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A new species of Riama Gray, 1858 (Squamata: Gymnophthalmidae) from the Tropical Andes
Abstract and Figures
A new species of Riama lizard from the western slopes of the Andes in northern Ecuador is described herein. Morpholog-ically, Riama yumborum sp. nov. can be distinguished from all other congenerics by having an incomplete nasoloreal su-ture and a cylindrical hemipenial body with diagonally orientated flounces on its lateral aspect. Phylogenetic analyses of mitochondrial and nuclear DNA support the monophyly of the new species and its sister taxon relationship with R. labi-onis, which occurs allopatrically.
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... obs.; present study). Although the molecular studies by Castoe et al. (2004; see also , Aguirre-Peñafiel et al. (2014) and advanced understanding of the phylogeny of Riama, the current delimitation of this genus is suspect. As recognized by , Castoe et al. (2004) sampled five of the 24 species of Riama only. ...
... Subsequently, many new species were referred to Proctoporus s.s. Goicoechea et al., 2013;Mamani et al., 2015), Petracola Echevarr ıa and Venegas, 2015) and Riama (Rivas et al., 2005;Arredondo and S anchez-Pacheco, 2010;S anchez-Pacheco, 2010a;S anchez-Pacheco et al., 2011S anchez-Pacheco et al., , 2012Aguirre-Peñafiel et al., 2014). In their phylogenetic study of Proctoporus s.s., Goicoechea et al. (2012) transferred into Proctoporus two species of Euspondylus Tschudi, 1845, a genus whose monophyly had also been questioned , and the monotypic genus Opipeuter Uzzell, 1969, which have prefrontals. ...
... Novel sequences were deposited in Gen-Bank. Our own data were augmented with sequences in GenBank from Fu (2000), Pellegrino et al. (2001), Castoe et al. (2004), Goicoechea et al. (2012), Kok et al. (2012), Torres-Carvajal and Mafla-Endara (2013), Aguirre-Peñafiel et al. (2014), Kok (2015) and . Voucher specimens and GenBank accession numbers are listed in Table 2. DNA isolation, sequencing, and editing. ...
Riama is the most speciose genus of the Neotropical lizard family Gymnophthalmidae. Its more than 30 montane species occur throughout the northern Andes, the Cordillera de la Costa (CC) in Venezuela, and Trinidad. We present the most comprehensive phylogenetic analysis of Riama to date based on a total evidence (TE) approach and direct optimization of molecular and morphological evidence. Analyses use DNA sequences from four loci and 35 phenotypic characters. The dataset consists of 55 ingroup terminals representing 25 of the 30 currently recognized species of Riama plus five undescribed taxa, including an endemic species from the Sierra Nevada de Santa Marta (SNSM) in Colombia, and 66 outgroup terminals of 47 species. Analysis results in a well supported hypothesis in which Riama is polyphyletic, with its species falling into three clades. The Tepuian Anadia mcdiarmidi nests within one clade of Riama, and the recently resurrected Pantodactylus nests within Cercosaura. Accordingly, we propose a monophyletic taxonomy that reflects historical relationships. Analysis of character evolution indicates that the presence/absence of prefrontals—a cornerstone of the early genus-level taxonomy of cercosaurines—is optimally explained as having been plesiomorphically present in the most recent common ancestor of Cercosaurinae and lost in that of the immediately less inclusive clade. Multiple independent reversals to present and subsequent returns to absent occur within this clade. To evaluate the impact of phenotypic evidence on our results, we compare our TE results with results obtained from analyses using only molecular data. Although phenotypic evidence comprises only 1.2% of the TE matrix, its inclusion alters both the topology and support values of the clades that do not differ. Finally, current phylogenetic evidence reveals a SNSM–CC–Trinidad–tepuis biogeographical link. We hypothesize that an ancient connection facilitated the exchange of species between the SNSM and the CC.
... Lepidoblepharis conolepis Avila-Pires, 2001 is a moderately large Lepidoblepharis species that is endemic to Ecuador. This species is only known from a few localities on the western slopes of Cotopaxi and Pichincha provinces (Avila-Pires 2001;Yánez-Muñoz et al. 2009;Aguirre-Peñafiel et al. 2014;Torres-Carvajal and Lobos 2014;Torres-Carvajal et al. 2015). We report a range extension for L. conolepis to the province of Carchi in northern Ecuador. ...
