Phylogenetic Classification at Generic Level in the Absence of Distinct Phylogenetic Patterns of Phenotypical Variation: A Case Study in Graphidaceae (Ascomycota)

Molecular phylogenies often reveal that taxa circumscribed by phenotypical characters are not monophyletic. While re-examination of phenotypical characters often identifies the presence of characters characterizing clades, there is a growing number of studies that fail to identify diagnostic characters, especially in organismal groups lacking complex morphologies. Taxonomists then can either merge the groups or split taxa into smaller entities. Due to the nature of binomial nomenclature, this decision is of special importance at the generic level. Here we propose a new approach to choose among classification alternatives using a combination of morphology-based phylogenetic binning and a multiresponse permutation procedure to test for morphological differences among clades. We illustrate the use of this method in the tribe Thelotremateae focusing on the genus Chapsa, a group of lichenized fungi in which our phylogenetic estimate is in conflict with traditional classification and the morphological and chemical characters do not show a clear phylogenetic pattern. We generated 75 new DNA sequences of mitochondrial SSU rDNA, nuclear LSU rDNA and the protein-coding RPB2. This data set was used to infer phylogenetic estimates using maximum likelihood and Bayesian approaches. The genus Chapsa was found to be polyphyletic, forming four well-supported clades, three of which clustering into one unsupported clade, and the other, supported clade forming two supported subclades. While these clades cannot be readily separated morphologically, the combined binning/multiresponse permutation procedure showed that accepting the four clades as different genera each reflects the phenotypical pattern significantly better than accepting two genera (or five genera if splitting the first clade). Another species within the Thelotremateae, Thelotrema petractoides, a unique taxon with carbonized excipulum resembling Schizotrema, was shown to fall outside Thelotrema. Consequently, the new genera Astrochapsa, Crutarndina, Pseudochapsa, and Pseudotopeliopsis are described here and 39 new combinations are proposed.     

Systematic implication of seed micromorphology in Arenaria (Caryophyllaceae) and allied genera

Variations in seed micromorphology of 64 taxa of Arenaria and allied genera were examined using scanning electron microscopy (SEM) in order to evaluate their diagnostic value for systematic studies in the genus and providing additional evidence on delimitation of natural groups. Significant diversity was found in seed coat morphology both among and within genera. Size, color, and shape of seeds appear to be of low importance at species level but provide useful evidence at generic rank. Cerastium and Stellaria are well distinguished from Arenaria by serrate seed margins and solid columellae on testa cells. The sculpturing pattern renders most characters potentially applicable in taxonomy and phylogeny assessment in Arenaria. The studied taxa show four basic types of sculpturing, viz. colliculate, rugose, papillate, and smooth. The colliculate type which is characteristic for the core group of Arenaria (A. subg. Arenaria) can be further divided into five subtypes. The smooth seed surface with indistinct cell boundaries represents a synapomorphy for A. subg. Leiosperma, whereas the winged seeds along with rugose surface characterize subg. Odontostemma. Otherwise seed micromorphology rarely provides adequate evidence for separation of natural groups, especially at sectional rank. We hypothesize that the seeds are highly polymorphic in certain groups of taxa showing recent adaptive radiation, such as the annual species of Arenaria, and do not provide strong support for natural groups within the genus.     

Phylogeny of the Archiborborinae (Diptera: Sphaeroceridae) Based on Combined Morphological and Molecular Analysis

The Archiborborinae is a diverse Neotropical subfamily of Sphaeroceridae, with many undescribed species. The existing generic classification includes three genera consisting of brachypterous species, with all other species placed in the genus Archiborborus. We present the first phylogenetic hypothesis for the subfamily based on morphological, molecular, and combined datasets. Morphological data include 53 characters and cover all valid described taxa (33 species in 4 genera) in the subfamily, as well as 83 undescribed species. Molecular data for five genes (mitochondrial 12S rDNA, cytochrome c oxidase subunit I, and cytochrome B, and nuclear alanyl-tRNA synthetase and 28S rDNA) were obtained for 21 ingroup taxa. Data support the separation of the Archiborborinae from the Copromyzinae, with which they were formerly combined. Analyses support consistent groups within the subfamily, but relationships between groups are poorly resolved. The validity of the brachypterous genera Penola Richards and Frutillaria Richards is supported. The former genus Archiborborus Duda is paraphyletic, and will be divided into monophyletic genera on the basis of this work. Aptery and brachyptery have evolved multiple times in the subfamily. Antrops Enderlein, previously including a single brachypterous species, is a senior synonym of Archiborborus.     

