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Mata-Silva, Vicente, Wilson, Larry David (2016): The taxonomic status of Tantilla marcovani Lema 2004 (Squamata: Colubridae). Zootaxa 4092 (3): 421-425, DOI: http://doi.org/10.11646/zootaxa.4092.3.7

The taxonomic status of Tantilla marcovani Lema 2004 (Squamata: Colubridae) VICENTE MATA-SILVA 1 & LARRY DAVID WILSON 2 1 Department of Biological Sciences, The University of Texas at El Paso, El Paso, Texas 79968 -0500, U.S.A. E-mail: vmata@utep.edu 2 Centro Zamorano de Biodiversidad, Escuela Agr��cola Panamericana Zamorano, Departmento de Francisco Moraz��n, Honduras; 16010 SW 207 th Avenue, Miami, Florida 33187 - 1056, U.S.A. E-mail: bufodoc@aol.com The colubrid snake genus Tantilla currently comprises 62 valid species distributed from the southern United States of America to southern Bolivia in the west, and Uruguay and northern Argentina in the east (Wilson & Mata-Silva 2014, 2015). Of these 62 species, 13 are found in South America (Wilson & Mata-Silva 2014, 2015). The most recently described of these Southamerican species is Tantilla marcovani Lema, 2004. While we were working on a checklist and key to the members of the Tantilla clade (sensu Holm 2008, including the genera Geagras Cope 1876, Scolecophis Fitzinger 184 3, Tantilla Baird & Girard 1853, and Tantillita Smith 1941; Wilson & Mata-Silva 2015), it became apparent that the diagnosis of T. marcovani entirely falls to distinguish this nominal taxon from the widespread and extensively variable T. melanocephala Linnaeus 1758. Tantilla melanocephala, as presently envisioned, is the most broadly distributed member of its genus (Wilson & Mena 1980; Savage 2002; Greenbaum et al. 2004), occurring from Panama southward into the major portion of the South American continent, apart from Chile and southern Argentina. The concept of Tantilla melanocephala has changed somewhat since the work of Wilson & Mena (1980). This conceptual change was effected mainly by two studies. Initially, Savage (2002) resurrected two names (T. armillata Cope ��� 1875 ��� [1876] and T. ruficeps Cope 1894) from the synonymy of T. melanocephala, as predicted by Wilson & Mena (1980). Savage (2002: 693) examined the status of the populations allocated to T. melanocephala by Wilson & Mena (1980) and concluded that ���in Costa Rica, where the ranges of T. armillata and T. ruficeps meet, the two behave as distinct species [and that] the populations referred to T. melanocephala from Guatemala to western Panama are considered to constitute two distinct species, T. armillata and T. ruficeps, that are marginally sympatric in central Costa Rica.��� Subsequently, Greenbaum et al. (2004) undertook a detailed study of the variation of 42 specimens allocated to T. melanocephala from Panama, Colombia, and Ecuador west of the Andes, as compared to that exhibited by four specimens allocated to T. equatoriana Wilson & Mena 1980, including the holotype of this nominal species. Greenbaum et al. (2004: 457) concluded that, based on their analysis of several characters of color pattern and the numbers of subcaudals, ���and a Principal Components Analysis of morphometric variation of T. equatoriana and T. melanocephala,��� the characters used by Wilson & Mena (1980) to diagnose T. equatoriana all overlap with those evidenced by the material of T. melanocephala they examined; thus, they treated ��� T. equatoriana as a junior synonym of T. melanocephala.��� As a result of the actions taken by Savage (2002) and Greenbaum et al. (2004), the range of T. melanocephala became restricted to South America and central and eastern Panama. Lema (2004) described Tantilla marcovani, based on a single specimen (Figs. 1���2), an adult male from ���Pico do Jabre, highest point of Paraiba State, Brazil at 50 km SW from Teixeira city (07�� 07��� 48 ������S, 37 �� 08��� 60 ������W), Matureia locality��� (Lema 2004: 269). He further noted that ���the region is located in highlands within Caatinga, with 1090 m above sea level.��� Since the original description, the name Tantilla marcovani has appeared infrequently in the literature (Freitas & Silva 2007; Hamdan & Lira-da-Silva 2012; Guedes et al. 2014). Hamdan & Lira-da-Silva (2012) reported a specimen identified as T. marcovani from Santo In��cio in the state of Bahia, which they indicated constituted the second record for the species from a locality ca. 456 km N of the type-locality. They did not state, however, how they identified their specimen as belonging to T. marcovani, as opposed to T. melanocephala. Guedes et al. (2014) indicated that ���the determination of this species is problematic and we thus opted to restrict the name T. marcovani only to specimens collected in Pico do Jabre��� (the type-locality). Thus, their distribution map (Guedes et al. 2014: figure 26.1) for this species and T. melanocephala indicates a single locality for T. marcovani (Pico do Jabre) located within the range of T. melanocephala, which is recorded from throughout the Caatinga. Based on the opinion expressed about the recognizability of T. marcovani by Guedes et al. (2014) and our own reading of the diagnosis of this nominal taxon in Lema (2004), we have undertaken an examination of the utility of this diagnosis to discriminate between the restricted Tantilla marcovani and the widespread Tantilla melanocephala. The description and diagnosis of Tantilla marcovani are based on the data of their holotype. The comparative statements in the diagnosis are based, apparently, on the features exhibited by a series of 20 additional specimens of T. melanocephala from Argentina, Brazil, and Uruguay, although Lema (2004) did not state this explicitly to be the case. Regardless, Lema did not examine material from elsewhere in the range of T. melanocephala in constructing the diagnosis for T. marcovani. In attempting to decipher the T. marcovani diagnosis, we constructed a table that indicates the condition of nine features of head shape and coloration. In doing so, we have hewed to the language used by Lema (2004), which as will be noted below, is, in some cases, indecipherable. Most of our commentaries are based on an extensive sample of Tantilla melanocephala examined by one of us (LDW), previously listed in Wilson & Mena (1980). In order to establish the taxonomic status of Tantilla marcovani we reviewed the features formally employed in its diagnosis, as following: (1) Head width, Lema (2004: 269) stated the head of T. marcovani is ���wider��� than that of T. melanocephala, which is indicated to be ���elongated��� (Table 1). Since neither of these conditions is defined, they cannot be used to distinguish T. marcovani from T. melanocephala; (2) Snout shape, Lema (2004: 269) indicated the snout to be ���wide and straight��� in T. marcovani and ���slender and long��� in T. melanocephala. Neither of the terms ���wide��� nor ���slender��� are defined and, so, cannot be used for discrimination. The terms ���straight��� and ���long��� are also not defined and, furthermore, do not describe conditions of the same character and also cannot be used for species discrimination; (3) Head diameter, Lema (2004: 269���270) described the head ���diameter��� (presumably the head width) as ���wider than neck��� in T. marcovani and ���equal to neck��� in T. melanocephala. The junior author of this paper (LDW) examined a large number of specimens of T. melanocephala during the writing of Wilson & Mena (1980) and found none in which the head was anything other than ���wider than the neck.��� Even so, this comparison is not supported by any mensural data and is based on a comparison of the holotype of T. marcovani with only 20 specimens of T. melanocephala from a restricted portion of its total range of distribution and, in our opinion, also is not usable to discriminate between the two taxa; (4) Snout color, Lema (2004: 270) indicated the snout color of T. marcovani to be ���light immaculate��� meaning presumably ���immaculate cream��� and that of T. melanocephala as ���black or black blotched.