Comparative Evolution of an Archetypal Adaptive Radiation: Innovation and Opportunity in Anolis Lizards.

Adaptive radiation is a widely recognized pattern of evolution wherein substantial phenotypic change accompanies rapid speciation. Adaptive radiation may be triggered by environmental opportunities resulting from dispersal to new areas or via the evolution of traits, called key innovations, that allow for invasion of new niches. Species sampling is a known source of bias in many comparative analyses, yet classic adaptive radiations have not been studied comparatively with comprehensively sampled phylogenies. In this study, we use unprecedented comprehensive phylogenetic sampling of Anolis lizard species to examine comparative evolution in this well-studied adaptive radiation. We compare adaptive radiation models within Anolis and in the Anolis clade and a potential sister lineage, the Corytophanidae. We find evidence for island (i.e., opportunity) effects and no evidence for trait (i.e., key innovation) effects causing accelerated body size evolution within Anolis. However, island effects are scale dependent: when Anolis and Corytophanidae are analyzed together, no island effect is evident. We find no evidence for an island effect on speciation rate and tenuous evidence for greater speciation rate due to trait effects. These results suggest the need for precision in treatments of classic adaptive radiations such as Anolis and further refinement of the concept of adaptive radiation.

A Phylogenetic, Biogeographic, and Taxonomic study of all Extant Species of Anolis (Squamata; Iguanidae).

Anolis lizards (anoles) are textbook study organisms in evolution and ecology. Although several topics in evolutionary biology have been elucidated by the study of anoles, progress in some areas has been hampered by limited phylogenetic information on this group. Here, we present a phylogenetic analysis of all 379 extant species of Anolis, with new phylogenetic data for 139 species including new DNA data for 101 species. We use the resulting estimates as a basis for defining anole clade names under the principles of phylogenetic nomenclature and to examine the biogeographic history of anoles. Our new taxonomic treatment achieves the supposed advantages of recent subdivisions of anoles that employed ranked Linnaean-based nomenclature while avoiding the pitfalls of those approaches regarding artificial constraints imposed by ranks. Our biogeographic analyses demonstrate complexity in the dispersal history of anoles, including multiple crossings of the Isthmus of Panama, two invasions of the Caribbean, single invasions to Jamaica and Cuba, and a single evolutionary dispersal from the Caribbean to the mainland that resulted in substantial anole diversity. Our comprehensive phylogenetic estimate of anoles should prove useful for rigorous testing of many comparative evolutionary hypotheses. [Anoles; biogeography; lizards; Neotropics; phylogeny; taxonomy].

Characters of external morphology used in Anolis taxonomy-definition of terms, advice on usage, and illustrated examples.

With the aim of establishing a reference for taxonomists, I describe and discuss characters of external morphology that have been used widely in descriptions of Anolis (sensu lato) species. For most characters, images are provided to illustrate the terms and definitions. Where appropriate, I give advice on how the counts and measurements should be done.

Anolis boulengerianus Thominot, 1887, a senior synonym of Anolis isthmicus Fitch, 1978 (Squamata: Dactyloidae).

For most of its history, the name Anolis boulengerianus Thominot, 1887 has been regarded as a junior synonym of Anolis nebulosus (Wiegmann, 1834) or Anolis nebuloides Bocourt, 1873. However, a comparison of the syntypes and additional topotypical specimens of A. boulengerianus with three topoparatypes and additional specimens of the species currently referred to as Anolis isthmicus Fitch, 1978 showed that these two names pertain to the same species. Because of the priority principle, A. isthmicus becomes a junior synonym of A. boulengerianus. We provide a redescription of the type series of A. boulengerianus and an updated diagnosis for this taxon relative to all other beta anoles.

A new species of pine anole from the Sierra Madre del Sur in Oaxaca, Mexico (Reptilia, Squamata, Dactyloidae: Anolis).