Systematic implications of flavonoid pigments in the fern genus hemionitis (adiantiaceae)

The closely related fern generaHemionitis L. andGymnopteris Bernhardi are separated primarily on differences in leaf architecture and venation. Studies indicate that these characters are highly variable and unreliably diagnostic. Further, the type species of the two genera readily hybridize with each other. Spore morphology, as exhibited by SEM, does not support the traditional alignment of the species in these two genera: some species ofHemionitis andGymnopteris have the same rugose to papillate spores, while other species from both genera possess crested spores. The flavonoid chemistry of these taxa coincides with spore type, i.e., taxa from both genera which possess crested spores produce kaempferol and quercetin 3-0-glycosides, while species with tuberculate spores produce only quercetin 3,4′-0-glycosides. The spore and chemical data suggest a realignment of these taxa within a single genus, which would avoid the rather tenuous dependence on a single vegetative character for generic distinctions.     

Partitioning core and satellite taxa from within cystic fibrosis lung bacterial communities

Cystic fibrosis (CF) patients suffer from chronic bacterial lung infections that lead to death in the majority of cases. The need to maintain lung function in these patients means that characterising these infections is vital. Increasingly, culture-independent analyses are expanding the number of bacterial species associated with CF respiratory samples; however, the potential significance of these species is not known. Here, we applied ecological statistical tools to such culture-independent data, in a novel manner, to partition taxa within the metacommunity into core and satellite species. Sputa and clinical data were obtained from 14 clinically stable adult CF patients. Fourteen rRNA gene libraries were constructed with 35 genera and 82 taxa, identified in 2139 bacterial clones. Shannon–Wiener and taxa-richness analyses confirmed no undersampling of bacterial diversity. By decomposing the distribution using the ratio of variance to the mean taxon abundance, we partitioned objectively the species abundance distribution into core and satellite species. The satellite group comprised 67 bacterial taxa from 33 genera and the core group, 15 taxa from 7 genera (including Pseudomonas (1 taxon), Streptococcus (2), Neisseria (2), Catonella (1), Porphyromonas (1), Prevotella (5) and Veillonella (3)], the last four being anaerobes). The core group was dominated by Pseudomonas aeruginosa. Other recognised CF pathogens were rare. Mantel and partial Mantel tests assessed which clinical factors influenced the composition observed. CF transmembrane conductance regulator genotype and antibiotic treatment correlated with all core taxa. Lung function correlated with richness. The clinical significance of these core and satellite species findings in the CF lung is discussed.     

Phylogenetically balanced evidence for structural and carbon isotope responses in plants along elevational gradients

We tested three hypotheses related to the functioning of mountain plants, namely their reproductive effort, leaf surface structure and effectiveness of CO₂ assimilation, using archive material from contrasting elevations. Analysis of elevational trends is at risk of suffering from two major biases: a phylogenetic bias (i.e. an elevational change in the abundance of taxonomic groups), and covariation of different environmental drivers (e.g. water, temperature, atmospheric pressure), which do not permit a mechanistic interpretation. We solved both problems in a subcontinental survey of elevational trends in key plant traits in the European Alps and the high Arctic (northern Sweden, Svalbard), using herbarium samples of 147 species belonging to the genera Carex, Saxifraga and Potentilla. We used both species and phylogenetically independent contrasts as data points. The analysis revealed enhanced reproductive efforts at higher elevation in insect-pollinated taxa (not in wind-pollinated taxa), no increase in leaf pubescence at high elevation (as is often assumed), and a strong correlation between ¹³C discrimination and elevation. Alpine taxa operate at a smaller mesophyll resistance to CO₂ uptake relative to diffusive resistance (stomata). By comparison with congeneric low altitude polar taxa (low temperature, but high atmospheric pressure), the response could be attributed to the elevational decline in atmospheric pressure rather than temperature (a mean increase in δ¹³C by 1.4‰ km^?1). The signal is consistent within and across genera and within species, suggesting rapid adjustment of leaf physiology to reduced partial pressure of CO₂. These results offer answers to long-debated issues of plant responses to high elevation life conditions.     