��� Wilson & Mena (1980), however, characterized both color pattern types C and D, which are the two found in Brazil, as having ���pale pigment on internasals and prefrontals��� (Table 1). Therefore, this distinction does not hold up to scrutiny; (5) Black head cap, Lema (2004: 270) described the dark head cap as ���small, restricted to frontal and parietal region��� in T. marcovani and ���over all the head from the snout to the nape��� in T. melanocephala. Apparently, this distinction reflects Lema���s contention that T. marcovani differed from T. melanocephala in having a pale snout. As we noted above, however, the pattern types found throughout Brazil also have pale snouts, so this feature will not operate for discrimination either; (6) Color of head cap, Lema (2004: 270) stated that the dark head cap of T. marcovani was ���grayish brown��� (presumably in life) and that of T. melanocephala is ���black.��� The color photograph of the holotype of T. marcovani (figure 2 of Lema 2004 and Fig. 2 of present study), however, demonstrates the head cap to be dark brown in color with some paler marbling. Furthermore, Wilson & Mena (1980; Table 1) stated that the dark head cap in color pattern type C is ���brown to dark brown��� and ���brown��� in color pattern type D and not black; (7) Oblique black band over eyes, such a feature described by Lema (2004: 270) was indicated by Wilson & Mena (1980; Table 1) to be the ���lateral extension of dark head cap.��� Lema (2004) stated that in T. marcovani it is ���attaining first supralabials, with remainder cream��� and in T. melanocephala there is ���none present on eyes and last supralabial.��� Neither of these descriptions makes sense to us and otherwise do not describe the actual appearance of this feature in these snakes. The color photograph of the holotype of T. marcovani (Fig. 2) illustrates that the lateral extension of the dark head cap does not reach the lip, differing slightly from the condition indicated in Table 1 of Wilson & Mena (1980) for color pattern types C and D, in which the lateral extension is said to reach the lip (as is evident, for example in the drawing of the head of a member of color pattern type D from Venezuela in figure 3 from Wilson & Mena 1980). Given that Lema���s description of this feature in the two taxa is incomprehensible, it is not possible to use it for the purposes of distinction; (8) Posterior end of head, it is not clear how Lema (2004) defined the ���posterior end of [the] head,��� but perhaps he meant the posterior end of the parietal scales. Otherwise, the posterior end of the head is defined usually as the point to which the posterior end of the lower jaw reaches. Regardless, Lema stated that the ���posterior end of the head��� is ���cream attaining nuchal light collar��� in T. marcovani and ���not black with pair of narrow cream blotches��� in T. melanocephala. These descriptions also are indecipherable, so their utility is indeterminable. Nonetheless, the drawing of the head of the holotype of T. marcovani in Lema (2004) and our photograph (Fig. 2) indicates that the pale nuchal band begins at about the mid-point of the parietals and extends to about the mid-point of the first nuchal scale behind the posteriormost point of the parietal scales. The dark head cap is also almost in contact with an anteriomedial extension of the dark nape band, thus almost completely dividing the pale nuchal band medially; (9) Black cervical collar, this is the feature described by Wilson & Mena (1980) as the dark nape band. Lema (2004) described it in T. marcovani as ���evident, only vertebral, three vertebral scales long��� and in T. melanocephala as ���not narrow, feeble or almost indistinct dark collar.��� These statements do not function to describe the actual conditions in these snakes and, therefore, cannot be used to discriminate between them. Nonetheless, in the holotype of T. marcovani, the dark nape band is almost connected to the dark head cap along the common parietal suture and otherwise extends for four middorsal scales posterior to the posterior end of the common parietal suture (Fig. 2). Wilson & Mena (1980: Table 1) indicated the mean length of the dark nape band in the members of color pattern type C to be 3.9 middorsal scales and that of those belonging to color pattern type D to be 3.6 middorsal scales. Thus, the condition in the holotype of T. marcovani is close to these mean values. Based on our analysis of the diagnosis for Tantilla marcovani, there are no features included within it that can act to distinguish (= unique state or combination of characters) this putative taxon from the widespread and highly variable T. melanocephala in its current concept. Although the number of ventral scales in the male holotype of T. marcovani was not used by Lema (2004) in his diagnosis, it is worthwhile to compare it to the data supplied by Wilson & Mena (1980, Table 3). This segmental count is the only one usable for such a comparison, since the tail of the holotype is incomplete. The ventral count given by Lema for this specimen is 145. This number is reasonably close to the mean number of ventrals given by Wilson & Mena (1980) for male T. melanocephala from Brazil, which is 143.4 (range 136���154). Thus, this feature would not have served either to differentiate T. marcovani from T. melanocephala. In conclusion, while attempting to construct an identification key for the 66 members of the Tantilla clade (sensu Holm 2008) we discovered that the diagnosis proposed by Lema (2004) for Tantilla marcovani would not serve to distinguish it from the widespread and extensively variable T. melanocephala and, thus, would not allow for the design of a couplet in the key to accomplish this. We undertook, therefore, to examine Lema���s diagnosis in detail. This analysis demonstrated that no single feature of this diagnosis would work to distinguish T. marcovani from T. melanocephala. Therefore, we conclude that Tantilla marcovani Lema, 2004, should be placed into the synonymy of Tantilla melanocephala (Linnaeus, 1758)., Published as part of Mata-Silva, Vicente & Wilson, Larry David, 2016, The taxonomic status of Tantilla marcovani Lema 2004 (Squamata: Colubridae), pp. 421-425 in Zootaxa 4092 (3) on pages 421-424, DOI: 10.11646/zootaxa.4092.3.7, http://zenodo.org/record/258595

A new species of arboreal lizard of the genus Celestus is deseribed from north-eentra1 Honduras. This new species appears to be most elosely related to C. bivittatus and is easily distinguished from that species by haying longer digits on both the fore- and hindlimbs. Se describe un·a especie nueva de Celestus arborea de el centro del norte de Honduras. Esta nueva especie parece ser mas relacionada a C. bivíttatus y es mas facil de distinguir esta especie ya que tien e dígitos mas largos en ambas extremidades del frente y traseras.

The conservation status of the members of the Honduran herpetofauna is discussed. Based on current and projected future human population growth, it is posited that the entire herpetofauna is endangered. The known herpetofauna of Honduras currently consists of 334 species, including 117 amphibians and 217 reptiles (including six marine reptiles, which are not discussed in this paper). The greatest number of species occur at low and moderate elevations in lowland and/or mesic forest formations, in the Northern and Southern Cordilleras of the Serranía, and the ecophysiographic areas of the Caribbean coastal plain and foothills. Slightly more than one-third of the herpetofauna consists of endemic species or those otherwise restricted to Nuclear Middle America. Honduras is an area severely affected by amphibian population decline, with close to one-half of the amphibian fauna threatened, endangered, or extinct. The principal threats to the survival of members of the herpetofauna are uncontrolled human population growth and its corollaries, habitat alteration and destruction, pollution, pest and predator control, overhunting, and overexploitation. No Honduran amphibians or reptiles are entirely free of human impact. A gauge is used to estimate environmental vulnerability of amphibian species, using measures of extent of geographic range, extent of ecological distribution, and degree of specialization of reproductive mode. A similar gauge is developed for reptiles, using the