We describe the new species Anolis peucephilus sp. nov. from the Pacific versant of southern Mexico. Anolis peucephilus differs from all congeners by having a combination of (1) smooth ventral scales; (2) usually a patch of three greatly enlarged supraocular scales; (3) extremely short hind legs, longest toe of adpressed hind leg reaching to a point between levels of axilla and ear opening, ratio shank length/snout-vent length 0.18-0.21; (4) circumnasal usually in contact with first supralabial; and (5) a large yellowish orange dewlap in males and a very small to small white dewlap in females. In external morphology, A. peucephilus is most similar to A. omiltemanus from which it differs by having even shorter hind legs with the longest toe of adpressed hind leg reaching to a point between levels of axilla and ear opening (versus usually to ear opening, occasionally to slightly beyond ear opening or to a point between shoulder and ear opening in A. omiltemanus), a slightly larger dewlap in females, to 64 mm2 (versus to 41 mm2 in A. omiltemanus), the circumnasal usually in contact with the first supralabial (versus those scales separated by the presence of a subnasal in A. omiltemanus), and 4-6 internasal scales in the new species (versus usually 6-7 in A. omiltemanus). Furthermore, A. peucephilus differs from A. omiltemanus in hemipenial morphology (no finger-like processus on asulcate side in A. peucephilus versus such a processus present in A. omiltemanus). Also, in a preliminary molecular genetic analysis of the mitochondrial CO1 gene fragment, A. peucephilus has a genetic distance of 11.5% from A. omiltemanus. Anolis peucephilus was collected at night while the lizards were sleeping in pine trees, 2-10 m above the ground.

A new species of Bolitoglossa (Caudata, Plethodontidae) from the continental divide of western Panama.

We describe the new salamander species Bolitoglossa jugivagans from the Atlantic slopes of the Fortuna depression in western Panama on the basis of morphological and molecular data. Based on mtDNA data, the new species seems to be closely related to B. aureogularis and B. robinsoni, with which it forms a subclade within the subgenus Eladinea.

Distribution and variation of the giant alpha anoles (Squamata: Dactyloidae) of the genus Dactyloa in the highlands of western Panama, with the description of a new species formerly referred to as D. microtus.

Six species of giant alpha anoles of the genus Dactyloa are known to occur in western Panama: Dactyloa casildae, D. frenata, D. ibanezi, D. insignis, D. kunayalae, and D. microtus. Based on own material collected along the highlands in Bocas del Toro, Chiriquf, and Veraguas provinces and the Comarca Ngöbe-Buglé of western Panama, we review their variation in morphological characters and the 16S rRNA mitochondrial gene. Our results support all six nominal taxa, but reveal considerable genetic differentiation between populations of the two highland species, D. casildae and D. microtus, respectively, from different localities. Correlated morphological differences confirm the existence of a cryptic species among populations currently assigned to D. microtus, which we describe as Dactyloa ginaelisae sp. nov. We provide point distribution maps, morphology and color descriptions, photographs in life, conservation status assessments, and an identification key for all seven species.

Two new species of semiaquatic Anolis (Squamata: Dactyloidae) from Costa Rica.

We describe two new species of Draconura-clade semiaquatic anoles from the central Pacific versant of Costa Rica. The two new species are similar to Anolis aquaticus in external appearance and ecology but differ from this species in male dewlap coloration and scalation. Anolis robinsoni sp. nov. and A. riparius sp. nov. differ from each other mainly in male dewlap color. All three species are distinct according to diagnostic morphological traits and a phylogenetic analysis of mitochondrial DNA sequences (669 bases of COI gene). We discuss the distribution and ecology of Anolis aquaticus and the new species.

A taxonomic revision of Boiga multomaculata (Boie, 1827) and B. ochracea (Theobald, 1868), with the description of a new subspecies (Squamata, Serpentes, Colubridae).

