tree of Life.

Elevational surveys of Sulawesi herpetofauna 1: Gunung Galang, Gunung Dako Nature Reserve.

The Indonesian island of Sulawesi has a unique geology and geography, which have produced an astoundingly diverse and endemic flora and fauna and a fascinating biogeographic history. Much biodiversity research has focused on the regional endemism in the island's Central Core and on its four peninsulas, but the biodiversity of the island's many upland regions is still poorly understood for most taxa, including amphibians and reptiles. Here, we report the first of several planned full-mountain checklists from a series of herpetological surveys of Sulawesi's mountains conducted by our team. In more than 3 weeks of work on Gunung Galang, a 2,254 m peak west of the city of Tolitoli, Sulawesi Tengah Province, on Sulawesi's Northern Peninsula, we recovered nearly fifty species of reptiles and amphibians, more than a dozen of which are either new to science or known but undescribed. The incompleteness of our sampling suggests that many more species remain to be discovered on and around this mountain.

A new extinct species of alligator lizard (Squamata: Elgaria) and an expanded perspective on the osteology and phylogeny of Gerrhonotinae.

BACKGROUND: Alligator lizards (Gerrhonotinae) are a well-known group of extant North American lizard. Although many fossils were previously referred to Gerrhonotinae, most of those fossils are isolated and fragmentary cranial elements that could not be placed in a precise phylogenetic context, and only a handful of known fossils are articulated skulls. The fossil record has provided limited information on the biogeography and phylogeny of Gerrhonotinae. RESULTS: We redescribe a nearly complete articulated fossil skull from the Pliocene sediments of the Anza-Borrego Desert in southern California, and refer the specimen to the alligator lizard genus Elgaria. The fossil is a representative of a newly described species, Elgaria peludoverde. We created a morphological matrix to assess the phylogeny of alligator lizards and facilitate identifications of fossil gerrhonotines. The matrix contains a considerably expanded taxonomic sample relative to previous morphological studies of gerrhonotines, and we sampled two specimens for many species to partially account for intraspecific variation. Specimen-based phylogenetic analyses of our dataset using Bayesian inference and parsimony inferred that Elgaria peludoverde is part of crown Elgaria. The new species is potentially related to the extant species Elgaria kingii and Elgaria paucicarinata, but that relationship was not strongly supported, probably because of extensive variation among Elgaria. We explored several alternative biogeographic scenarios implied by the geographic and temporal occurrence of the new species and its potential phylogenetic placements. CONCLUSIONS: Elgaria peludoverde is the first described extinct species of Elgaria and provides new information on the biogeographic history and diversification of Elgaria. Our research expands the understanding of phylogenetic relationships and biogeography of alligator lizards and strengthens the foundation of future investigations. The osteological data and phylogenetic matrix that we provided will be critical for future efforts to place fossil gerrhonotines. Despite limited intraspecific sampled sizes, we encountered substantial variation among gerrhonotines, demonstrating the value of exploring patterns of variation for morphological phylogenetics and for the phylogenetic placement of fossils. Future osteological investigations on the species we examined and on species we did not examine will continue to augment our knowledge of patterns of variation in alligator lizards and aid in phylogenetics and fossil placement.

Variation in the skulls of Elgaria and Gerrhonotus (Anguidae, Gerrhonotinae) and implications for phylogenetics and fossil identification.

BACKGROUND: There are limited data on intra- and interspecific osteological variation for many squamate clades. Those data are relevant for phylogenetic analyses that use osteological characters and for apomorphic identifications of fossils. We investigate whether morphological features in the skulls of extant gerrhonotine lizards can be used to distinguish taxa at the species- and genus-level and assess whether newly discovered intra- and interspecific osteological variation alters the utility of previously reported apomorphic features. We examined skulls of species belonging to the gerrhonotine genera Elgaria and Gerrhonotus. These genera contain 17 extant species, but the cranial osteology of only a few species was previously examined. As a result, intra- and interspecific osteological variation of these gerrhonotines is poorly understood. METHODS: We employed high-resolution x-ray computed tomography (CT) to scan 25 alcohol-preserved specimens. We provide data on the skulls of all eight species of Elgaria, four for the first time, and five species of Gerrhonotus, three for the first time. We examined 3-D reconstructed skulls of the scanned specimens as well as dry, traditionally prepared skeletons (when they were available). RESULTS: We found that the purported diagnostic utility of many previously described morphological features is impacted because of substantial morphological variation between and within species. We present an assessment of osteological differences that may be useful to differentiate species of Elgaria and Gerrhonotus, many of which are present on isolated cranial elements commonly recovered as fossils, including the premaxilla, maxilla, parietal, pterygoid, prootic, dentary, and surangular. We demonstrate the importance of documenting patterns of osteological variation using large sample sizes, and the utility of examining disarticulated cranial elements of the squamate skull to identify diagnostic morphology. This study adds to a growing body of literature suggesting that extensive documentation of morphological variation is needed to further our understanding of the phylogenetic and diagnostic utility of morphological features across vertebrate clades. Efforts in that direction likely will benefit from examination of disarticulated skeletal elements.

