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.
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.
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.
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DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
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
, 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 (
), stem Pleurodonta in the Cretaceous of North America (
) and a stem anole in the Eocene of North America (
). 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.
ABSTRACT Motivated by the ongoing Spitzer observational campaign, and the forthcoming K2 one, we revisit, working in an heliocentric reference frame, the geometrical foundation for the analysis of ...the microlensing parallax, as measured with the simultaneous observation of the same microlensing event from two observers with relative distance of order au. For the case of observers at rest, we discuss the well-known fourfold microlensing parallax degeneracy and determine an equation for the degenerate directions of the lens trajectory. For the case of observers in motion, we write down an extension of the Gould relationship between the microlensing parallax and the observable quantities and, at the same time, highlight the functional dependence of these same quantities from the timescale of the underlying microlensing event. Furthermore, through a series of examples, we show the importance of taking into account themotion of the observers to correctly recover the parameters of the underlying microlensing event. In particular, we discuss the cases of the amplitude of the microlensing parallax and that of the difference of the timescales between the observed microlensing events, which are key to understand the breaking of the microlensing parallax degeneracy. Finally, we consider the case of the simultaneous observation of the same microlensing event from the ground and two satellites, a case relevant for the expected joint K2 and Spitzer observational programs in 2016.
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.
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DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
We present a new analysis of the results of the Expérience pour la Recherche d'Objets Sombres (EROS)-2, Optical Gravitational Lensing Experiment (OGLE)-II and OGLE-III microlensing campaigns towards ...the Small Magellanic Clouds (SMC). Through a statistical analysis we address the issue of the nature of the reported microlensing candidate events, whether to be attributed to lenses belonging to known population the SMC luminous components or the Milky Way disc, to which we broadly refer to as 'self-lensing' or to the would be population of dark matter compact halo objects (MACHOs). To this purpose, we present profiles of the optical depth and, comparing to the observed quantities, we carry out analyses of the events position and duration. Finally, we evaluate and study the microlensing rate. Overall, we consider five reported microlensing events towards the SMC (one by EROS and four by OGLE). The analysis shows that in terms of number of events the expected self-lensing signal may indeed explain the observed rate. However, the characteristics of the events, spatial distribution and duration (and for one event, the projected velocity) rather suggest a non-self-lensing origin for a few of them. In particular, we evaluate, through a likelihood analysis, the resulting upper limit for the halo mass fraction in form of MACHOs given the expected self-lensing and MACHO lensing signal. At 95 per cent CL, the tighter upper limit, about 10 per cent, is found for MACHO mass of 10−2 M, upper limit that reduces to above 20 per cent for 0.5 M MACHOs.
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.
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.
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 (
and
) and bog lemmings (
), 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.
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.
We present high-precision photometry of two transit events of the extrasolar planetary system WASP-5, obtained with the Danish 1.54-m telescope at European Southern Obseratory La Silla. In order to ...minimize both random and flat-fielding errors, we defocused the telescope so its point spread function approximated an annulus of diameter 40 pixel (16 arcsec). Data reduction was undertaken using standard aperture photometry plus an algorithm for optimally combining the ensemble of comparison stars. The resulting light curves have point-to-point scatters of 0.50 mmag for the first transit and 0.59 mmag for the second. We construct detailed signal-to-noise ratio calculations for defocused photometry, and apply them to our observations. We model the light curves with the jktebop code and combine the results with tabulated predictions from theoretical stellar evolutionary models to derive the physical properties of the WASP-5 system. We find that the planet has a mass of Mb= 1.637 ± 0.075 ± 0.033 MJup, a radius of Rb= 1.171 ± 0.056 ± 0.012 R Jup, a large surface gravity of gb= 29.6 ± 2.8 m s−2 and a density of ρb= 1.02 ± 0.14 ± 0.01 ρJup (statistical and systematic uncertainties). The planet's high equilibrium temperature of Teq= 1732 ± 80 K makes it a good candidate for detecting secondary eclipses.
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.
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.
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.
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
and
. 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.
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
, four for the first time, and five species of
, 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).
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
and
, 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.
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