The analyses of molecular genetic data (mtDNA markers 16S, ND4, CYTB, and the nuclear marker c-mos) provided evidence that the Asian cat snake taxa Boiga multomaculata and B. ochracea actually represent a single species. They form mixed clades of low intraclade genetic differentiation. This evidence for conspecificy is supported by the lack of differentiation in all examined pholidotic and morphometric characters. Therefore, we formally place Dipsas ochracea Theobald, 1868 in the synonymy of Dipsas multomaculata Boie, 1827. We provide a summary of the tangled taxonomic history of the taxa involved in this study. Also, we resurrect Dipsadomorphus stoliczkae Wall, 1909 from synonymy of B. ochracea, for specimens exhibiting 21 midbody dorsal scale rows. Boiga stoliczkae is found in the Himalayas north and west of the Brahmaputra valley. Finally, based on the detection of historical genetic lineages within the newly defined species Boiga multomaculata we recognize three subspecies: Boiga multomaculata multomaculata (Boie, 1827), Boiga multomaculata ochracea (Theobald, 1868), and Boiga multomaculata septentrionalis n. ssp. which is distributed in northern Myanmar and Assam and Nagaland, India. We designate BMNH 1946.1.2.60 (1) as neotype of Dipsas ochracea Theobald, (2) as lectotype of D. ochraceus Günther, and (3) as lectotype of Boiga ochracea walli Smith, thereby making these names objective synonyms. Finally, we designate BMNH 94.12.31.55 as lectotype of Dipsadomorphus stoliczkae Wall.

Distribution and identification of the species in the genus Helicops Wagler, 1830 (Serpentes, Colubridae, Xenodontinae).

Background: The aquatic snakes of the genus Helicops are widely distributed throughout northern South America, but understudied concerning some aspects, including morphological traits and distribution. The most recent publication that provided an identification key to all species of Helicops is over 50 years old. This key is of limited value today since it includes taxa no longer recognised and lacks 8 of the 19 species currently recognised. There never was a publication trying to summarise distributional and morphological information of all species of Helicops. Most knowledge of these species is distributed throughout many small publications, such as short observation notes. New information: Here, we present distribution maps (point records), an identification key and comments on identification for all species in this genus. We base our results on a comprehensive literature review of over 300 scientific publications and own examinations. Our examinations comprise 190 specimens of 10 of the 19 currently recognised species and one Helicops sp. We report range extensions for the species H.danieli, H.infrataeniatus, H.leopardinus, H.pastazae and H.polylepis.

A study on the genetic population structure and the tetrodotoxin content of rough-skinned newts, Taricha granulosa (Salamandridae), from their northern range of distribution.

Rough-skinned newts, Taricha granulosa, which contain tetrodotoxin (TTX), a potent neurotoxin, are widely distributed along the west-coast of North America up to British Columbia (BC), Canada, and Southeast Alaska. Their genetic population structure using DNA-microsatellites and the TTX-content of specimens from British Columbia (Prince Rupert area) and Alaska (Revillagigedo Island, Shelter Island, and Juneau) were analysed. TTX-concentrations were low in newts from BC and Revillagigedo Island, but high in specimens from mainland Juneau, which had been deliberately introduced from Shelter Island, where TTX was not detectable in the individuals sampled. No significant genetic differences were detected between these populations, which may correlate with the high intraspecies variability of TTX. It is still an open question, which factors favour or induce the toxin production in the newts.

A new genus and species of mud snake from Myanmar (Reptilia, Squamata, Homalopsidae).

