Hemipenial Morphology Research Articles

First Report of Hemiclitores in Females of South American Liolaemid Lizards

Abstract Studies about copulatory organs in Squamata were restricted to the morphology of hemipenes until Bohme reported homologous paired structures in females of a species of Varanus, which he called hemiclitores. We report the presence of hemiclitores in females of Phymaturus and of two species of Liolaemus and describe observations on the interspecific variation in hemipenis morphology in Phymaturus. Phymaturus and Liolaemus belong to Liolaemidae, a species-rich family of lizards; research about hemipenis morphology is scarce and limited to a few species of those genera. We found the retractor clitoridis magnus in all of the species analyzed; however, the transverse penis was not present in all of them. The general structure of hemiclitores of Phymaturus and Liolaemus females resembled that described for other squamate species; they were smaller than hemipenes and exhibited a sulcus spermaticus. The variation found in different features of these organs (shape, size, pigmentation), as well as the gener...

Anolis marsupialis Taylor 1956, a valid species from southern Pacific Costa Rica (Reptilia, Squamata, Dactyloidae).

The examination of the holotype of Anolis marsupialis Taylor 1956 along with recently collected specimens reveals that A. marsupialis is a valid species. It differs from its closest congeners A. humilis Peters 1863 and A. quaggulus Cope 1885, in male dewlap coloration, scalation, body size, and hemipenial morphology. These findings are supported by preliminary molecular genetic analysis.

A revision of the Mexican Anolis (Reptilia, Squamata, Dactyloidae) from the Pacific versant west of the Isthmus de Tehuantepec in the states of Oaxaca, Guerrero, and Puebla, with the description of six new species.

We revise the species of anoles occurring along the Pacific versant of Mexico west of the Isthmus de Tehuantepec in the states of Oaxaca, Guerrero, and Puebla. Based on our analyses of morphological and molecular genetic data, we recognize 21 species, six of which we describe as new (i.e., Anolis carlliebi sp. nov., A. immaculogularis sp. nov., A. nietoi sp. nov., A. sacamecatensis sp. nov., A. stevepoei sp. nov., and A. zapotecorum sp. nov.). Furthermore, we synonymize Anolis forbesi Smith & Van Gelder 1955 with Anolis microlepidotus Davis 1954. Of the recognized species, six have smooth ventral scales (i.e., Anolis dunni, A. gadovii, A. liogaster, A. omiltemanus, A. peucephilus, and A. taylori) and 14 have keeled ventral scales (i.e., A. boulengerianus, A. carlliebi, A. immaculogularis, A. megapholidotus, A. microlepidotus, A. nebuloides, A. nebulosus, A. nietoi, A. quercorum, A. sacamecatensis, A. stevepoei, A. subocularis, A. unilobatus, and A. zapotecorum). In one species, A. macrinii, the ventral scales vary from smooth to weakly keeled. For each species we provide color descriptions in life, color photographs in life, descriptions and illustration of hemipenis morphology (if available), description of external morphology, distribution maps based on the specimensexamined, comments on the conservation status, and natural history notes. Finally, we provide a dichotomous key for the identification of the 21 species of anoles occurring along the Pacific versant of Mexico west of the Isthmus de Tehuantepec in the states of Oaxaca, Guerrero, and Puebla.

Systematics of the Neotropical SnakeDendrophidion paucicarinatum(COPE), with a Description of its Hemipenis (Serpentes: Colubridae)

Abstract We present the first modern comprehensive systematic treatment of the Central American snake Dendrophidion paucicarinatum (Cope). We redescribe the species on the basis of a large sample from throughout its range, redescribe and illustrate the holotype, and provide a detailed treatment of its hemipenial morphology. Dendrophidion paucicarinatum is distinguished from other species of the genus by a combination of having more ventrals (>175), fewer subcaudals (<140), very weak keels on the dorsal scales, and distinct transverse dark lines across each ventral in adults. It is an upland species found almost exclusively above 1,000 m in mountainous regions of Costa Rica and western Panama, primarily in premontane and montane humid to wet forests and cloud forests. It is a large species attaining a total length of 1.5 m and is diurnal and anurophagous like other species of Dendrophidion. Known clutch sizes are 3–12. Evidence from the Monteverde region in Costa Rica suggests that populations of D. paucic...