Genic differentiation and phylogenetic relationships within two New World bat genera

Phylogenetic placement of taxa within the two New World leaf-nosed bat genera, Tonatia and Micronycteris was carried out using allozymic characters. Of particular interest were the relationships between the karyotypically megaevolved species and their proposed congeners. Neither of the two karyotypically megaevolved Tonatia species shared synapomorphic allozymes with the other Tonatia taxa. In contrast, the three Micronycteris species identified as having undergone this unusual mode of karyotypic evolution fall within a common evolutionary assemblage with all but one of their congeners. Measures of genic similarity among the Tonatia and Micronycteris species yield relatively low Nei identity values. No correlation is found when the number of karyotypic and allozymic changes since a common ancestor are compared.     

Evaluating DNA barcoding criteria using African duiker antelope (Cephalophinae) as a test case

African duikers in the subfamily Cephalophinae (genera Cephalophus, Philantomba and Sylvicapra) constitute an important target for DNA barcoding efforts because of their importance to the bushmeat trade and protection under the Convention for International Trade in Endangered Species (CITES). Duikers also make a challenging test case of barcoding methods due to their recent diversification, substantial intra-specific genetic variation and high species richness. However, no study to date has evaluated how well DNA barcoding methods can be used to delineate all of the taxa within this group. To address this question, cytochrome c oxidase subunit 1 (COX1) sequences from all eighteen species within this subfamily and an outgroup taxon (genus Tragelaphus) were used to build a neighbor-joining tree, identify species-specific diagnostic synapomorphies, and determine whether species exceed a given pair-wise genetic distance threshold commonly employed in DNA barcoding studies. Tree-based analyses of the data indicate that several species within two clusters of closely related taxa consistently failed to form reciprocally monophyletic clades and similarly lack species-specific synapomorphies. Furthermore, one additional taxon failed to constitute a diagnosable clade and another occupied an unresolved position in the tree. Of the two genetic distance criteria evaluated, the 3% threshold was far more effective in delimiting species than a threshold level based on the ratio of inter- to intra-specific distances. However, neither approach could effectively delineate all sister species. While the taxonomy of this group might be open to question, the fact that barcodes consistently failed to differentiate several currently recognized sister taxa challenges the routine application of this approach in forensic studies of duiker species. Future barcoding work of this group should always include a complete taxonomic sampling and strive to include a broader geographic sampling of sequence diversity than has been carried out to date. Lastly, this work highlights the need to re-examine the taxonomy of this group, which may illuminate why some barcoding criteria fail to reliably differentiate species.     

Predicting Plastid Marker Variation: Can Complete Plastid Genomes from Closely Related Species Help?

Rapidly evolving non-coding plastid regions (NCPs) are currently widely used in evolutionary biology especially in plant systematic studies where NCPs have become one of the most commonly used tools in clarifying species relationships. Currently, the generally small amount of sequence variation provided by NCPs compared to nuclear regions makes plastid phylogeny reconstruction challenging at the species-level, especially so in species rich clades such as Solanum with c. 1,200 species. Previous studies have established that the set of most highly variable NCPs vary between major plant families, and here we explore whether this variation extends beyond family level to genera and major clades within genera. Using full plastome data, we identify the most highly variable plastid markers in the Potato clade of Solanum. We then compare sequence variation between the Potato and the closely related Morelloid clade. Results show that whilst a narrow set of NCPs show consistently high variation, levels of sequence variation in most NCPs differ greatly between the two closely related clades. The high variation detected between closely related groups implies that repeated screening studies will be needed for individual projects despite the potential availability of results from closely related taxa, and indicates a narrower applicability of family-specific screening studies than previously thought.     

Cryptomyrus: a new genus of Mormyridae (Teleostei,Osteoglossomorpha) with two new species from Gabon,West-Central Africa

We use mitochondrial and nuclear sequence data to show that three weakly electric mormyrid fish specimens collected at three widely separated localities in Gabon, Africa over a 13-year period represent an unrecognized lineage within the subfamily Mormyrinae and determine its phylogenetic position with respect to other taxa. We describe these three specimens as a new genus containing two new species. Cryptomyrus, new genus, is readily distinguished from all other mormyrid genera by a combination of features of squamation, morphometrics, and dental attributes. Cryptomyrus ogoouensis, new species, is differentiated from its single congener, Cryptomyrus ona, new species, by the possession of an anal-fin origin located well in advance of the dorsal fin, a narrow caudal peduncle and caudal-fin lobes nearly as long as the peduncle. In Cryptomyrus ona, the anal-fin origin is located only slightly in advance of the dorsal fin, the caudal peduncle is deep and the caudal-fin lobes considerably shorter than the peduncle. Continued discovery of new taxa within the “Lower Guinea Clade” of Mormyridae highlights the incompleteness of our knowledge of fish diversity in West-Central Africa. We present a revised key to the mormyrid genera of Lower Guinea.     