first two measures for amphibian vulnerability, and a third scale for the degree of human persecution. Based on these gauges, amphibians and reptiles show an actual range of Environmental Vulnerability Scores (EVS) almost as broad as the theoretical range. Based on the actual EVS, both amphibian and reptilian species are divided into three categories of low, medium, and high vulnerability. There are 24 low vulnerability amphibians and 47 reptiles, 43 medium vulnerability amphibians and 111 reptiles, and 50 high vulnerability amphibians and 53 reptiles. Theoretical EVS values are assessed against available information on current population status of endemic and Nuclear Middle American taxa. Almost half (48.8%) of the endemic species of Honduran amphibians are already extinct or have populations that are in decline. Populations of 40.0% of the Nuclear Middle American amphibian species are extirpated or in decline. A little less than a third (27.0%) of the endemic reptiles are thought to have declining populations. Almost six of every ten (54.5%) of the Nuclear Middle American reptilian species are thought to have declining populations. EVS values provide a useful indicator of potential for endangerment, illustrating that the species whose populations are currently in decline or are extinct or extirpated have relatively high EVS. All high EVS species need to be monitored closely for changes in population status. A set of recommendations are offered, assuming that biotic reserves in Honduras can be safeguarded, that it is hoped will lead to a system of robust, healthy, and economically self-sustaining protected areas for the country’s herpetofauna. These recommendations will have to be enacted swiftly, however, due to unremitting pressure from human population growth and the resulting deforestation. Se discute el estatus de conservación de los miembros de la herpetofauna de Honduras. Basados en el crecimiento presente y proyectado de la población del ser humano, se propone que toda la fauna herpetológica de Honduras está en peligro de extinción. Lo que se conoce de la fauna herpetológica hondureña en el presente consiste de 334 especies, incluyendo 117 anfibios y 217 reptiles (incluyendo seis reptiles marinos, que no se discuten en este artículo). La mayoria de las especies se presentan en bajas y moderadas elevaciones en formaciones forestales de tierras bajas y/o húmedas, en las Cordilleras Septentrional y Meridional de la Serranía, y las áreas ecofisiográficas de la costa y las faldas de la montaña del Caribe. Un poco mas de un tercero de la fauna herpetológica consiste de especies endémicas o sino de esas especies restringidas al Mesoamérica Nuclear. Honduras es una área severemente afectada por la disminución de las poblaciones de anfibios, con cerca de la mitad de la fauna anfibia amenazada, en peligro, o extinta. Las principales amenazas a la sobreviviencia de los miembros de la fauna herpetológica son el crecimiento sin control de la población humana y sus vástagos, la alteración y destructión de habitación, polución, el control de pestes y predadores, el exceso de caza y explotación. Ningun anfibio o reptil hondureño está totalmente libre de el impacto humano. Se ha desarrollado una regla de medir para estimar la vulnerabilidad ambiental de las especies de anfibios, usando medidas de extensión del rango geografíco, amplitud de distribución ecológica, y estado de especialización del modo de reproducción. Se ha desarrollado una medida similar para los reptiles, usando las dos primeras medidas de vulnerabilidad usados con los anfibios, y una tercera medida para el grado de persecusión humana. Basados en estas medidas, los anfibios y reptiles muestran un rango actual de una marca de vulnerabilidad medioambiental (EVS) casi tan amplia como el rango teorético. Basados en la EVS, ambas especies de anfibios y reptiles están divididas en tres categorías, de baja, media, y alta vulnerabilidad. Hay 24 especies de anfibios y 47 de reptiles de baja vulnerabilidad, 43 especies de anfibios y 111 de reptiles de media vulnerabilidad, y 50 especies de anfibios y 53 de reptiles de alta vulnerabilidad. Teoréticamente, los valores de EVS son determinados de acuerdo de información disponible del estado presente de las taxas endémicas de Mesoamérica Nuclear. Casi la mitad (48.8%) de las especies endémicas de anfibios hondureños están ya extintos o tienen poblaciones en disminución. Poblaciones de 40.0% de las especies de anfibios de Mesoamérica Nuclear están extintas o en disminución. Un poco menos de un tercio (27.0%) de los reptiles endémicos se piensa que tienen poblaciones en disminución. Casi seis de cada diez (54.5%) de las especies de reptiles de Mesoamérica Nuclear se piensa que tienen poblaciones en disminución. Los valores de EVS proporcionan un indicador utíl del riesgo potencial, el cual muestra que las especies cuyas poblaciones actuales están disminuyendo, o son extintos o extirpados tienen EVS relativamente altos. Todas las especies con un EVS alto necesitan ser observadas de cerca para anotar los cambios en el estado de las poblaciones. Ofrecemos un grupo de recomendaciones, asumiendo que las reservas bióticas de Honduras pueden ser preservadas, se espera que esto resulte en un sistema de áreas protegidas que es robusta, saludable, y sostenible economicamente para la fauna herpetológica del país. Estas recomendaciones tienen que ser observados rapidamente, debido a la presión continua causada por el crecimiento de la población humana y la resultante destrucción de los bosques.

Nototriton tomamorum new species (Figs. 2, 3) Holotype: A female (UF 155377) from 2.5 km NNE of La Fortuna (15 �� 25.965 ���N, 87 �� 18.556 ���W), 1550 m, Refugio de Vida Silvestre Texiguat, Departamento de Yoro, Honduras, collected 9 April 2008 by J. Slapcinsky and the field team of J. M. Butler, L. P. Ketzler, N. M. Stewart, J. H. Townsend, and L. D. Wilson. Original field number JHT 2437. Diagnosis. A small member of the genus Nototriton (SVL= 26.9 mm; Table 2) based on having 13 costal grooves (> 16 costal grooves in Oedipina), hands and feet longer than broad (hands and feet broader than long in Bolitoglossa), and nares that are smaller than most Cryptotriton and Dendrotriton (Fig. 3 A, B; 0.018 NL/ SVL; 0.020���0.029 NL/SVL in Cryptotriton [except some individuals of C. veraepacis] and Dendrotriton). Cryptotriton veraepacis has nares ranging from 0.017���0.027 NL/SVL (mean 0.022), and can be differentiated from N. tomamorum by having a uniformly dark gray ventral surface (ventral surface pale with gray flecks in N. tomamorum; Fig. 2). Generic placement in Nototriton is also strongly supported by sequence data from the mitochondrial genes 16 S and cob (Table 3; Fig. 4). This new species is distinguished from all other Nototriton, except N. richardi and N. tapanti, by having syndactylous hands and feet (Fig. 3 C, D; hands and feet with free, differentiated toes in all other species) and relatively large nares (Fig. 3 A, B; 0.018 NL/SVL versus 0.010���0.016 in N. picadoi, 0.003���0.014 in N. abscondens, 0.012 in N. stuarti, 0.005���0.011 in N. barbouri, 0.006���0.009 in N. lignicola, 0.004���0.009 in N. guanacaste, 0.004���0.005 in N. brodiei, 0.003 in N. limnospectator, 0.003 in N. major, and 0.002���0.003 in N. saslaya). Nototriton tomamorum can be further differentiated from members of the N. barbouri group by having a broader head (0.145 HW/SVL versus 0.138 in N. stuarti, 0.104���0.132 in N. barbouri, 0.120 in N. brodiei, 0.103���0.118 in N. lignicola, and 0.095���0.118 in N. limnospectator) and fewer maxillary teeth (26, versus 36 in N. stuarti, 41���54 in N. barbouri, 42���55 in N. limnospectator, 46���54 in N. lignicola, and 60���62 in N. brodiei), from members of the N. picadoi group by having a relatively shorter tail (0.911 TL/SVL, versus 1.441 in N. major, 1.123���1.344 in N. picadoi, 1.013��� 1.365 in N. abscondens, 1.210���1.337 in N. guanacaste, and 1.10���1.30 in N. gamezi) and narrower feet (0.037 HFW /SVL, versus 0.058���0.071 in N. abscondens, 0.059 in N. major, 0.060���0.070 in N. picadoi, and 0.066��� 0.072 in N. guanacaste), and from N. saslaya by having shorter forelimbs (0.160 