Reclassification of Oligodon ningshaanensis Yuan, 1983 (Ophidia: Colubridae) into a New Genus, Stichophanes gen. nov. with Description on Its Malacophagous Behavior

The complete mitochondrial cytb gene and the partial nuclear c-mos gene of Oligodon ningshaanensis Yuan, 1983 were sequenced and used for reconstructing the phylogenetic relationship of this taxon amongst alethinophidian snakes. Its strong affinity to the New World subfamily Dipsadinae and the Old World species Thermophis baileyi was inferred. Hemipenial morphology found by authors conflicts with the original description and its similarity with those of the dipsadid snakes is in accordance with our molecular results. Feeding tests show that O. ningshaanensis is a malacophagous predator, which is another matchless character for this species. This behavior is described and compared with other known slug- and snail-feeding snakes. The discovery of the particular position of our subject indicates that erecting a new genus is necessary accommodate this unique species.

Hemipenial Morphology in the Semifossorial Snakes of the GenusNiniaand a New Species from Trinidad, West Indies (Serpentes: Dipsadidae)

The hemipenial morphology of nine species in Ninia is described for the first time. A total of 65 hemipenes of all recognized species of Ninia were examined. As a result of the evaluation of the hemipenial and caudal morphology, Ninia diademata labiosa is elevated to full species status and a new species is described from 240 m above sea level in the Northern Range of Trinidad. The new species exhibits several diagnostic features, including (1) a primary temporal scale that is extremely long and contacts the upper labial scales 5–7; (2) a temporal scale formula 1 2, with the primary temporal scale three times longer than the lower posterior temporal; and (3) forked hemipenes. The male genitalia of Ninia are excellent indicators of species limits and an effective tool to clarify the identity of cryptic species in the genus.

Systematics of the blindsnakes (Serpentes: Scolecophidia: Typhlopoidea) based on molecular and morphological evidence.

The blindsnake superfamily Typhlopoidea (Gerrhopilidae, Typhlopidae, and Xenotyphlopidae) is a diverse, widespread part of the global snake fauna. A recent systematic revision based on molecular phylogenetic analyses and some morphological evidence presented a preliminary solution to the non-monophyly of many previously recognized genera, but additional clarification is needed regarding the recognition of some species and genera. We rectify these problems here with a new molecular phylogenetic analysis including 95 of the 275 currently recognized, extant typhlopoids, incorporating both nuclear and mitochondrial loci. We supplement this with data on the external, visceral, and hemipenial morphology of nearly all species to generate a revised classification for Typhlopoidea. Based on morphological data, we re-assign Cathetorhinus from Typhlopidae to Gerrhopilidae. Xenotyphlopidae maintains its current contents (Xenotyphlops). In Typhlopidae, one monotypic genus is synonymized with its larger sister-group as it cannot be unambiguously diagnosed morphologically (Sundatyphlops with Anilios), and two genera are synonymizedwith Typhlops (Antillotyphlops and Cubatyphlops), as they are not reciprocally monophyletic. The genus Asiatyphylops is renamed Argyrophis, the senior synonym for the group. We erect one new genus (Lemuriatyphlops) for a phylogenetically distinct species-group in Asiatyphlopinae. Fourteen of eighteen recognized typhlopid genera are maintained in four subfamilies: Afrotyphlopinae (Afrotyphlops, Grypotyphlops [re-assigned from Asiatyphlopinae], Letheobia, and Rhinotyphlops), Asiatyphlopinae (Acutotyphlops, Anilios, Cyclotyphlops, Indotyphlops, Malayotyphlops, Ramphotyphlops, and Xerotyphlops), Madatyphlopinae (Madatyphlops), and Typhlopinae (Amerotyphlops and Typhlops), some with altered contents. Diagnoses based on morphology are provided for all 19 typhlopoid genera, accounting for all 275 species. This taxonomy provides a robust platform for future revisions and description of new species.