... The specimen shows a dorsal pattern of dark olive with irregular dark brown marks and light spots ( Figure 2), in contrast to the previous records that report a uniform dark brown dorsum with light or turquoise spots (Avila-Pires 2001;Yánez-Muñoz et al. 2009;Torres-Carvajal et al. 2015). Lepidoblepharis conolepis was known to occur only at six localities in Pichincha province and two localities in Cotopaxi province (Table 1; Avila-Pires 2001;Yánez-Muñoz et al. 2009;Aguirre-Peñafiel et al. 2014;Torres-Carvajal and Lobos 2014;Torres-Carvajal et al. 2015). Because of this restricted distribution, the species is recognized as Endangered (EN) under criteria B1ab(i,iii,iv) (Carrillo et al. 2005;IUCN 2012); the minimum convex polygon (MCP) encompassing all previous sites is 1,732 km 2 . ...
We document Lepidoblepharis conolepis from El Cielito, Carchi province, northern Ecuador. This record , the first record for Carchi province, represents a northward range extension of 90 km from the previously known occurences. All known occurrences of L. conolepis, including our new record, are restricted to the western slopes of the Ecuadorian Andes at sites within fragmented natural forest habitats.
... En el transcurso de la última década, la taxonomía, diversidad críptica y relaciones filogenéticas de los saurios neotropicales de la familia Gymnophthalmidae, han sido activamente estudiadas, con una tasa de descripción promedio de cuatro especies nuevas por año [e.g. [1][2][3][4][5][6]). Varios estudios han provocado un cambio en el entendimiento de las relaciones filogenéticas entre los géneros que componen este clado, con la definición de subfamilias, cambios en los géneros y análisis biogeográficos [7][8][9][10][11][12]. ...
We performed a systematic review of the genus Anadia in Ecuador to delimit species based on several lines of evidence: external morphology, hemipenes, coloration, DNA sequences and geographical distribution. We describe a new species, Anadia buenaventura sp. nov., which is distinguished from other species of Anadia by presenting a pattern of coloration with light dorsolateral bands and paravertebral rows of dark spots, as well as fewer scales around the body. We also present new data on the distribution of Anadia petersi Oftedal and a molecular phylogeny that suggests that Anadia rhombifera (Günther) is a complex of species that requires further taxonomic work that should include material from Colombia.
... Moreover, basically every new phylogenetic study published to date brought evidence for cryptic species being present (Goicoechea et al. 2012;Torres-Carvajal et al. 2015Sánchez-Pacheco et al. 2017b). The amount of overall cryptic diversity within all cercosaurines is not straightforward to gauge because most previous phylogenies had a narrow focus on particular genera Goicoechea et al. 2012;Torres-Carvajal and Mafla-Endara 2013;Aguirre-Peñafiel et al. 2014) or regions (Kok et al. 2012), or had species represented by a single sample (Pellegrino et al. 2001b;Kok 2015). The most comprehensive phylogenetic reconstructions of higher clades were published recently by Torres-Carvajal et al. (2015, Goicoechea et al. (2016), and Sánchez-Pacheco et al. (2017b). ...
