Tantilla planiceps (Blainville, 1835). This species is the nominotypical member of the Tantilla planiceps group. The westernmost member of this group, its distribution extends from southern California, United States, to the southernmost tip of the Baja California Peninsula, in Mexico. Pictured here is an individual found in the Anza-Borrego Desert State Park in San Diego County, California, United States, which is located in the Colorado Desert ecosystem of the Sonoran Desert Ecoregion. The snake was found crawling on the ground at night in an extremely rocky area in desert scrub vegetation that included brittle bush, ocotillo, and barrel cactus. We determined its EVS as 9, placing it ' at the upper end of the low vulnerability category. Its IUCN status is of Least Concern. © Jack Goldfarb (www.JackGoldfarb.com) 441177 www.mesoamericanherpetology.com www.eaglemountainpublishing.com A checklist and key to the snakes of the Tantilla clade (Squamata: Colubridae), with comments on taxonomy, distribution, and conservation Larry DaviD WiLson1 anD vicente Mata-siLva2 1Centro Zamorano de Biodiversidad, Escuela Agrícola Panamericana Zamorano, Departamento de Francisco Morazán, Honduras. E-mail: [email protected] (Corresponding author) 2Department of Biological Sciences, The University of Texas at El Paso, El Paso, Texas 79968-0500, United States. E-mail: [email protected] abstract: We provide a checklist and identification key for the 66 members of the Tantilla clade, as defined by Holm (2008), which consists of the genera Geagras (one species), Scolecophis (one species), Tantilla (61 species), and Tantillita (three species). We provide summary information for these genera, and the species accounts contain information on a basic synonymy, the holotype, the type locality, geographic distribution, the assigned species group, critical references, and remarks. Comments on color pattern com- ponents in members of the Tantilla clade precede an identification key, which we largely based on color pattern and scutellation. We discuss the distribution of clade members with respect to country, physiog- raphy, and vegetation, and also examine their conservation status. We provide illustrations in life for 39 members of the clade, the largest number ever accumulated in a single source. Key Words: Endemism, EVS and IUCN categorizations, Geagras, physiographic and vegetational distri- bution, Scolecophis, Tantilla, Tantillita resuMen: Proporcionamos una lista y una clave de identificación para los 66 miembros del clado Tantilla, como está definido por Holm (2008), y el cual consiste de los géneros Geagras (una especie), Scolecophis (una especie), Tantilla (61 especies) y Tantillita (tres especies). Incluimos un resumen de información para cada uno de estos géneros, y la ficha para cada especie en la lista proporciona información básica sobre sinonimia, holotipo, localidad tipo, distribución geográfica, grupo de especie asignado, referencias más relevantes y comentarios pertinentes. Comentarios sobre los componentes del patrón de coloración en los miembros del clado Tantilla preceden una clave de identificación, la cual está basada ampliamente en el patrón de coloración y escutelación. Discutimos la distribución de los miembros del clado con respecto al país, fisiografía y vegetación, y también examinamos el estatus de conservación. Proporcionamos ilustra- ciones en vida de 39 miembros del clado, el número más grande hasta ahora acumulado en una sola fuente. Palabras Claves: Categorizaciones de EVS y UICN, distribución fisiográfica y vegetacional, endemismo, Geagras, Scolecophis, Tantilla, Tantillita Citation: Wilson, L. D., and V. Mata-Silva. 2015. A checklist and key to the snakes of the Tantilla clade (Squamata: Colubridae), with comments on taxonomy, distribution, and conservation. Mesoamerican Herpetology 2: 418–498. Copyright: Wilson and Mata-Silva, 2015. This work is licensed under a Creative Commons Attribution-NoDerivatives 4.0 International License. Received: 14 October 2015; Accepted: 23 November 2015; Published: 26 December 2015. Mesoamerican Herpetology 418 December 2015 | Volume 2 | Number 4 Wilson and Mata-Silva Tantilla clade checklist and key DEDICATIONS To my wife, Elizabeth To my dear mother, Elena (Betty) Pineda Wilson Beatriz Silva Sandoval (1946–2015), who bat- (1935–2007), who always tled an implacable en- supported my siblings and emy to the end, an incur- me in our search for knowl- able disease, and who I edge and our efforts to be- know would agree that come better people. She the best thing in our always was interested and 47-year marriage was mystified to see me study- to give life to our two ing a group of animals that wonderful daughters, Robin Lee and Tayra Barbara. were not the usual cows and horses. – Larry David Wilson – Vicente Mata-Silva INTRODUCTION In an unpublished dissertation, Holm (2008) defined the Tantilla clade as comprised of an expanded Tantilla plus its putative sister genus Scolecophis. Based on an analysis of morphological data, Holm’s expanded concept of the ge- nus Tantilla included the nominal genera Geagras and Tantillita, and he recognized the Tantilla clade as one of three (the other two are the Sonora and Ficimia clades) in the tribe Sonorini within the family Colubridae. Unfortunately, these important taxonomic decisions likely will not be published (P. Holm, pers. comm.). Thus, in updating the taxonomy of the genus Tantilla documented in Wilson (1999), herein we decided to include the other generic-level members of the Tantilla clade and maintained the traditional taxonomy for these other members. The Tantilla Clade Following the conclusions of Holm (2008) on the composition of “the Tantilla clade,” we include the nominal genera Geagras (one species), Scolecophis (one species), Tantilla (61 species, including the poorly understood T. trilineata), and Tantillita (three species) in this clade, for a total of 66 members. Herein, we maintain the usual arrangement of these four genera, until Holm’s (2008) conclusions can be tested through the use of molecular tech- niques. Nonetheless, we predict that Holm’s taxonomy will be upheld whenever this work is completed. The