FLL/SVL, versus 0.194��� 0.210 in N. saslaya) and hind limbs (0.197 HLL /SVL, versus 0.217���0.244 in N. saslaya) and narrower feet (0.037 HFW /SVL, versus 0.075���0.091 in N. saslaya). The new species also differs from N. richardi and N. tapanti in having a pale ventral surface mottled with gray chromatophores (ventral surface brown with dark flecks in N. richardi and dark brown in N. tapanti), by having a tail that is shorter than the snout-vent length (0.91 TL/SVL, versus 1.072���1.482 in N. richardi and 1.205 in N. tapanti), longer forelimbs (0.160 FLL/SVL, versus 0.140���0.146 in N. richardi and 0.147 in N. tapanti), longer hind limbs (0.197 HLL /SVL, versus 0.174��� 0.187 in N. richardi and 0.174 in N. tapanti), and narrower feet (0.037 HFW /SVL, versus 0.044���0.050 in N. richardi and 0.041 in N. tapanti). This new species is also well differentiated from all other species of Nototriton based on mitochondrial sequence data (Table 3), and is 3.6 ���6.0% divergent on 16 S and 10.2��� 15.1 % divergent on cob from all other congeners. Description of holotype. Nototriton tomamorum is known only from a single, presumably female (mental gland and cloacal papillae absent) specimen, preserved with its mouth open and tongue extended, and is a relatively small member of the genus (SVL= 26.9 mm, total length= 51.4 mm) with a slender body and reduced limbs. Its head is rounded and slightly broader than the body, and the nostrils are relatively large (NL/ SVL= 0.018), and the snout is rounded and of moderate length (Fig. 3 A, B). The nasolabial protuberances are apparent but not well developed, and barely extend below the upper lip line. The eyes are relatively large and protuberant, and the parotoid glands appear large but not well defined. The teeth are exceedingly small; there are approximately 26 maxillaries, 4 premaxillaries set slightly forward from the maxillary teeth, and 11 vomerines; vomerine teeth are arranged in two short medially-positioned arches. The limbs are short (CLL/ SVL= 0.36), with the adpressed limbs being separated by approximately 5.5 costal grooves. The hands and feet are narrow and have poorly-developed, poorly-differentiated digits that are fused and lack subdigital pads (Fig. 3 C, D). The free tips of digits III on the hands and III and IV on the feet are pointed, and digits I, II and IV on the hands and I, II, and V on the feet are very short and essentially completely fused, being demarcated by shallow grooves on the dorsal side of the feet. The relative length of the digits is IMeasurements of holotype (in mm). SVL 26.9; AG 15.2; TW 3.6; HL 4.8; HW 3.9; TL 24.5; HLL 5.3; FLL 4.3; CLL 9.6; HFL 1.7; HFW 1.2; NL 0.5; eye length 1.6; eye width 1.2; interorbital distance 1.4; anterior rim of orbit to snout 1.1; distance separating internal nares 0.8; distance separating external nares 1.8; snout projection beyond mandible 0.6; tip of snout to axilla 7.7; distance from axilla to groin 15.2; snout to anterior edge of vent 24.9; tail depth at basal constriction 2.8; tail width at basal constriction 2.4. Coloration of holotype. Dorsal surfaces of head, body, and tail medium grayish brown, with profuse pale chromatophores laterally, becoming less abundant dorsolaterally. The head has some pale mottling on the top of the snout, and two irregular lines of pale chromatophores extending from the lateral region above the forelimbs onto the posterior portion of the head and parotoid glands. There is a very thin, pale middorsal stripe, with a herringbone pattern with lines extending from the middorsal stripe posteriorly. There is an indistinct dark dorsolateral stripe starting about one-third the way down the trunk and extending onto the proximal one-third of the tail. Ventral surface of head, body, and tail cream, mottled with dark gray chromatophores, becoming somewhat more profuse toward the distal end of the tail. Etymology. The specific name ��� tomamorum ��� means ���belonging to the Tomams.��� Tomams are the highest level of deities recognized in the belief system of the indigenous Tolupan of Honduras, of which there are four: Tomam Pones Popawai (Grandfather Tomam), his wife Tomam Pones Namawai (mother of all that exists), and their children Tomam the Elder and Tomam the Younger (Chapman 1992). This name is given in recognition that the Tolupan are the traditional inhabitants of this area and that this new species is known only from the Cordillera Nombre de Dios, or ���Name of God Mountains,��� a name which, ironically, was given by 15 th century Spanish explorers. Natural history. The single known specimen of Nototriton tomamorum was collected during the daytime from leaf litter packed onto a rock ledge alongside a small creek at about 1550 m elevation in the Lower Montane Wet Forest formation (Fig. 5). This species is presumably endemic to the vicinity of the type locality, and is likely restricted to the Lower Montane Wet Forest around the neighboring peaks of Cerro San Francisco and Cerro Texiguat. The locality can be characterized as a narrow, steep-sided canyon with riparian forest, and more xeric upland pine-oak forest dominating the terrain above the canyon. The pine-oak forest transitions to mesic cloud forest (or remnant cloud forest) uphill from this locality. This species is sympatric with a congener, N. barbouri, and three other plethodontids: Bolitoglossa dofleini, Bolitoglossa porrasorum and Oedipina gephyra. The canyon where the holotype of N. tomamorum was collected is also the type locality of the hylid frog Isthmohyla insolita, and a known locality for the anurans Lithobates maculatus, Plectrohyla guatemalensis, and Ptychohyla spinipollex. Conservation status. Based on the criteria of the IUCN (2001), Nototriton tomamorum should be classified as Critically Endangered (B 1 ab[iii]+ 2 ab[iii]) due to the extremely limited extent of its known distribution and the direct threat to the remaining habitat around that locality. The primary threat to the type and only known locality for this species is from illegal timber extraction and wholesale slash-and-burn clearing of forest for bean cultivation once it has been depleted of valuable hardwoods, such as mahogany (Fig. 6). Systematic relationships. Nototriton tomamorum bears the strongest morphological resemblance to members of the N. richardi group (Table 2), which are endemic to the central highlands of Costa Rica. The two species, N. richardi and N. tapanti, are the most morphologically distinctive members of the genus, differing from all other Nototriton by having syndactylous feet, relatively large nostrils, and frontal processes of the premaxillary arising separately from the maxillary (Good & Wake 1993; Papenfuss & Wake 1987; Savage 2002). According to Savage (2002), N. richardi is distributed from 1370 to 1800 m elevation along the Atlantic slope of the Cordillera Central, and N. tapanti at a single locality at 1300 m elevation in the northern portion of the Cordillera de Talamanca. These localities are situated in the central montane spine of Costa Rica in Lower Central America and are separated widely from the type locality of N. tomamorum by over 670 airline km and much of the central highlands of Honduras and the lowland Nicaraguan Depression. Despite the morphological similarities between N. tomamorum and members of the N. richardi group, our phylogenetic analysis (Fig. 4) does not support the inclusion of N. tomamorum in the N. richardi group. Preliminary phylogenetic analysis of fragments from two mitochondrial genes (16 S and cob) weakly supports (posterior probability = 0.55, bootstrap support = 58 %) N. tomamorum as sister taxon to the rest of the N. barbouri group (Fig. 4). We also recovered N. richardi as the weakly supported sister (posterior probability = 0.58, bootstrap support = 59 %) to the N. picadoi group, a finding consistent with previous phylogenetic hypotheses for Nototriton and N. richardi (Garc��a-Par��s & Wake 2000; Wiens et al. 2007; Adams et al. 2009). Sequence data from the holotype of Nototriton tomamorum are at least 12.5 % (16 S) and 20.6 % (cob) divergent from representatives of morphologically-similar genera of Central American plethodontids (Cryptotriton, Dendrotriton, and