Effects of phylogenetic uncertainty on fossil identification illustrated by a new and enigmatic Eocene iguanian.

Fossil identifications made in a phylogenetic framework are beholden to specific tree hypotheses. Without phylogenetic consensus, the systematic provenance of any given fossil can be volatile. Paleobiogeographic and divergence time hypotheses are contingent on the accurate systematic placement of fossils. Thus, fossil diagnoses should consider multiple topologies when phylogenetic resolution or clear apomorphies are lacking. However, such analyses are infrequently performed. Pleurodonta (Squamata: Iguania) is an ancient and frequently-studied lizard clade for which phylogenetic resolution is notoriously elusive. I describe a skull fossil of a new pleurodontan lizard taxon from the Eocene deposits of the Willwood Formation, Wyoming, and use the new taxon as a case-study to explore the effects of phylogenetic uncertainty on fossil identification. The relationships of the new taxon differ considerably among analyses, and resulting interpretations are correspondingly disparate. These results illustrate generalizable and severe issues with fossil interpretations made without consideration of alternative phylogenetic hypotheses.

Unusual lizard fossil from the Miocene of Nebraska and a minimum age for cnemidophorine teiids.

Teiid lizards are well represented in the fossil record and are common components of modern ecosystems in North and South America. Many fossils were referred to the cnemidophorine teiid group (whiptails, racerunners and relatives), particularly from North America. However, systematic interpretations of morphological features in cnemidophorines were hampered by the historically problematic taxonomy of the clade, and the biogeography and chronology of cnemidophorine evolution in North America is poorly understood from the fossil record. Few fossil cnemidophorines were identified with an apomorphy-based diagnosis, and there are almost no fossil cnemidophorines that could be used to anchor node calibrations. Here, I describe a cnemidophorine from the Miocene Ogallala Group of Nebraska and diagnose the fossil using apomorphies. In that process, I clarify the systematic utility of several morphological features of cnemidophorine lizards. I refer the fossil to the least inclusive clade containing Aspidoscelis, Holcosus and Pholidoscelis. The most conservative minimum age of the locality of the fossil is a fission-track date of 6.3 Ma, but mammal biochronology provides a more refined age of 9.4 Ma, which can be used as a minimum age for the crown cnemidophorine clade in divergence time analyses. The fossil indicates that a cnemidophorine lineage that does not live in Nebraska today inhabited the area during the Miocene. I refrain from naming a new taxon pending discovery of additional fossil material of the lineage to which the fossil belonged.

Combined-evidence analyses of ultraconserved elements and morphological data: an empirical example in iguanian lizards.

Genomic datasets generated by next-generation sequencing are increasingly prevalent in phylogenetics, but morphological data are required to phylogenetically place fossils, corroborate molecular hypotheses and date phylogenies. Combined-evidence analyses provide an integrative assessment of tree topology. However, no attempt has been made to simultaneously analyse next-generation genomic datasets and morphological data, and the future of morphology in the context of genomic data is uncertain. I conducted combined-evidence analyses that include genomic and morphological datasets, specifically, with ultraconserved elements and two morphological matrices. In unweighted maximum-likelihood and Bayesian combined-evidence analyses, morphological signal was dwarfed by the ultraconserved elements, and some node support values were reduced relative to ultraconserved element-only analyses. Increasing the weight of morphological characters allowed those data to influence the tree, but weighting subjectivity should be considered in future analyses. More attempts should be made to simultaneously analyse genomic and morphological datasets.