Based on two male and two female individuals, we describe a new genus and species of mud snake, Myanophis thanlyinensis gen. nov., sp. nov., from the vicinity of the campus of East Yangon University, Yangon, Thanlyin, Myanmar. This species differs from every other homalopsid species by the following combination of characters: (1) dorsal scales smooth, row formula 21-21-19 or 21-21-17; (2) tail short, ratio tail length/SVL 0.185-0.204 in males, 0.160-0.167 in females; (3) nasal scales separated; (4) 125-126 ventral scales in males, 120-122 in females; (5) 38-39 subcaudal scales in males, 32-34 in females; and (6) hemipenis bilobed. Its matrilineal genealogy (based on analyses of 16S and cytochrome b sequences), associates Myanophis thanlyinensis gen. nov., sp. nov. most closely with species of the genera Myrrophis and Gyiophis. The new taxon differs from the species of Myrrophis and Gyiophis by having a bilobed hemipenis (vs. unilobed). Myanophis thanlyinensis gen. nov., sp. nov. differs further from the species of Myrrophis by having 125-126 ventral scales in males and 120-122 in females (vs. 137-162 and 137-164, respectively), and 38-39 subcaudal scales in males and 32-34 in females (vs. 39-55 and 37-52, respectively). Myanophis thanlyinensis gen. nov., sp. nov. differs further from the species of Gyiophis by lacking dark blotches along flank (vs. present), and by having 21 dorsal scales rows at midbody (vs. 25). We provide an identification key to the homalopsid species known to occur in Myanmar. As a novelty to the classic holotype description and characterization, the individual has been genome sequenced by Illumina short-read technology and its genome has been assembled into a draft nuclear genome and a complete, annotated mitochondrial genome. This innovative approach comprehensively and permanently characterizes the genomic variation of the holotype.

The earless monitor lizard Lanthanotus borneensis - A venomous animal?

Based on its mandibular gland secretion, the earless monitor lizard, Lanthanotus borneensis, has been considered a venomous animal like other members of the Toxicofera group, including Heloderma. In the present study, the gland structure and teeth of L. borneensis were examined by micro-tomography (µCT) and scanning electron microscopy (SEM), respectively, and proteomic analysis of the gland extract was performed. The mandibular gland consists of six compartments with separate ducts. The pleurodont teeth of the lower and upper jaw are not grooved but possess a sharp ridge on the anterior surface. Proteomic analysis of the gland extract confirmed previous studies that kallikrein enzymes are the major biologically active components. In view of the lizard's biology, its mandibular gland secretion is obviously not needed for prey capture or defence. It seems not justified the labelling of L. borneensis as a venomous animal. However, definitively answering this question requires toxinological studies on natural prey.

A new endemic species of Hemidactylus (Squamata: Gekkonidae) from São Nicolau Island, Cabo Verde.

A new species of gecko of the genus Hemidactylus (Squamata: Gekkonidae) is described from São Nicolau Island, Cabo Verde Archipelago, and the Sal and Boavista island populations of Hemidactylus boavistensis (i.e., Hemidactylus boavistensis boavistensis comb. nov. and Hemidactylus boavistensis chevalieri comb. nov.) are recognized as subspecies. Hemidactylus nicolauensis sp. nov. is genetically distinct from H. bouvieri, to which it has previously been referred, and from all other closely related endemic Hemidactylus from Cabo Verde Islands in mitochondrial (12S cyt b) and nuclear (RAG2, MC1R) markers. It is characterized morphologically by its distinct colouration and a diagnostically different arrangement of digital lamellae. With the description of this new species, São Nicolau is now known to harbour three single-island endemic gecko species, and the documented reptile diversity in Cabo Verde is raised to 23 endemic species. As a result of our taxonomic changes, existing conservation regulations should be updated and the conservation status of these taxa should be re-evaluated.

A new species of anole from Parque Nacional Volcán Arenal, Costa Rica (Reptilia, Squamata, Dactyloidae: Norops).