Oedipina), but less than 6.0% (16 S) and 15.1 % (cob) divergent from other species of Nototriton, supporting placement of N. tomamorum in this genus (Table 3). Within the genus Nototriton, the new species demonstrates differing patterns of sequence divergence on the two genes sampled. For 16 S, N. tomamorum is closest to N. limnospectator (3.6���3.8 %), a biogeographically-logical situation given the relative geographical proximity of RVS Texiguat to the type locality of N. limnospectator, approximately 100 airline km to the WSW in Parque Nacional Monta��a de Santa B��rbara; however N. tomamorum is 12.4 % divergent from N. limnospectator for cob, and for that gene is closer to the Costa Rican species N. abscondens (10.2 %) and N. gamezi (11.5 %) than to any other member of the N. barbouri group (Table 3). Given the morphological differentiation and lack of strong phylogenetic support, we refrain from assigning N. tomamorum to any recognized species group pending acquisition and analysis of additional molecular data from mitochondrial and nuclear genes. Our phylogenetic analyses included taxa and populations of Nototriton not available in previous studies (Garc��a-Par��s & Wake 2000; Wiens et al. 2007), including a sample from the vicinity of the type locality of N. barbouri (���Portillo Grande, Yoro, Honduras ��� [Schmidt 1936]). Inclusion of these data reveals the taxon N. barbouri to be paraphyletic (Fig. 4), with strong support for N. barbouri sensu stricto as the sister species of N. limnospectator and for the remaining populations assigned to N. barbouri (N. sp. in Fig 4) from the Cordillera Nombre de Dios to be sister to N. brodiei from NE Guatemala and NW Honduras. We are currently preparing an intergrative systematic revision of the N. barbouri group that will use molecular and morphological analyses to resolve the taxonomic status of populations assigned to N. barbouri. The herpetofauna of the RVS Texiguat is characterized by a remarkable degree of endemism (McCranie & Casta��eda 2004 a; Wilson & McCranie 2004 b). Currently, 39 species are known from 1550 m in elevation and above in this refuge. Of this number, 27 are considered to be conservation priority species (Table 4), defined as those species judged to occupy one of three IUCN categories Critically Endangered, Endangered, or Vulnerable. Nine of the 27 are considered Critically Endangered, 12 are Endangered, and six are Vulnerable; 19 (70.4 %) of these species (four salamanders, seven anurans, four lizards, and four snakes) are endemic to Honduras (Table 4). Of the remainder, six are endemic to Nuclear Central America and two have widespread distributions. As indicated by Townsend & Wilson (In press), 91 species of amphibians and reptiles are endemic to Honduras, thus the Texiguat figure is 20.9 % of the total. Therefore, RVS Texiguat qualifies as a major center of endemism in Honduras, as well as in the whole of eastern Nuclear Central America (as defined by Campbell, 1999). The Environmental Vulnerability Scores, as tabulated by Townsend and Wilson (in press) and indicated in Table 4, indicate that the values for the Texiguat conservation priority species range from 9 to 17. Most of these species involved (16) are considered high vulnerability species (59.3 %), nine (33.3 %) are medium vulnerability species, and only two (7.4 %) have scores lying at the upper end of the range for low vulnerability species (Table 4). In summary, of the 27 species judged to be of conservation priority, 19 are endemic to Honduras and 16 are of high vulnerability. Based on the analysis above, Refugio de Vida Silvestre Texiguat is an area of immense importance for the conservation of the Honduran herpetofauna, especially its endemic component. Unfortunately, little attention has been paid to RVS Texiguat by conservation authorities past its initial designation as a protected area in 1987 (Wilson et al. 2001). Since 1991, ten species (including the present one) with type localities in the area have been described: Anolis kreutzi, A. purpurgularis, A. yoroensis, Celestus scansorius, Geophis damiani, Isthmohyla insolita, Nototriton tomamorum, Oedipina gephyra, Omoadiphas texiguatensis, and Rhadinaea tolpanorum (McCranie et al. 1993 a, b; Wilson et al. 1998; McCranie & Casta��eda 2004 a). Given the pattern of these finds, we expect that additional new taxa remain to be described from RVS Texiguat. During this period, significant environmental damage has been observed, most of it apparently as a consequence of the illegal extraction of mahogany and the subsequent conversion of the remaining forest to cropland for use in subsistence agriculture (Fig. 6). Most recently during our visit in April 2008, we observed that a large stretch of forest upstream from the type locality of Nototriton tomamorum and Isthmohyla insolita had been clear-cut down to the streamside and planted to beans (Fig. 6 C). Damage to the forests in this refuge also has occurred by natural means. The stream that constitutes the type locality of Nototriton tomamorum and Isthmohyla insolita (Fig. 5) was severely damaged by Hurricane Mitch in 1998. At that time, the habitat supporting I. insolita was scoured by high waters in the narrow canyon. The moss masses clinging to the branches of small trees overhanging the stream, within which the adults laid their eggs, were stripped away. Our trip in 2008 evidenced that the moss has yet to be reestablished as we found it in 1991 when I. insolita was discovered. Whether the reestablishment of these conditions will occur is doubtful. Isthmohyla insolita has been located in one or two nearby streams (McCranie & Casta��eda 2007), but the same anthropogenic activities detailed above also are occurring there. It is apparent that the natural habitat of this hotspot for herpetofaunal endemism will continue to succumb to the forces of habitat destruction unless steps are taken to protect it. Therefore, we use this opportunity not only to describe yet another endemic species from this reserve, but to attempt to raise RVS Texiguat���s international profile and issue an urgent call for its conservation. We offer the following recommendations, and emphasize the need for immediate and sustained action to prevent further loss of habitat: Carry out intensive reconnaissance of RVS Texiguat on the Caribbean side of Cerro Texiguat and Cerro San Francisco, to search for and identify previously unknown areas supporting populations of conservation priority species. The majority of work in RVS Texiguat has taken place in the vicinity of La Fortuna, which is accessed by means of a logging road established sometime before 1991. The Caribbean slope of RVS Texiguat remains virtually unexplored and has no known access routes, and so exploratory work is needed immediately to determine if this areas supports target species. Establish a habitat and species monitoring program led by Honduran biologists and local guardarecursos. This program has been initiated in cooperation with PROLANSATE (the organization tasked with managing RVS Texiguat). Initial plans are to hire six guardarecursos in early 2010 and begin training and capacity building for dev, Published as part of Townsend, Josiah H., Butler, Michael, Wilson, Larry David & Austin, James D., 2010, A distinctive new species of moss salamander (Caudata: Plethodontidae: Nototriton) from an imperiled Honduran endemism hotspot, pp. 1-16 in Zootaxa 2434 on pages 4-14, DOI: 10.5281/zenodo.194769, {"references":["Chapman, A. M. (1992). Masters of Animals: Oral Traditions of the Tolupan Indians, Honduras. Gordon and Breach Scientific Publishers, Philadelphia, Pennsylvania. xxi + 259 pp.","Good, D. A. & Wake, D. B. (1993) Systematic studies of the Costa Rican moss salamanders, genus Nototriton, with descriptions of three new species. Herpetological Monographs, 7, 131 - 159.","Papenfuss, T. J. & Wake, D. B. (1987) Two new species of plethodontid salamanders (genus Nototriton) from Mexico. Acta Zoologica Mexicana, 21, 1 - 16.","Savage, J. M. (2002) The Amphibians and Reptiles of Costa Rica: A Herpetofauna Between Two Continents, Between Two Seas. University of Chicago Press. Chicago, Illinois. xx + 934 