A new species of triadal coral snake of the genus Micrurus Wagler, 1824 (Serpentes: Elapidae) from northeastern Brazil.

The genus Micrurus comprises 123 currently recognized taxa (species and subspecies) that are traditionally arranged in four species groups diagnosable mainly by color pattern characteristics. Here, we describe a new species of triadal coral snake from northeastern Brazil. The new species is distinguished from other sympatric triadal congeners (M. lemniscatus carvalhoi, M. ibiboboca and M. brasiliensis) mainly by the entirely black parietals and by a suite of external characters and hemipenial morphology. The new species appears to be restricted to tropical ombrophilous lowland coastal forests of northeastern Brazil and all recently collected specimens are known to occur in small forest patches surrounded by periurban environment, which calls for an urgent evaluation on its conservation status.

A new Andean species of Philodryas (Dipsadidae, Xenodontinae) from Ecuador.

We describe a new species of Philodryas from the highlands of southern Ecuador. The new species is distinguished from all known species of Philodryas by a unique combination of coloration, scalation, and hemipenial characters. The new species resembles Philodryas simonsii in color pattern. However, they differ notoriously by their hemipenial morphology. The three other trans-Andean members of the genus (Philodryas simonsii, Philodryas chamissonis, and Philodryas tachymenoides), along with the new species, compose a probably monophyletic group that may be characterized by the presence of ungrooved postdiastemal teeth in the maxilla. Unlike most species of the genus Philodryas, the new species shows a restricted distribution, being apparently endemic to a small region of high-altitude (3150-4450m) grasslands in the southern Andes of Ecuador.

Are Hemipenial Spines Related to Limb Reduction? A Spiny Discussion Focused on Gymnophthalmid Lizards (Squamata: Gymnophthalmidae)

Calcified spines in the hemipenial surface occur convergently in several gymnophthalmid lizard species and in advanced snakes. Based on the pronounced degrees of limb reduction in these distantly related lineages, such hemipenial structures were suggested to play a functional role in couple-anchoring during copulation, partly assuming the function of the limbs during mating. Herein, we assessed the hemipenial morphology of virtually all the valid genera of the family Gymnophthalmidae to test for a phylogenetic correlation between limb reduction and the presence of calcified hemipenial spines. The occurrence of calcified structures was mapped on the two most comprehensive phylogenies of the family. We concluded that spiny hemipenes are by no means necessarily associated with reduction of limbs. Conversely, the presence of well-developed hemipenial spines in specific limb-reduced taxa does not allow one to disregard the possibility that in some instances such structures might indeed be functionally associated with couple-anchoring, improving the success of mating.

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.

Description of the hemipenial morphology of Tupinambis quadrilineatus Manzani and Abe, 1997 (Squamata, Teiidae) and new records from Piauí, Brazil

Few data are available on the morphology of the hemipenis of teiid lizards, especially those of the recently-defined genus Tupinambis, a widely-distributed group of large-bodied lizards. This study provides an illustrated description of the hemipenis of Tupinambis quadrilineatus, which is similar to that of other representatives of the Tupinambinae subfamily. New records of the species from the state of Piauí, in northeastern Brazil, are also presented.

A Contribution to the Knowledge ofAnolis macriniiSmith, 1968 (Reptilia: Squamata: Dactyloidae)

Abstract During three short visits to the coffee-growing region in the hills north of Pochutla (Oaxaca, Mexico), we observed Anolis macrinii in its natural habitat. The species appeared to be relatively abundant, and we collected 12 individuals, including several adult males. The holotype of this species was reported erroneously to be an adult male but actually is a female. The confusion might have arisen from the moderate-sized dewlap present in adult females. However, males have a very large dewlap and a pair of moderately to greatly enlarged postcloacal scales. We provide color descriptions in life for three individuals, color photographs in life, description and illustration of hemipenis morphology, and some natural history notes. Finally, we discuss the conservation status of this species.