The composition and structure of the benthic diatom taxocene (Bacillariophyta) near Cape Fiolent (the Crimea,the Black Sea)

For the first time in the waters of the Cape Fiolent Reserve, a high species richness of benthic diatoms was revealed on natural substrates, 290 species and infraspecific taxa (IT), belonging to 281 species, 68 genera, 32 families, 19 orders, and 3 classes. A total of 68 species and 3 genera were previously registered as new for the flora of the Black Sea; 3 species and 1 taxonomic combination were previously described as new to science, as well as 4 species that did not occur in the Black sea in the past 50 and 100 years of research. Navicula (41 species and IT), Amphora (30), Nitzschia (29), Fallacia (20), Cocconeis (17) and Diploneis (16) were the most represented genera. Only one representative was identified in each of the orders Anaulales, Ardissoneales, Thalassionematales, Rhaphoneidales, and Rhopalodiales. The diversity assessment that was carried out using taxonomic distinctness indices, TaxDI, revealed that the structure of the phylogenetic diatom tree was formed by branches with various species saturation levels and different hierarchical subordination, but with the predominance of polyspecific taxa up to at genera level. The TaxDI values (Δ⁺ = 76.71; Λ⁺ = 361.52) indicate the low evenness and high variability of the structure of the benthic diatom taxocene near Cape Fiolent.     

Phylogenetic interrelationships of ginglymodian fishes (Actinopterygii: Neopterygii)

The Ginglymodi is one of the most common, though poorly understood groups of neopterygians, which includes gars, macrosemiiforms, and “semionotiforms.” In particular, the phylogenetic relationships between the widely distributed “semionotiforms,” and between them and other ginglymodians have been enigmatic. Here, the phylogenetic relationships between eight of the 11 “semionotiform” genera, five genera of living and fossil gars and three macrosemiid genera, are analysed through cladistic analysis, based on 90 morphological characters and 37 taxa, including 7 out-group taxa. The results of the analysis show that the Ginglymodi includes two main lineages: Lepisosteiformes and †Semionotiformes. The genera †Pliodetes, †Araripelepidotes, †Lepidotes, †Scheenstia, and †Isanichthys are lepisosteiforms, and not semionotiforms, as previously thought, and these taxa extend the stratigraphic range of the lineage leading to gars back up to the Early Jurassic. A monophyletic †Lepidotes is restricted to the Early Jurassic species, whereas the strongly tritoral species previously referred to †Lepidotes are referred to †Scheenstia. Other species previously referred to †Lepidotes represent other genera or new taxa. The macrosemiids are well nested within semionotiforms, together with †Semionotidae, here restricted to †Semionotus, and a new family including †Callipurbeckia n. gen. minor (previously referred to †Lepidotes), †Macrosemimimus, †Tlayuamichin, †Paralepidotus, and †Semiolepis. Due to the numerous taxonomic changes needed according to the phylogenetic analysis, this article also includes formal taxonomic definitions and diagnoses for all generic and higher taxa, which are new or modified. The study of Mesozoic ginglymodians led to confirm Patterson’s observation that these fishes show morphological affinities with both halecomorphs and teleosts. Therefore, the compilation of large data sets including the Mesozoic ginglymodians and the re-evaluation of several hypotheses of homology are essential to test the hypotheses of the Halecostomi vs. the Holostei, which is one of the major topics in the evolution of Mesozoic vertebrates and the origin of modern fish faunas.     