Novel and Bizarre Abnormalities of the Tooth Row in Side-Blotched Lizards (Uta) and Rock Lizards (Petrosaurus).

Multiple tooth rows along the dentary or maxilla are present in many bony and cartilaginous fishes but occur infrequently in amniotes. Although two partially overlapping rows of teeth or a few doubled tooth positions were both previously reported in extant and extinct lizards, multiple rows of teeth across the entire dental or maxillary shelf were not previously documented in any lizard taxon. Here, we report one specimen of the side-blotched lizard Uta stansburiana and two specimens of the rock lizard Petrosaurus mearnsi that have two or more marginal tooth rows that overlap for most of the dental shelf. We also describe several other phrynosomatid lizard specimens with one or two doubled tooth positions per marginal tooth-bearing skeletal element. We examined 520 total specimens of pleurodont iguanian lizards and verified that aberrant tooth row and tooth position morphologies were almost exclusively present in phrynosomatid lizards. We hypothesize that developmental irregularities in the number of odontogenic bands or in the function of zones of inhibition may have resulted in the observed tooth row abnormalities. The expression of multiple tooth rows or doubled tooth positions may be phylogenetically informative morphologies of phrynosomatid lizards. This bears further investigation from developmental, genetic, ecological, and phylogenetic perspectives. Anat Rec, 2019. © 2019 American Association for Anatomy Anat Rec, 303:2014-2025, 2020. © 2019 American Association for Anatomy.

The first known fossil Uma: ecological evolution and the origins of North American fringe-toed lizards.

BACKGROUND: Fossil evidence suggests that extant North American lizard genera (north of Mexico) evolved during the Miocene. Although fossils of the clade Phrynosomatidae (spiny lizards and sand lizards) have been reported, there have been no previously described fossils of the fringe-toed sand lizards (Uma). In the extant biota, Uma inhabit arid deserts, and members of the western clade of Uma are restricted to sand dunes or other habitats containing fine-grained sand. RESULTS: I describe the first known fossil of Uma and refer the fossil to the total clade of Uma with an apomorphy-based diagnosis. The fossil is a partial premaxilla that was found in the Miocene strata of the Dove Spring Formation of southern California, dating to 8.77 Ma. The paleoenvironment of the Dove Spring Formation was semiarid and contained ephemeral streams that facilitated deposition, and there is no evidence of sand dune deposits in the strata containing the locality from which the Uma fossil was found. Divergence time analyses of a concatenated molecular dataset with four fossil calibrations support a Neogene origin of the total clade of Uma and of the crown clade of Uma. Those analyses also estimated a Neogene divergence between Uma scoparia and the Uma notata complex. Multispecies coalescent analyses with one fossil calibration inferred a Paleogene origin for the total clade of Uma and a Pliocene or Pleistocene divergence between Uma scoparia and the Uma notata complex. The fossil and the total and crown clades of Uma precede the evolution of modern desert ecosystems in the southwestern United States and northern Mexico by millions of years. CONCLUSIONS: The total clade and the crown clade of Uma were not restricted to arid deserts throughout their evolutionary histories. I demonstrate that an apomorphy-based diagnosis can be used to identify fossils of isolated skeletal elements for at least one clade of phrynosomatid lizard, and suggest exercising caution when using environmental tolerances of extant taxa to hypothesize paleoecological reconstructions.

The skull of the gerrhonotine lizard Elgaria panamintina (Squamata: Anguidae).

We provide the first description of the skull, osteoderms, and hyoid apparatus of the poorly known alligator lizard Elgaria panamintina, and compare the cranial osteology of that species to the widespread and well-studied taxon Elgaria multicarinata. Patterns of morphological variation resulting from ontogenetic transformations and pathology are discussed. We employed x-ray computed tomography (CT) scans to examine two adult specimens of Elgaria panamintina and two adult specimens of Elgaria multicarinata, in addition to examining multiple traditionally prepared skeletal specimens of the latter species. CT scans provide simultaneous study of both articulated and disarticulated elements, allowing us to describe and document the morphology of the skull with exceptional precision and detail. The description of the skull of Elgaria panamintina serves as a generalization for all Elgaria; here we provide the first complete description of the skull of this genus for future uses in morphological and phylogenetic studies of both extant species and fossils.