Rediscovery of the earless microteiid lizard Anotosaura collaris Amaral, 1933 (Squamata: Gymnophthalmidae): a redescription complemented by osteological, hemipenial, molecular, karyological, physiological and ecological data.

More than a century after its discovery by Ernest Garbe, and almost 80 years after its original description, we obtained a series of specimens of the earless gymnophthalmid Anotosaura collaris, the type species of the genus, up to now known only by a single specimen. On the basis of the material obtained at and close to the type locality we redescribe the species, adding information about the external and hemipenial morphology, osteology and karytoype. Molecular data confirm its sister relationship with Anotosaura vanzolinia as well as the close relationship of Anotosaura with the Ecpleopodini Colobosauroides and Dryadosaura. We supplement this information with thermophysiological, ecogeographical, karyotypic and ecological data.

Taxonomic revision of Drymoluber Amaral, 1930 (Serpentes: Colubridae).

The present study is a taxonomic revision of the genus Drymoluber Amaral, 1930, using meristic and morphometric characters, aspects of external hemipenial morphology and body coloration. Sexual dimorphism occurs in D. dichrous and D. brazili but was not detected in D. apurimacensis. Morphological variation within D. dichrous is related to geographic distance between populations. Furthermore, variation in the number of ventrals and subcaudals in D. dichrous and D. brazili follows latitudinal and longitudinal clinal patterns. Drymoluber dichrous is diagnosed by the presence of 15-15-15 smooth dorsal scale rows with two apical pits, and 157-180 ventrals and 86-110 subcaudals; it occurs along the eastern versant of the Andes, in the Amazon forest, on the Guiana Shield, in the Atlantic forest, and its transitional areas with the Caatinga and Cerrado. Drymoluber brazili has 17-17-15 smooth dorsal scale rows with two apical pits, 182-202 ventrals and 109-127 subcaudals, and ranges throughout the Caatinga, Cerrado, Atlantic forest and transitional areas between these last two domains. Drymoluber apurimacensis has 13-13-13 smooth dorsal scale rows without apical pits, 158-182 ventrals and 84-93 subcaudals, and occurs in the Apurimac Valley, south of the Apurimac and Pampas rivers in Peru.

A new species of Kukri Snake (Oligodon Fitzinger, 1826; Squamata: Colubridae) from the Cat Tien National Park, southern Vietnam.

We describe a new species of the genus Oligodon from the lowland forests of Cat Tien National Park, Dong Nai Province, in southern Vietnam. Oligodon cattienensis sp. nov. is distinguished from the remaining Southeast Asian kukri snakes by the combination of the following characters: medium-sized, deeply forked hemipenes without spines, 17-17-15 dorsal scale rows, nasal entire, 2 small postoculars, almost equal in size, 167-178 ventrals, 31-35 subcaudals, 24-35 + 5 large dark-edged vertebral blotches in combination with a yellow-orange or red vertebral stripe between blotches, head pattern including ocular band, temporal bands and elongated chevron, ventrals pink or whitish (reddish in juveniles) in life, some bearing a quadrangular dark blotch on each lateral side, or ventrals being entirely dark. Based on the hemipenial morphology the new species is assigned to the Oligodon cyclurus species group. A comparison table for all Indochinese Oligodon is provided.

The genus Cnemidophorus (Squamata: Teiidae) in State of Piauí, northeastern Brazil, with description of a new species.