We describe the new species Norops arenal sp. nov. from Parque Nacional Volcán Arenal, north-central Costa Rica. In external morphology and genetic similarity of the 16S DNA barcode, Norops arenal is most similar to N. altae, N. fortunensis, N. fuscoauratus, N. gruuo, N. kemptoni, N. monteverde, N. pseudokemptoni, and N. tenorioensis. In morphology it shares with these species the following characteristics: (1) short hind limbs; (2) a single elongate prenasal scale; (3) tiny, smooth, often juxtaposed body scales; and (4) a slender habitus, often delicate. Norops arenal differs from these species, among several scalation details, by having a blackish central area in the male dewlap in life and in preservative (vs. no suffusion of black pigment on male dewlap in the other species), and a small red female dewlap in life (vs. dirty white, cream colored, or orange); extremely short hind legs with the tip of fourth toe of the adpressed hind leg reaching only to level of shoulder (vs. usually at least to level of ear in the other species); a short tail with a tail length/SVL ratio of 1.53 in single specimen with complete tail (vs. this ratio >1.6 in the other species); and a tiny size with 41.5 mm in single known adult male and 38.5 mm in single known adult female (vs. SVL of adults usually >42.0 mm). It further differs from N. altae, N. fuscoauratus, N. gruuo, N. pseudokemptoni, and N. tenorioensis by having a unilobed hemipenis (vs. bilobed in these five species).

Cryptic Diversity, but to What Extent? Discordance Between Single-Locus Species Delimitation Methods Within Mainland Anoles (Squamata: Dactyloidae) of Northern Central America.

Single-locus molecular barcoding is a useful method for identifying overlooked and undescribed biodiversity, providing the groundwork for further systematic study and taxonomic investigation. A variety of methods for delimiting species from barcoding libraries have been developed and applied, allowing for rapid estimates of species diversity in a broad range of taxa. However, tree-based and distance-based analyses can infer different group assignments, potentially over- or underestimating the number of putative species groups. Here, we explore diversity of mainland species of anole lizards from the Chortís Block biogeographical province of northern Central America using a DNA barcoding approach, generating and analyzing cytochrome oxidase subunit I (COI) sequences for over 400 samples assignable to 33 of 38 (86.8%) native and one introduced mainland species. We subsequently tested the effects different models of nucleotide substitution, different species-delimitation algorithms, and reducing our dataset had on species delimitation estimates. We performed of two distance-based (ABGD, RESL) and three tree-based (bPTP, mPTP, GMYC) analyses on both the full dataset and a dataset consisting only of unique halotypes. From 34 nominal taxa, analyses of the full dataset recovered between 34 and 64 operational taxonomic units (OTUs), while analyses of the reduced dataset inferred between 36 and 59. Reassigning individuals to either mPTP-inferred or ABGD clustered (7.2% threshold) groups improved the detection of a barcoding gap across three different models of nucleotide substitution, removing overlap between intra- and interspecific distances. Our results highlight the underestimated diversity of mainland Chortís Block anoles, but the lack of congruence between analyses demonstrates the importance of considering multiple analytical methods when dealing with single-locus datasets. We recommend future studies consider the effects of different models of nucleotide substitution on proposed barcoding gaps, as well as the effect reducing a dataset to unique haplotypes may have on proposed diversity estimates.

A global catalog of primary reptile type specimens.

We present information on primary type specimens for 13,282 species and subspecies of reptiles compiled in the Reptile Database, that is, holotypes, neotypes, lectotypes, and syntypes. These represent 99.4% of all 13,361 currently recognized taxa (11,050 species and 2311 subspecies). Type specimens of 653 taxa (4.9%) are either lost or not located, were never designated, or we did not find any information about them. 51 species are based on iconotypes. To map all types to physical collections we have consolidated all synonymous and ambiguous collection acronyms into an unambiguous list of 364 collections holding these primary types. The 10 largest collections possess more than 50% of all (primary) reptile types, the 36 largest collections possess more than 10,000 types and the largest 73 collections possess over 90% of all types. Of the 364 collections, 107 hold type specimens of only 1 species or subspecies. Dozens of types are still in private collections. In order to increase their utility, we recommend that the description of type specimens be supplemented with data from high-resolution images and CT-scans, and clear links to tissue samples and DNA sequence data (when available). We request members of the herpetological community provide us with any missing type information to complete the list.