Cercosaurine lizards (subfamily Cercosaurinae of the family Gymnophthalmidae) represent a substantial component of the reptile fauna in the Neotropics. Several attempts have been made to reconstruct the phylogenetic relationships within this group, but most studies focused on particular genera or regions and did not cover the subfamily as a whole. In this study, material from the montane forests of Peru was newly sequenced. In combination with all cercosaurine sequences available on GenBank, an updated phylogeny of Cercosaurinae is provided. Monophyly was not supported for three of the currently recognised genera ( Echinosaura , Oreosaurus , and Proctoporus ). The genus Proctoporus is formed by five monophyletic groups, which should be used in future taxonomic revisions as feasible entities. Recognition of two previously identified undescribed clades (Unnamed clades 2 and 3) was supported and yet another undescribed clade (termed here Unnamed clade 4), which deserves recognition as an independent genus, was identified herein. Selvasaurabrava , a new genus and new species of arboreal gymnophthalmid lizard is described from the montane forests of the Pui Pui Protected Forest, Provincia de Chanchamayo, Región Junín, Peru. The new species is characterised by its small size (SVL 42.1–45.9 mm), slender body, smooth head shields, presence of paired prefrontal shields, fused anteriormost supraocular and anteriormost superciliary shields, transparent not divided lower palpebral disc, slightly rugose subimbricate rectangular dorsal scales in adults (slightly keeled in juveniles), distinctly smaller but non-granular lateral scales, smooth squared to rectangular ventral scales, and hemipenial lobes large, distinct from the hemipenial body. Phylogenetic affinities of the new genus to the other cercosaurine genera, as well as basal phylogenetic relationships between the other cercosaurine genera remain unresolved.
... PALABRAS CLAVE: Lagarto criptozoico, Microteiido, Andes venezolanos, estado Trujillo. Doan and Castoe (2005) retrieved the name Riama Gray, 1858, for a group of semifossorial/cryptozoic microteiids distributed in the central Andes of Peru, Ecuador, Colombia, Venezuela (Trinidad and Tobago and Cordillera de Mérida) (Sánchez-Pacheco 2010, , Aguirre-Peñafiel et al. 2014. The name of Proctoporus Tschudi, 1845 was limited to the central and southern regions of Peru, Bolivia and northern Argentina (see Goicoechea et al. 2013, Mamani et al. 2015. ...
The presence of microteiid Oreosaurus cephalolineatus is documented for the first time in the lacustrine slope of the Cordillera de Mérida, Venezuela. In addition, the collected material has allowed the relocation of the species within the genus (previously "Proctoporus"), which had already been suggested by other authors. In the other hand, this assumption suggests a review of other Cis-Andean microteiid "Riama inanis = Oreosaurus inanis", whose geographical distribution and morphological characteristics are very similar to O. cephalolineatus.
... As currently defined , Riama is composed of 31 species that occur throughout the Andes of central Peru, Ecuador, Colombia, Venezuela, the Cordillera de la Costa of Venezuela and Trinidad and Tobago (Aguirre-Peñafiel et al., 2014). Here we show that this definition does not reflect phylogeny, because ''Riama" laudahnae is nested within ''Proctoporus" as sister to P. ''bolivianus" (Fig. 2). ...
Gymnophthalmids are a highly diverse group of Neotropical lizards and its species richness is still in process of discovery. The incorporation of molecular evidence and a noticeable increase in taxon and geographic sampling in systematic studies has led to the description of numerous new genera and species of gymnophthalmids (particularly in Cercosaurinae) in recent years. Herein we describe a new genus and species of cercosaurine lizard with crocodile-like morphology, from the Venezuelan Guiana Shield on the basis of molecular phylogenetic and morphological evidence. Kataphraktosaurus gen. nov. can be readily distinguished from all other genera of Cercosaurinae by a unique combination of morphological characters that includes heterogeneous dorsal scalation with enlarged and strongly keeled scales forming two paravertebral rows, ventral and subcaudal scales imbricated and strongly keeled, large and symmetrical cephalic scales, absence of postmental scale, palpebral disc translucent and divided, tail slightly compressed, all digits clawed, and only six femoral pores (three at each hindlimb) inserted in a clump of small scales. This genus is described as monotypic and only contains Kataphraktosaurus ungerhamiltoni sp. nov., which is known from one specimen and diagnosed by the same set of aforementioned characters. The secretive habits of this species and the remoteness of the locality may explain its singleton situation. Following the International Union for Conservation of Nature's criteria, we categorized the new species as Data Deficient.