The state of Piauí, northeastern Brazil, is covered mainly by 'cerrado' (33% of the total area of the state) and 'caatinga' (37%) vegetations, with another 19% occupied by ecotonal zones. Piauí lies within the known distribution of the Cnemidophorus ocellifer species group, currently composed of ten species. Until recently only C. ocellifer (Spix, 1825), a name recognized to cover several species, was recorded from Piauí. Currently two other species are registered from Piauí: C. confusionibus Arias, Carvalho, Rodrigues and Zaher 2011 and C. venetacaudus Arias, Carvalho, Rodrigues and Zaher 2011. We analyzed the external and hemipenial morphology of specimens from different localities in the state and our results indicate the presence of at least four species: C.cf. ocellifer, C. confusionibus, C. venetacaudus and C. pyrrhogularis sp. nov. Hemipenial analysis corroborates the existence of two subgroups within C. ocellifer species group. Cnemidophorus pyrrhogularis sp. nov., a member of the ocellifer subgroup, occurs in the northern part of Piauí, in ecotonal areas, and differs from the remaining species of the genus mainly on hemipenial morphology, number of femoral pores, number the lamellae under fourth finger and fourth toe, and color pattern. Diagnoses and hemipenis description of the four taxa are presented, as well as a detailed description of the new species.

A new species of Atractus (Serpentes: Dipsadidae) from Serra do Cipó, Espinhaço Range, Southeastern Brazil, with proposition of a new species group to the genus

We describe a new species of Atractus from Serra do Cipó, at the southeastern versant of Serra do Espinhaço, an inland mountain range nearly parallel to the Brazilian Atlantic coast. The new species is morphologically similar to A. paraguayensis and A. potschi with which it shares: 15 dorsal scale rows; white occipital region in juvenile specimens; dorsal ground color reddish pink to red (in life) with alternate black transversal spots, blotches or transversal bands; seven upper and lower labial scales; ventrals and subcaudals creamish white; hemipenis slightly bilobed, semicapitate and semicalyculate. The new species differs from both taxa mainly on the basis of exclusive coloration characters (number and shape of transversal bands) and hemipenial morphology (median face of the lobes without ornamentation), besides punctual meristic (number of ventral scales and maxillary teeth) and morphometric (adult snout-vent length) differences. Finally, we provide a discussion regarding the potential affinities of the new species and we redefine some Atractus species groups in order to better accommodate species sharing unique combinations of morphological characters.

A new species of gymnophthalmid lizard of the genus Anadia (Gymnophthalmidae: Cercosaurinae) from Northern Andes of Colombia

A new Colombian species of gymnophthalmid lizard is described from Porce River piedmont, in the northern region of the Cordillera Central of Colombia. The new species is allocated in the Anadia bitaeniata group and represents the westernmost member of the group. This species is distinguished primarily by the sub-hexagonal dorsal scales, large quadrangular ventral scales, ten femoral pores per leg not separated by inter-pore scales in both sexes, and by a combination of several characteristic scale counts, hemipenial morphology and color pattern. The threatened status of the new species is discussed.

Taxonomy of the South American Dwarf Boas of the Genus Tropidophis Bibron, 1840, With the Description of Two New Species from the Atlantic Forest (Serpentes: Tropidophiidae)

A taxonomic study on the South American dwarf boas of the genus Tropidophis revealed the existence of two new species in the Atlantic Forest biome. As a result, we recognize five mainland species, three in the Atlantic Forest and two in northwestern South America. Based on general distribution and morphological orientation, the type locality of T. paucisquamis is restricted to Estação Biológica de Boracéia (EBB), municipality of Salesópolis, state of São Paulo, Brazil; furthermore, a lectotype for T. taczanowskyi is designated. We provide data on the hemipenial morphology of two South American Tropidophis, showing that the quadrifurcate condition described for West Indian taxa also occurs in mainland congeners. The distributions of the three Atlantic Forest species are congruent with patterns of diversification of other vertebrate taxa associated with cold climates prevalent at high elevations. Refugial isolation and riverine barriers may account for such speciation events.