pp.","Garcia-Paris, M. & Wake, D. B. (2000) Molecular phylogenetic analysis of relationships of the tropical salamander genera Oedipina and Nototriton, with descriptions of a new genus and three new species. Copeia, 2000, 42 - 70.","Wiens, J. J., Parra-Olea, G., Garcia-Paris, M. & Wake, D. B. (2007) Phylogenetic history underlies elevational biodiversity patterns in tropical salamanders. Proceedings of the Royal Society B, 274, 919 - 928.","Adams, D. C., Berns, C. M., Kozak, K. H. & Wiens, J. J. (2009) Are rates of species diversification correlated with rates of morphological evolution? Proceedings of the Royal Society B, 276, 2729 - 2738.","Schmidt, K. P. (1936) New amphibians and reptiles from Honduras in the Museum of Comparative Zoology. Proceedings of the Biological Society of Washington 49, 43 - 50.","McCranie, J. R. & Castaneda, F. E. (2004 a) A new species of snake of the genus Omoadiphas (Reptilia: Squamata: Colubridae) from the Cordillera Nombre de Dios in northern Honduras. Proceedings of the Biological Society of Washington, 117, 311 - 316.","Wilson, L. D. & McCranie, J. R. (2004 b) The herpetofauna of the cloud forests of Honduras. Amphibian and Reptile Conservation, 3, 34 - 48.","Campbell, J. A. (1999) Distribution patterns of amphibians in Middle America. In: Duellman, W. E. (Ed.), Patterns of Distribution of Amphibians: a global perspective. Johns Hopkins University Press, Baltimore, Maryland, pp. 111 - 210.","Wilson, L. D., McCranie, J. R. & Espinal, M. R. (2001) The ecogeography of the amphibians and reptiles of Honduras and the design of herpetological reserves. In: Johnson, J. D., Webb, R. G., & Flores-Villela, O. A. (Eds.). Mesoamerican Herpetology: Systematics, Zoogeography, and Conservation. Centennial Museum, Special Publication No. 1, The University of Texas at El Paso, pp. 109 - 158.","McCranie, J. R., Wilson, L. D. & Williams, K. L. (1993 a) A new species of Oedipina (Amphibia: Caudata: Plethodontidae) from northern Honduras. Proceedings of the Biological Society of Washington, 106, 385 - 389.","McCranie, J. R., Wilson, L. D. & Williams, K. L. (1993 b) New species of tree frog of the genus Hyla (Anura: Hylidae) from northern Honduras. Copeia, 1993, 1057 - 1062.","McCranie, J. R. & Castaneda, F. E. (2007) Guia de Campo de los Anfibios de Honduras. Bibliomania!, Salt Lake City, Utah, 304 pp."]}

The cloud forest amphibians and reptiles constitute the most important herpetofaunal segment in Honduras, due to the prevalence of endemic and Nuclear Middle American-restricted species. This segment, however, is subject to severe environmental threats due to the actions of humans. Of the 334 species of amphibians and reptiles currently known from Honduras, 122 are known to be distributed in cloud forest habitats. Cloud forest habitats are found throughout the mountainous interior of Honduras. They are subject to a Highland Wet climate, which features annual precipitation of >1500 mm and a mean annual temperature of

Anolis rubribarbaris (K��hler, McCranie, & Wilson 1999) Figs. 2���3 Norops rubribarbaris K��hler et al. 1999: 280. Holotype. UF 90206, an adult male from 4 km S of San Lu��s de los Planes, 1700 m elevation, northern slope of Monta��a de Santa B��rbara, Parque Nacional Monta��a de Santa B��rbara, Depto. Santa B��rbara, Honduras, collected 3 March 1994 by Fred G. Thompson, original field number FGT 5659. Referred specimens. UF 152660 (Figs. 2 a, 2 b), a small adult male, and UF 152661 ��� 62 (Fig. 3), adult females, all from disturbed cloud forest ca. 1���2 km W of El Cedral (14 �� 54.49 ���N, 88 ��05.38���W), 1720���1740 m, E slope of Monta��a de Santa B��rbara, Parque Nacional Monta��a de Santa B��rbara, Depto. Santa B��rbara, Honduras, collected 28 January 2008 by Leonel Erazo Ch��vez, Ileana R. Luque-Montes, Josiah H. Townsend, and Larry David Wilson. Diagnosis. Anolis rubribarbaris differs from all other Mesoamerican anoles, except those in the crassulus group, by having the following combination of characteristics: red dewlap, moderately to strongly enlarged medial dorsal scales, strongly keeled ventral scales, no more than one scale separating the supraorbital semicircles, four to five rows of loreals, suboculars and supralabials in contact, no axillary pocket, and heterogeneous flank squamation. Of the members of the crassulus group, A. rubribarbaris differs from A. amplisquamosus in having a red dewlap and enlarged middorsal scales grading into the dorsolateral scales (dewlap bright orange and middorsal scales abruptly larger than adjacent dorsolateral scales in A. amplisquamosus), from A. heteropholidotus, A. muralla, A. sminthus, and A. wermuthi in having strongly keeled ventral scales (ventral scales smooth or weakly keeled in A. heteropholidotus, A. sminthus, and A. wermuthi and smooth in A. muralla), and from A. anisolepis, A. crassulus, and A. haguei by having 8���11 rows of enlarged dorsal scales (usually 12���15 and as high as 23 in A. anisolepis, A. crassulus and A. haguei). This species can be distinguished from A. sp. (Townsend and Wilson, In press), and further differentiated from A. wermuthi, by having a hemipenis with an undivided asulcate processus (asulcate processus divided in A. sp. and A. wermuthi). Definition. A medium sized (snout-vent length [SVL] 38.1���47.5 mm in two males, 40.2���51.6 mm in two females; tail length [TL] 84���120.5 mm in two males, 87���113 mm in two females; SVL/TL 2.16���2.54 in two males, 2.19���2.21 in two females) member of the Anolis crassulus species group with a laterally compressed tail (1.11���1.56 times as high as wide), head length/head width 1.36���1.66, head length/SVL 0.28���0.29, relative leg length 3���5, tibia length/SVL 0.23���0.26. Scales on snout keeled; 4���7 scales between internasals; canthal ridges well-defined, composed of 3���4 canthal scales, with 4���6 scales between second canthals; 20���25 weakly keeled to keeled loreal scales present in a maximum of 4���5 oblique rows; 6 ��� 6 supralabials to below center of eye; suboculars contacting supralabials; supraorbital semicircles well-developed, with 0���1 scales separating supraorbital semicircles at narrowest point; interparietal scale well-developed, larger than surrounding scales, separated from supraorbital semicircles by 2���3 scales at the most proximate point; mental wider than long, partially divided by a medial suture; 4���6 postmentals, outermost pair the largest; 6 ��� 6 / 7 infralabials to below center of eye; keeled granular scales on chin and throat; males have a medium to large dewlap, and females have a small dewlap similarly colored to that of the male; greatest depth of extended male dewlap 5.5���7.9 mm; male dewlap length/SVL 0.358���0.36, female dewlap length/SVL 0.103���0.114; male dewlap extending 5.78���8.42 % of SVL beyond level of axilla, bearing 30���50 large gorgetal scales in 5���10 oblique rows; skin between gorgetals lacking scales; female dewlap does not extend beyond level of axilla, bearing 30���50 large gorgetal scales in 5���10 oblique rows; skin between gorgetals lacking scales; dorsal scales keeled, with 8���11 middorsal rows irregularly enlarged; 25���30 dorsal scales in one head length; lateral scales heterogeneous, with slightly enlarged keeled scales scattered among smaller granular scales; ventral scales keeled, imbricate, largest at midbody; 25���29 ventral scales in one head length; caudal scales strongly keeled; enlarged postanal scales present; axillary pocket absent; limb scales keeled, imbricate, largest on anterior surface of upper limbs; 24���27 lamellae under phalanges II to IV of fourth toe; 8���9 lamellae under distal phalanx of fourth toe. Hemipenis. The everted hemipenis of UF 152660, a subadult male, is a somewhat stout organ; asulcus processus undivided; sulcus spermaticus bounded by moderately well-developed sulcal lips, terminates at the base of the apex; truncus bearing some shallows folds, otherwise lacking surface structures; apical region appearing relatively smooth, slightly calyculate. Color in life. Coloration in life of UF 152660 (Fig. 2 a) was recorded as follows: dorsum rust brown on enlarged middorsal scale rows, smudged middorsally with dark gray; lateral region of body