Plant diversity of Hyrcanian relict forests: An annotated checklist,chorology and threat categories of endemic and near endemic vascular plant speciesOA

In this paper a critical annotated checklist of 256 endemic and near endemic species belonging to 152 genera and 50 families of flowering plants known from Hyrcanian relict forests is presented. Distribution maps of taxa, elevational range, number of known records, chorotypes, life forms, IUCN threat categories and habitat types are also provided. The chorotypes are categorized into eight main patterns: 1) the Omni-Hyrcanian pattern(OH), 2) West Hyrcanian pattern(WH), 3) Manjil-Rudbar pattern(MR...     

The phylogeny and taxonomy of genera Cystoderma and Cystodermella (Agaricales) based on nuclear ITS and LSU sequences

Species delimitation in Cystoderma and Cystodermella was evaluated based on ITS and LSU rDNA sequences as well as morphological data. Two species of Cystoderma are synonymised with C. carcharias and three species with C. jasonis, distinguishing the synonymised taxa as varieties of these accepted species. Analyses of partial LSU rDNA sequences revealed Cystoderma and Cystodermella as distinct monophyletic genera, with Ripartitella representing a well-supported sister group of the latter. Phaeolepiota aurea represents either an unsupported sister group or member of Cystoderma in the phylogenies based on LSU and ITS sequences rDNA data, respectively. The tribe Cystodermateae sensu Singer did not appear monophyletic according to analyses of LSU sequences. On the basis of these data, the phylogenetic relationships among most of the analyzed genera could not be resolved unequivocally.     

Detailed cytomorphological investigations through male meiosis of polypetalous plants from the Kashmir Himalaya

Cytomorphological studies were made on 322 populations of 150 species/153 taxa belonging to 79 genera and 23 families of flowering plants. Out of these, 14 species were cytologically worked out for the first time on a worldwide basis along with B-chromosomes (0–1B) recorded for the first time in Agrimonia eupatoria. Similarly, 29 species were reported with varied chromosomal reports on a worldwide basis. The chromosome numbers ranged from 2n = 10 to 2n = 96 and posed high amount of intraspecific genetic diversity when seen in relation to other reports for the worked out species in light of previous chromosomal databases on India and world basis. The different levels of ploidy in these species/taxa vary from 2x to 12x with the overall polyploidy being 27.45 % and predominant in members belonging to the genera Ranunculus and Epilobium as well as families Ranunculaceae and Rosaceae. In the taxa showing anomalous meiotic behavior, cytomixis is found to be quite common and associated with various meiotic abnormalities in the meiocytes, which ultimately leads to pollen sterility and formation of variable sized pollen grains.     

Zeins as indicators of maize-Tripsacum introgression

A survey of zeins in tripsacoid and non-tripsacoid races of maize from Mesoamerica and from South America, annual teosinte, perennial species of Zea and species of Tripsacum revealed at least 33 zein proteins as determined by isoelectric focusing. Zea and Tripsacum and generally also species within these genera are characterized by distinct combinations of zein proteins. Maize is extensively heterogenous, and spans the complete spectrum of zeins present in wild Zea taxa. A comparison of zein proteins failed to distinguish between introgression of maize with Tripsacum or teosinte. The ease with which maize crosses naturally with wild Zea taxa, and the rarity of hybrids with Tripsacum essentially rule out natural Tripsacum introgression as a mode of racial evolution in maize.     

Helminth communities of herons (Aves: Ardeidae) in southern Italy

The helminth communities of nine species of herons from southern Italy were studied and compared. Of 24 taxa found including seven digeneans, seven nematodes, six cestodes and four acanthocephalans, only five taxa were found in more than one heron species, and five of the 21 taxa that could be identified to species level were classified as ‘heron specialists’. The total number of helminth species per heron species ranged from 1 in Botaurus stellaris to 9 in Ixobrychus minutus with infection levels generally low. A statistical comparison was carried out for herons with a sample size > 5. At the infracommunity level, only I. minutus clearly differed from other heron species. Diversity parameters of heminth infracommunities did not significantly differ among heron species. Species richness ranged from just 0.3 to 2.3 helminth taxa per individual host, and the Brillouin index, from 0 to 0.3. Total helminth abundance did not exceed 40 worms per host except in a single case of Ardeola ralloides. Infracommunities clearly were dominated by single helminth species. The present study confirms a depauperate helminth community in herons from southern Italy. Comparison with data from Spain and the Czech Republic showed strong quantitative similarities with values obtained in the present study. Results also suggest that the composition of local helminth communities are strongly variable depending on geographical location as is demonstrated by comparison with data from other European areas. However, whether herons in Europe naturally host depauperate helminth communities or these communities are depauperate because of other factors is unknown.     