Tissue sample databases housed in biodiversity archives represent a vast trove of genetic resources, and these tissues are often destructively subsampled and provided to researchers for DNA extractions and subsequent sequencing. While obtaining a sufficient quantity of DNA for downstream applications is vital for these researchers, it is also important to preserve tissue resources for future use given that the original material is destructively and consumptively sampled with each use. It is therefore necessary to develop standardized tissue subsampling and loaning procedures to ensure that tissues are being used efficiently. In this study, we specifically focus on the efficiency of DNA extraction methods by using anuran liver and muscle tissues maintained at a biodiversity archive. We conducted a series of experiments to test whether current practices involving coarse visual assessments of tissue size are effective, how tissue mass correlates with DNA yield and concentration, and whether the amount of DNA recovered is correlated with sample age. We found that tissue samples between 2 and 8 mg resulted in the most efficient extractions, with tissues at the lower end of this range providing more DNA per unit mass and tissues at the higher end of this range providing more total DNA. Additionally, we found no correlation between tissue age and DNA yield. Because we find that even very small tissue subsamples tend to yield far more DNA than is required by researchers for modern sequencing applications (including whole genome shotgun sequencing), we recommend that biodiversity archives consider dramatically improving sustainable use of their archived material by providing researchers with set quantities of extracted DNA rather than with the subsampled tissues themselves.
Tissue sample databases housed in biodiversity archives represent a vast trove of genetic resources, and these tissues are frequently destructively subsampled and provided to researchers for DNA extractions and subsequent sequencing. While obtaining a sufficient quantity of DNA for downstream applications is vital for these researchers, it is also important to preserve tissue resources for future use given that the original material is destructively and consumptively sampled with each use. It is therefore necessary to develop standardized tissue subsampling and loaning procedures to ensure that tissues are being used efficiently. In this study, we specifically focus on the efficiency of DNA extraction methods by using anuran liver and muscle tissues maintained at a biodiversity archive. We conducted a series of experiments to test whether current practices involving coarse visual assessments of tissue size are effective, how tissue mass correlates with DNA yield and concentration, and whether the amount of DNA recovered is correlated with sample age. We found that tissue samples between 2 mg and 8 mg resulted in the most efficient extractions, with tissues at the lower end of this range providing more DNA per unit mass and tissues at the higher end of this range providing more total DNA. Additionally, we found no correlation between tissue age and DNA yield. Because we find that even very small tissue subsamples tend to yield far more DNA than is required by researchers for modern sequencing applications (including whole genome shotgun sequencing), we recommend that biodiversity archives consider dramatically improving sustainable use of their archived material by providing researchers with set quantities of extracted DNA rather than with the subsampled tissues themselves.
Supplementary phylogenetic trees and table of analysed samples
Una nueva especie endémica de lagarto gymnophthalmido del género Riama es descrita para el norte de la Cordillera Occidental en Colombia. La nueva especie se diferencia de todas las especies conocidas de Riama por folidosis, morfología hemipenial y patrón de coloración. Las cuatro especies endémicas de Colombia, Riama columbiana, Riama laevis, Riama stellae, y Riama striata se distribuyen en diferentes bosques montanos, y no se conoce ningún registro de simpatría con la nueva especie. La nueva especie se asemeja más a R. laevis de la Cordillera Occidental de los Andes Colombianos, pero difiere de esta principalmente por tener 15 hileras longitudinales de escamas dorsales, 21–23 hileras transversas de escamas ventrales en machos, 22 en hembras y las extremidades no se sobreponen o solo se tocan una con la otra cuando son apoyadas contra el cuerpo.
A new species of a moderately large Lepidoblepharis from the western slopes of the Andes, provinces of Pichincha and Cotopaxi, Ecuador, is described. It is characterized by its high, conical dorsal scales; 14– 17 lamellae under the fourth toe; V-shaped mental scale followed by 5–7 small postmentals; and dark brown color (in preservative) with almost no pattern. The new species is compared with its congeners. A redescription of Lepidoblepharis grandis Miyata, 1985, also known from the province of Pichincha, is presented.