yellow-brown; anterior limbs yellow-brown; posterior limbs yellow-brown with narrow brown crossbars on lower limb; dorsum of head rust brown mottled with dark gray; tail yellow-brown with dark gray crossbars; venter peachcream; dewlap red with slight orange tinge (Fig. 2 b); iris rust brown. Coloration in life of UF 152661 was recorded as follows: dorsum uniform rust brown, lateral regions same; anterior limbs yellow-tan; posterior limbs rust brown; dorsum of head rust brown with dark gray smudging; tail rust brown; x-shaped dark brown mark at base of tail; venter pale peach with scattered black punctuations; small dewlap orangish red; iris rust brown. Coloration in life of UF 152662 (Fig. 3) was recorded as follows: enlarged middorsal scale rows gray brown with four dark brown chevrons; lateral regions yellow-gray with scattered black punctuations; anterior limbs yellow-tan; posterior limbs yellow-tan with brown crossbands on lower limb; dorsum of head graybrown; tail brown with slightly dark crossbands; venter yellowish cream; small dewlap orangish red; iris rust brown. Distribution and natural history. All known localities for Anolis rubribarbaris are in the Lower Montane Wet Forest formation, as defined by Holdridge (1967), in a relatively narrow elevational band between 1700 and 1740 m on the northern and eastern slopes of Monta��a de Santa B��rbara. According to the original field notes, the holotype (UF 90206) was collected on limestone boulders along a dry stream bed at 1700 m elevation, presumably during the daytime. UF 152660, a male, was collected at 2015 h while it slept on an orchid leaf growing approximately 3 m high on the trunk of a large tree at 1720 m elevation along a trail through disturbed cloud forest. UF 152661, a female, was asleep on an adjacent leaf of the same orchid directly under UF 152660. The large female, UF 152662, was collected further along the same trail in a patch of lightly disturbed cloud forest at 1740 m elevation, while asleep on a bracken fern (Pteridium sp.) approximately 1 m above the ground at 2045 h. Undisturbed cloud forest occurs at all points above these collection localities on Monta��a de Santa B��rbara, reaching a maximum elevation of 2744 m; however, herpetological survey work in the forests above 2000 m has not produced any additional material of A. rubribarbaris. Conservation status. Based on the extent of the remaining forest in Parque Nacional Monta��a de Santa B��rbara and the known distribution of Anolis rubribarbaris, this species would qualify as Endangered (EN B 2 ab[iii]) based on IUCN Red List Criteria (IUCN, 2001). This ranking assumes the distribution of A. rubribarbaris is not in fact limited to the 1700���1740 m elevational band described above and extends into the intact forest above this elevation. This species was consider to have a high degree of vulnerability to environmental degradation (EVS= 16) by Wilson and McCranie (2004), due to its occurrence in only a single forest formation, its status as a Honduran endemic, and low level of threat due to human persecution. FIGURE: 2 a) Subadult male Anolis rubribarbaris from above El Cedral, 1720 m elevation, Parque Nacional Santa B��rbara, Honduras; 2 b) dewlap of subadult male A. rubribarbaris shown in Fig. 2 a (UF 152660); 3) adult female Anolis rubribarbaris from above El Cedral, 1740 m elevation, Parque Nacional Santa B��rbara, Honduras (UF 152662)., Published as part of Townsend, Josiah H., Wilson, Larry David, Luque-Montes, Ileana R. & Ketzler, Lorraine P., 2008, Redescription of Anolis rubribarbaris (K��hler, McCranie, & Wilson 1999), a poorly-known Mesoamerican cloud forest anole (Squamata: Polychrotidae), pp. 39-44 in Zootaxa 1918 on pages 40-43, DOI: 10.5281/zenodo.274572, {"references":["Kohler, G., McCranie, J. R. & Wilson, L. D. (1999) Two new species of anoles of the Norops crassulus group from Honduras (Reptilia: Sauria: Polychrotidae). Amphibia-Reptilia, 20, 279 - 298.","Holdridge, L. R. (1967) Life Zone Ecology. Revised edition. Tropical Science Center, San Jose, Costa Rica. 206 p.","IUCN (2001) IUCN Red List Categories and Criteria: Version 3.1. IUCN Species Survival Commission, IUCN, Gland, Switzerland and Cambridge, United Kingdom. ii + 30 pp.","Wilson, L. D. & McCranie, J. R. (2004) The conservation status of the herpetofauna of Honduras. Amphibian and Reptile Conservation, 3, 6 - 33."]}

42. Geophis semidoliatus (Dum��ril, Bibron, and Dum��ril) Rabdosoma semi��doliatum Dum��ril, Bibron & Dum��ril 1854: 93. Catastoma semidoliatum: Cope 1860: 339. E (lapoides) semidoliatus: Jan 1862: 21. G (eophis) semidoliatus: Peters 1859: 276. Geophis (Parageophis) semidoliatus: Bocourt 1883: 534. Geophis semidoliata: G��nther 1893: 90. Atractus semidoliatus: Cope 1900: 1230. Syntypes. Originally five, Museum National d���Histoire Naturelle, Paris (MNHN) 3313 (1���3), 4522, fifth syntype apparently lost. Downs (1967: 133) designated MNHN 4522, an adult female, the lectotype. Type��locality. Mexico. Distribution. Central and southern Veracruz, Mexico, 500��1400 m elevation. Species group. semidoliatus (Downs 1967). Systematic references. Dum��ril et al. (1854), Downs (1967)., Published as part of Wilson, Larry David & Townsend, Josiah H., 2007, A checklist and key to the snakes of the genus Geophis (Squamata: Colubridae: Dipsadinae), with commentary on distribution and conservation, pp. 1-31 in Zootaxa 1395 on page 18, DOI: 10.5281/zenodo.273681, {"references":["Dumeril, A. M. C., Bibron, G. & Dumeril, A. H. A. (1854) Erpetologie Generale ou Histoire Naturelle Complete des Reptiles. Tome Septieme. - Premiere Partie. Comprenant l'Histoire Naturelle des Serpents non Venimeux. Librairie Encyclopedique Roret, Paris. vii + 1 - 780 pp.","Cope, E. D. (1860) Notes and descriptions of new and little known species of American reptiles. Proceedings of the Academy of Natural Sciences, Philadelphia, 12, 339 - 345.","Jan, G. (1862) Promodo dell Iconographie Generale degli Ofidi. Parte I. Calamaridae. Genoa, 76 pp.","Peters, W. (1859) Die von Hrn. Dr. Hoffmann in Costa Rica gesammelten Schlangen. Monatsberichte der Koniglichen Preuss. Akademie der Wissenschaften zu Berlin, 1859, 275 - 278.","Bocourt, F. M. (1883) In: Dumeril, A. H. A., Bocourt, M. F. & Mocquard, F. 1870 - 1909. Etudes sur les Reptiles. Mission Scientifique au Mexique et dans l'Amerique Centrale. Recherches Zoologiques pour servir a l'Histoire de la Fauna de l'Amerique Centrale et du Mexique. Troisieme Partie. - 1 re Section. Texte. Imprimerie Nationale, Paris. xiv + 1012 pp.","Gunther, A. C. L. F. (1893) Part 109, pages 89 - 96. In: Salvin, O. & Godman, F. D. (Eds.). Biologia Centrali-Americana; or, Contributions to the Knowledge of the Fauna and Flora of Mexico and Central America. R. H. Porter and Dulau & Co., London. xx + 326 pp. + 76 pls.","Cope, E. D. (1900) The crocodilians, lizards, and snakes of North America. Annual Report of the United States National Museum, 1898, 151 - 1294.","Downs, F. L. (1967) Intrageneric relationships among colubrid snakes of the genus Geophis Wagler. Miscellaneous Publications of the Museum of Zoology, University of Michigan, 131, 1 - 193."]}