Diversity of Tropidurus (Squamata: Tropiduridae) in Paraguay-an integrative taxonomic approach based on morphological and molecular genetic evidence.

Tropidurus is a Neotropical genus of iguanoid lizards characterized by a conspicuously enlarged interparietal plate, the presence of gular folds, presence of infradigital keels, and the absence of femoral pores. Currently, 29 species are recognized within the genus, seven of which are present in Paraguay: T. etheridgei, T. torquatus, T. guarani, T. lagunablanca, T. spinulosus, T. tarara, and T. teyumirim. We generated genetic data based on two DNA mitochondrial markers (16S and COI) and one nuclear (PRLR) marker for all the seven Paraguayan species with the goal to identify the taxonomic relationships among taxa based on the intra- and interspecific genetic variation and the construction of molecular clusters. ML and BI analyses match in the recognition of two main clusters: groups torquatus and spinulosus, and within the torquatus group the differentiation between T. catalanensis and T. etheridgei is highly supported. Nevertheless, there is a complete lack of congruence between mitochondrial and nuclear genes in the topology within the spinulosus group. Tropidurus guarani and T. spinulosus are more differentiated from the remaining species of the spinulosus group with genetic p-distances from 4.0 to 6.0. Low distances were found between T. lagunablanca and T. tarara (1.0-1.1%), and slightly higher, among T. teyumirim, T. lagunablanca, and T. tarara (2.0-2.6% respectively). From a morphological perspective, species of the Tropidurus torquatus group are easily distinguished; but we found strong overlaps of scalation characters in the spinulosus group. We interpret the low genetic distances documented among the nominal taxa Tropidurus lagunablanca, T. tarara, and T. teyumirim as evidence for conspecificity. This hypothesis is supported by the lack of morphological characters that would diagnose any of the three taxa. Similarly, we found low genetic distances among populations assigned to the nominal taxa T. guarani and T. spinulosus, including samples from near the type locality of the former, and therefore we recognize only two species of the T. spinulosus complex in Paraguay: T. spinulosus and T. lagunablanca.

The Only Known Jawed Vertebrate with Four Eyes and the Bauplan of the Pineal Complex.

The pineal and parapineal organs are dorsal outpocketings of the vertebrate diencephalon that play key roles in orientation and in circadian and annual cycles. Lampreys are four eyed in that both the pineal and parapineal form eyelike photosensory structures, but the pineal is the dominant or sole median photosensory structure in most lower vertebrate clades. The pineal complex has been thought to evolve in a single direction by losing photosensory and augmenting secretory function in the transitions from three-eyed lower vertebrates to two-eyed mammals and archosaurs [1-3]. Yet the widely accepted elaboration of the parapineal instead of the pineal as the primary median photosensory organ [4] in Lepidosauria (lizards, snakes, and tuataras) hints at a more complex evolutionary history. Here we present evidence that a fourth eye re-evolved from the pineal organ at least once within vertebrates, specifically in an extinct monitor lizard, Saniwa ensidens, in which pineal and parapineal eyes were present simultaneously. The tandem midline location of these structures confirms in a striking fashion the proposed homology of the parietal eye with the parapineal organ and refutes the classical model of pineal bilaterality. It furthermore raises questions about the evolution and functional interpretation of the median photosensory organ in other tetrapod clades.

Lack of alkaloids and tetrodotoxin in the neotropical frogs Allobates spp. (Aromobatidae) and Silverstoneia flotator (Dendrobatidae).

Frogs of the family Dendrobatidae are known to contain toxic alkaloids in their skin secretion, but Allobates species of the closely related Aromobatidae family are considered to lack toxic secretions. However, contradictory results have been reported. Analyses of alcohol extracts from three different Allobates species from South-America (Guiana Shield), Central America (Costa Rica), and from the dendrobatid frog Silverstoneia flotator confirm the absence of alkaloids and tetrodotoxin in aromobatids and in a dendrobatid of the subfamily Colostethinae.