Reverse evolution and cryptic diversity in putative sister families of the Oribatida (Acari)

The family Zetorchestidae is a morphologically and ecologically diverse group assigned to the higher oribatid mites (Brachypylina). We addressed the phylogeny of the family by including species of the genera Belorchestes, Litholestes, Microzetorchestes and Zetorchestes. We also analysed the affinities of the putative sister taxon (Eremaeidae), investigating Eremaeus and Eueremaeus. Zetorchestidae, Eremaeidae and Niphocepheidae were recently combined in one superfamily (Zetorchestoidea). These taxa were placed into a wider phylogenetic context by adding other presumably closely related taxa. Phylogenetic analyses based upon nuclear and mitochondrial DNA‐sequences revealed the monophyly of the Zetorchestidae as well as of all investigated species and genera of this family. Ancestral state reconstruction of jumping ability in latter family, moreover, suggested reverse character evolution within the studied zetorchestid taxa. Genetic diversity of the genera Eremaeus and Eueremaeus turned out to be higher than known, suggesting the existence of cryptic species. However, none of our analyses supported a sister group relationship among Zetorchestidae and Eremaeidae. Moreover, all calculated trees show a paraphyletic position between Zetorchestidae respectively Eremaeidae and Niphocepheidae.     

Dated Phylogenies of the Sister Genera Macaranga and Mallotus (Euphorbiaceae): Congruence in Historical Biogeographic Patterns?

Molecular phylogenies and estimates of divergence times within the sister genera Macaranga and Mallotus were estimated using Bayesian relaxed clock analyses of two generic data sets, one per genus. Both data sets were based on different molecular markers and largely different samples. Per genus three calibration points were utilised. The basal calibration point (crown node of all taxa used) was taken from literature and used for both taxa. The other three calibrations were based on fossils of which two were used per genus. We compared patterns of dispersal and diversification in Macaranga and Mallotus using ancestral area reconstruction in RASP (S-DIVA option) and contrasted our results with biogeographical and geological records to assess accuracy of inferred age estimates. A check of the fossil calibration point showed that the Japanese fossil, used for dating the divergence of Mallotus, probably had to be attached to a lower node, the stem node of all pioneer species, but even then the divergence time was still younger than the estimated age of the fossil. The African (only used in the Macaranga data set) and New Zealand fossils (used for both genera) seemed reliably placed. Our results are in line with existing geological data and the presence of stepping stones that provided dispersal pathways from Borneo to New Guinea-Australia, from Borneo to mainland Asia and additionally at least once to Africa and Madagascar via land and back to India via Indian Ocean island chains. The two genera show congruence in dispersal patterns, which corroborate divergence time estimates, although the overall mode and tempo of dispersal and diversification differ significantly as shown by distribution patterns of extant species.     

Phylogenetic relationships of phrynosomatid lizards based on nuclear and mitochondrial data,and a revised phylogeny for Sceloporus

Phrynosomatid lizards are among the most common and diverse groups of reptiles in western North America, Mexico, and Central America. Phrynosomatidae includes 136 species in 10 genera. Phrynosomatids are used as model systems in many research programs in evolution and ecology, and much of this research has been undertaken in a comparative phylogenetic framework. However, relationships among many phrynosomatid genera are poorly supported and in conflict between recent studies. Further, previous studies based on mitochondrial DNA sequences suggested that the most species-rich genus (Sceloporus) is possibly paraphyletic with respect to as many as four other genera (Petrosaurus, Sator, Urosaurus, and Uta). Here, we collect new sequence data from five nuclear genes and combine them with published data from one additional nuclear gene and five mitochondrial gene regions. We compare trees from nuclear and mitochondrial data from 37 phrynosomatid taxa, including a “species tree” (from BEST) for the nuclear data. We also present a phylogeny for 122 phrynosomatid species based on maximum likelihood analysis of the combined data, which provides a strongly-supported hypothesis for relationships among most phrynosomatid genera and includes most phrynosomatid species. Our results strongly support the monophyly of Sceloporus (including Sator) and many of the relationships within it. We present a new classification for phrynosomatid lizards and the genus Sceloporus, and offer a new tree with branch lengths for use in comparative studies.