37. Geophis rhodogaster (Cope) Colophrys rhodogaster Cope 1868: 130. Geophis rhodogaster: Bocourt 1883: 531. Geophis chalybaea: G��nther 1893: 87. Catastoma rhodogaster: Amaral 1929: 192. Geophis fulvoguttatus: K��hler 1996: 36 (in error). Syntypes. Three: Academy of Natural Sciences of Philadelphia (ANSP) 3316 �� 17 and National Museum of Natural History (USNM) 12425, collected by Van Patten. Downs (1967: 92) designated ANSP 3317, an adult female, the lectotype. Type��locality. ������elevated country in the neighborhood of the city of Guatemala.��� Distribution. The mountains of southwestern Chiapas, Mexico, east through southern Guatemala to the Cordillera Metap��n, Santa Ana, El Salvador, 1500��2744 m elevation. Species group. dubius (Downs 1967). Systematic references. Cope (1868), Downs (1967), Smith (1995), Townsend (2006). Remarks. G��nther (1893) reported G. rhodogaster from Costa Rica. Boulenger (1894) apparently followed G��nther (1893) and included Costa Rica in the range of G. rhodogaster. Taylor (1951) also included G. rhodogaster in his review of Costa Rica snakes, but noted ���I regard this record as somewhat doubtful.��� K��hler (1996) gave measurements and scutellation data for a specimen of G. rhodogaster (SMF 77413) under the name G. fulvoguttatus., Published as part of Wilson, Larry David & Townsend, Josiah H., 2007, A checklist and key to the snakes of the genus Geophis (Squamata: Colubridae: Dipsadinae), with commentary on distribution and conservation, pp. 1-31 in Zootaxa 1395 on pages 16-17, DOI: 10.5281/zenodo.273681, {"references":["Cope, E. D. (1868) An examination of the reptilia and batrachia obtained by the Orton expedition to Ecuador; and the upper Amazon; with notes on other species. Proceedings of the Academy of Natural Sciences, Philadelphia, 20, 96 - 140.","Bocourt, F. M. (1883) In: Dumeril, A. H. A., Bocourt, M. F. & Mocquard, F. 1870 - 1909. Etudes sur les Reptiles. Mission Scientifique au Mexique et dans l'Amerique Centrale. Recherches Zoologiques pour servir a l'Histoire de la Fauna de l'Amerique Centrale et du Mexique. Troisieme Partie. - 1 re Section. Texte. Imprimerie Nationale, Paris. xiv + 1012 pp.","Gunther, A. C. L. F. (1893) Part 109, pages 89 - 96. In: Salvin, O. & Godman, F. D. (Eds.). Biologia Centrali-Americana; or, Contributions to the Knowledge of the Fauna and Flora of Mexico and Central America. R. H. Porter and Dulau & Co., London. xx + 326 pp. + 76 pls.","Amaral, A. d. (1929) Estudos sobre ophidios neotropicos XVIII. Lista remissive dos ophidios da regiao neotropico. Memorias Do Instituto Butantan, 4, 129 - 271.","Kohler, G. (1996) Notes on a collection of reptiles from El Salvador collected between 1951 - 1956. Senckenbergiana Biologica, 76, 29 - 38.","Downs, F. L. (1967) Intrageneric relationships among colubrid snakes of the genus Geophis Wagler. Miscellaneous Publications of the Museum of Zoology, University of Michigan, 131, 1 - 193.","Smith, E. N. (1995) Geophis rhodogaster (Colubridae), an addition to the snake fauna of Mexico. Southwestern Naturalist, 40, 123 - 124.","Boulenger, G. A. (1894) Catalogue of snakes in the British Museum (Natural History). London, British Museum 2, i - xi, 1 - 382, 20 pls.","Taylor, E. H. (1951) A brief review of the snakes of Costa Rica. University of Kansas Science Bulletin, 34, 1 - 188."]}

24. Geophis juliai P��rez��Higareda, Smith, & L��pez��Luna Geophis juliai P��rez��Higareda, Smith & L��pez��Luna 2001: 42. Geophis semidoliatus: P��rez��Higareda et al. 1987: 16. Holotype. Estaci��n de Biolog��a Tropical Los Tuxtlas (UNAM��LT) 2775, adult male collected 10 August 1984. Type��locality. Estaci��n de Biolog��a Tropical Los Tuxtlas, 150 m elevation, Veracruz, Mexico. Distribution. Known only from mesic forest in the Los Tuxtlas region of southern Veracruz, Mexico, 150�� 600 m elevation. Species group. semidoliatus (P��rez��Higareda et al. 2001). Systematic references. P��rez��Higareda et al. (2001)., Published as part of Wilson, Larry David & Townsend, Josiah H., 2007, A checklist and key to the snakes of the genus Geophis (Squamata: Colubridae: Dipsadinae), with commentary on distribution and conservation, pp. 1-31 in Zootaxa 1395 on page 12, DOI: 10.5281/zenodo.273681, {"references":["Perez-Higareda, G., Smith, H. M. & Lopez-Luna, M. A. (2001) A new Geophis (Reptilia: Serpentes) from southern Veracruz, Mexico. Bulletin of the Maryland Herpetological Society 37, 42 - 48.","Perez-Higareda, G., Vogt, R. C. & Flores-Villela, O. (1987) Lista Anotada de los Anfibios y Reptiles de la Region de Las Tuxtlas, Veracruz. Instituto de Biologia, Universidad Nacional Autonoma de Mexico. 23 pp."]}

10. Geophis championi Boulenger Geophis championi Boulenger 1894: 321. Geophis chalybaea: G��nther 1893: 87. Catastoma championi: Amaral 1929: 191. Geophis brachycephalus: Dunn 1942: 4. Holotype. The Natural History Museum, London (BMNH) 1946.1.1.77, collected by J.G. Champion and F.D. Godman. Type��locality. Chiriqui, Panama. Distribution. The type locality simply lists ���Chiriqui, Panama ���, but Downs (1967) reported an additional specimen (CAS 78977) from Boquete, 1370 m elevation, eastern slope of Volcan Chiriqui, Chiriqui, Panama. Species group. championi (Downs 1967). Systematic references. Boulenger (1894), Dunn (1942), Downs (1967)., Published as part of Wilson, Larry David & Townsend, Josiah H., 2007, A checklist and key to the snakes of the genus Geophis (Squamata: Colubridae: Dipsadinae), with commentary on distribution and conservation, pp. 1-31 in Zootaxa 1395 on page 8, DOI: 10.5281/zenodo.273681, {"references":["Boulenger, G. A. (1894) Catalogue of snakes in the British Museum (Natural History). London, British Museum 2, i - xi, 1 - 382, 20 pls.","Gunther, A. C. L. F. (1893) Part 109, pages 89 - 96. In: Salvin, O. & Godman, F. D. (Eds.). Biologia Centrali-Americana; or, Contributions to the Knowledge of the Fauna and Flora of Mexico and Central America. R. H. Porter and Dulau & Co., London. xx + 326 pp. + 76 pls.","Amaral, A. d. (1929) Estudos sobre ophidios neotropicos XVIII. Lista remissive dos ophidios da regiao neotropico. Memorias Do Instituto Butantan, 4, 129 - 271.","Dunn, E. R. (1942) New and noteworthy snakes from Panama. Notulae Naturae, 108, 1 - 8.","Downs, F. L. (1967) Intrageneric relationships among colubrid snakes of the genus Geophis Wagler. Miscellaneous Publications of the Museum of Zoology, University of Michigan, 131, 1 - 193."]}

1. Geophis anocularis Dunn Geophis anocularis Dunn 1920: 127. Catostoma anoculare: Amaral 1929: 190. Geophis dubius: Downs 1967: 84. Holotype. National Museum of Natural History (USNM) 46556, adult male collected 26 July 1894 by E. W. Nelson and E. A. Goldman. Type��locality. Totontepec, Oaxaca, Mexico. Distribution. Known only from 1850��1900 m elevation in the vicinity of the type locality in the Sierra Mije, Oaxaca, Mexico. Species group. dubius (Downs 1967; Campbell et al. 1983). Systematic references. Dunn (1920), Downs (1967), Campbell et al. (1983). Remarks. Synonymized with G. dubius by Downs (1967: 84). Resurrected by Campbell et al. (1983), based on the collection of 21 additional specimens from Totontepec, Oaxaca, Mexico., Published as part of Wilson, Larry David & Townsend, Josiah H., 2007, A checklist and key to the snakes of the genus Geophis (Squamata: Colubridae: Dipsadinae), with commentary on distribution and conservation, pp. 1-31 in Zootaxa 1395 on pages 4-5, DOI: 10.5281/zenodo.273681, {"references":["Dunn, E. R. (1920) A new Geophis from Mexico. Proceedings of the Biological Society of Washington, 33, 27 - 28.","Amaral, A. d. (1929) Estudos sobre ophidios neotropicos XVIII. Lista remissive dos ophidios da regiao neotropico. Memorias Do Instituto Butantan, 4, 129 - 271.","Downs, F. L. (1967) Intrageneric relationships among colubrid snakes of the genus Geophis Wagler. Miscellaneous Publications of the Museum of Zoology, University of Michigan, 131, 1 - 193.","Campbell, J. A., Ford, L. S. & Karges, J. P. (1983) Resurrection of Geophis anocularis Dunn with comments on its relationships and natural history. Transactions of the Kansas Academy of Sciences, 86, 38 - 47."]}

31. Geophis nephodrymus Townsend & Wilson Geophis nephodrymus Townsend & Wilson, 2006: 151. Holotype. Florida Museum of Natural History (UF) 142577, a female collected 11 July 2004 by S. M. Hughes and J. H. Townsend. Type��locality. Sendero Las Minas (15 �� 29.525 ���N, 88 �� 12.705 ���W), 1580 m elevation, Parque Nacional El Cusuco, Cort��s, Honduras. Distribution. Known only from the vicinity of the type locality in the Sierra de Omoa, Cort��s, Honduras. Species group. dubius (Townsend & Wilson 2006). Systematic references. Townsend (2006), Townsend & Wilson (2006)., Published as part of Wilson, Larry David & Townsend, Josiah H., 2007, A checklist and key to the snakes of the genus Geophis (Squamata: Colubridae: Dipsadinae), with commentary on distribution and conservation, pp. 1-31 in Zootaxa 1395 on page 14, DOI: 10.5281/zenodo.273681, {"references":["Townsend, J. H. & Wilson, L. D. (2006) A new species of snake of the Geophis dubius group (Reptilia: Squamata: Colubridae) from the Sierra de Omoa of northwestern Honduras. Proceedings of the Biological Society of Washington, 119, 150 - 159."]}