The Yutajé-Corocoro massif is a highly eroded sandstone table mountain, with internal drainage mainly to the central valley of the Río Corocoro, a stream in the headwater drainage of the Río Manapiare - some 100 km east of the middle Río Orinoco, at the northern edge of the State of Amazonas in southern Venezuela. The rocky soil supports a mosaic of diverse scrubland and forest, with small tepui meadows at the higher elevations. The herpetofauna is depauperate, as is typical of the Venezuelan tepuis. Eight species of amphibians and reptiles were collected during a 7-day period in the dry season (February). This sample includes two new frogs (Hyalinobatrachium eccentricum, n. sp., Centrolenidae; Colostethus undulatus, n. sp., Dendrobatidae) and a new genus and species of lizards (Adercosaurus vixadnexus, n. gen. & sp., Teiidae), all of which were found in humid montane mossy forest at 1700-1750 m elevation. Another new lizard (Tropidurus panstictus, n. sp., Tropiduridae) was discovered at lower elevations (180-1220 m), especially in dry scrub. The fauna also includes a widespread lowland frog (Pseudopaludicola llanera Lynch), two tepui frogs (Eleutherodactylus cantitans Myers and Donnelly; E. yaviensis Myers and Donnelly), a tepui lizard (Prionodactylus goeleti [Myers and Donnelly], new combination), a snake (Liophis?) that escaped capture, and another snake (Thamnodynastes corocoroensis Gorzula and Ayarzagüena) obtained by S. Gorzula in 1987. The two Eleutherodactylus and the Prionodactylus also occur on neighboring Cerro Yaví (the type locality), although one of the frogs (E. yaviensis) and the lizard show evidence of differentiation. Based on the original description, the snake Thamnodynastes corocoroensis appears to be distinct from a related species on Cerro Yaví. Two of the new species exhibit characters that are novel or not previously noted. The dendrobatid frog Colostethus undulatus, n. sp. has a glandular supracarpal pad atop the wrist, being best developed in males. This species, which also has the parasphenoid bone curiously concealed, seems to be unusual among tepui Colostethus in lacking the recently described median lingual process. The centrolenid frog Hyalinobatrachium eccentricum, n. sp. has a peculiar bicolored iris, with a dark median sector that conceals the pupil and which apparently dilates with the pupil. This character is retained in preservative and differentiates H. eccentricum from H. crurifasciatum Donnelly and Myers. Both species share a previously overlooked bubblelike structure in the web between the third and fourth fingers, herein termed bulla (possibly parasite induced?).
We describe two new species of Proctoporus and a new species of Euspondylus (all from central Peru, Departamento de Huánuco, 2545 to 3010 m elevation). The new species are distinguished from all species presently assigned to Proctoporus and Euspondylus by unique characteristics of pholidosis, morphometrics, and color pattern. A key to the Peruvian species of Proctoporus and Euspondylus is provided. The current allocation of species to Proctoporus and Euspandylus may not reflect the phylogenetic relationships of the species under consideration. Based on the available data there appears to be no evidence to justify the separation of these genera. However, because of nomenclatural problems that would arise from synonymizing the two genera (e.g., producing a secondary homonymy in the case of Proctoporus guentheri Boettger and Ecpleopus guentheri O'Shaughnessy), we chose to retain the two genera until phylogenetic information is available.
A molecular phylogeny was reconstructed for 26 recognized genera of the Gymnophthalmidae using a total of 2379 bp of mitochondrial (12S, 16S and ND4) and nuclear (18S and c-mos) DNA sequences. We performed maximum parsimony (MP) and maximum likelihood (ML) analyses, and data partitions were analysed separately and in combination under MP. ML analyses were carried out only on the combined sequences for computational simplicity. Robustness for the recovered nodes was assessed with bootstrap and partitioned Bremer support (PBS) analyses. The total molecular evidence provided a better-resolved hypothesis than did separate analysis of individual partitions, and the PBS analysis indicates congruence among independent partitions for support of some internal nodes. Based on this hypothesis, a new classification for the family is proposed. Alopoglossus, the sister group of all the other Gymnophthalmidae was allocated to a new subfamily Alopoglossinae, and Rhachisaurus (a new genus for Anotosaura brachylepis) to the new Rhachisaurinae. Two tribes are recognized within the subfamily Gymnophthalminae: Heterodactylini and Gymnophthalmini, and two others within Cercosaurinae (Ecpleopini and Cercosaurini). Some ecological and evolutionary implications of the phylogenetic hypothesis are considered, including the independent occurrence of limb reduction, body elongation, and other characters associated with fossoriality.