Genus Geagras Cope (1876) described the genus Geagras and its single member, G. redimitus, based on a holotype from the “west side of the State of Tehuantepec, Mexico” (see below). Wilson (1987d: 430.1) defined Geagras as “a colubrid ge- nus characterized by: slender body with head not noticeably distinct from body; head cuneiform, head scutellation accordingly modified and simplified, consisting of a cuneiform rostral, two internasals and prefrontals, a broad scutiform frontal, two parietals, divided nasal, no loreal, single tiny preocular widely separated from postnasal, sin- gle postocular, temporals 1+1; five supralabials, with the third entering the orbit; six infralabials, with four touching anterior chin shields and fourth the largest; dorsal scales smooth, in 15 rows throughout; ventrals 113–124; anal plate divided; subcaudals 26–33, paired; maximum known total length 235 mm; maxillary teeth 10 (one count), separated by a short diastema from two distinctly enlarged grooved fangs; hemipenis simple [Smith, 1943, indicated the organ to be capitate, which appears unlikely] with single sulcus spermaticus, distal third calyculate, median third spinose, proximal third bare; dorsal color pattern of narrow diffuse dark lines or stripes coursing the length of all but two lower scale rows; head pattern of a dark median spatulate blotch bounded laterally by narrow pale markings which unite on snout, these in turn bounded below by a dark facial stripe.” This genus has been recognized in the herpetological literature since its description (see accounts at The Reptile Database and IUCN Red List websites). As indicated by Wilson (1987d: 430.1), “Smith [1943] noted the Mesoamerican Herpetology 419 December 2015 | Volume 2 | Number 4 Wilson and Mata-Silva Tantilla clade checklist and key relationship of Geagras to the large genus Tantilla. Wilson and Meyer [1981] supported this contention and, further- more, concluded that its relationships most closely lie with the members of the T. calamarina group, in particular the atypical T. calamarina. Wilson and Meyer [1981] noted the greater specialization of Geagras for fossorial life over the condition seen in Tantilla calamarina, itself a highly specialized member of its genus.” In the introduction to his dissertation on the phylogenetic biology of the snake tribe Sonorini, however, Holm (2008) stated that, “Wilson and Meyer (1981) suggested that Geagras redimitus is related to the Tantilla calamarina species group,” and proposed the question “Is Geagras nested within Tantilla?” Based on his analysis of morpho- logical features in the Sonorini, his answer was yes. His phylogenetic tree (his Fig. 4) indicated that G. redimitus is the sister species of T. calamarina, and these two taxa successively are linked to T. cascadae, T. coronadoi, and T. deppei. Holm (2008) did not consider T. sertula in his analysis. Apparently, the original description of T. ceboruca (Canseco-Márquez et al., 2007) was not available to him. Holm’s (2008) conclusions about the phylogeny of the T. calamarina group agree with those of Wilson and Meyer (1981) and Wilson and Campbell (2000). His conclusions about the relationship of G. redimitus to members of the T. calamarina group also agree with those of Wilson and Meyer (1981). He stated (2008: 97–98) that, “Wilson and Meyer (1981)…noted the similarity and probable relationship to the highly fossorial species Geagras redimi- tus. Wilson and Meyer (1981) also noted that the degree of fossoriality increased from T. deppei to T. calamarina, culminating with Geagras.” Ramírez-Bautista et al. (2014b) amplified these conclusions by constructing a phylogenetic hypothesis using data on scutellation for the seven species of Tantilla currently placed in the T. calamarina group, and the species G. redimitus, and used these data to develop an index of fossoriality. Based on the resulting application of this index, these authors (2014b: 803) concluded that “the arrangement of species, from least to more adapted to fossoriality, is as follows: T. sertula…T. deppei…T. ceboruca…T. coronadoi…T. cascadae…T. vermiformis…T. calamarina… and G. redimitus…” Nonetheless, they (2014: 804) urged caution in the application of their conclusions “to sup- port either of Holm’s (2008) contentions, i.e., that T. vermiformis is not a member of the calamarina group or that G. redimitus should be allocated to the genus Tantilla. Answering these two questions, as well as those relating to the phylogenetic relationships of the remainder of the species included in this discussion, will have to await the assembly and evaluation of sufficient molecular material to allow for a robust analysis.” Given the apparent rarity of most of these species, however, it might take considerable time to assemble the necessary molecular material to test these hypotheses. Geagras redimitus Cope Geagras redimitus Cope 1876: 141. Holotype: National Museum of Natural History (USNM) 30115, probably a juvenile or subadult male, collected on an unknown date by Dr. Francis Sumichrast. Type-locality: “West side of the State of Tehuantepec, Mexico” (= the Pacific side of the region of the Isthmus of Tehuantepec, Oaxaca, Mexico). Distribution: Low elevations (0–400 m) of the Plains of Tehuantepec on the Pacific versant of southeastern Oaxaca, Mexico. Casas-Andreu et al. (1996) gave an elevational range of 100–1,000 m, but did not indicate the source of these records. References: Cope (1876); Sumichrast (1882); Hartweg and Oliver (1940); Wilson and Meyer (1981); Wilson (1987d); Holm (2008); Wilson and Johnson (2010); Wilson and Townsend (2010); Wilson et al. (2013a); Ramírez- Bautista et al. (2014b); Wallach et al. (2014); Mata-Silva et al. (2015). Remarks: Wilson (1987d) corrected geographic distribution errors in this taxon that continue to be promul- gated (see The Reptile Database website for G. redimitus) with the following information: “Records of this species from Michoacán (Duellman, 1961) are based on Tantilla calamarina (Wilson and Meyer, 1981). The type-locality of Sphenocalamus lineolatus is Mazatlán. As noted by Hardy and McDiarmid (1969), the state was not indicated. They listed G. redimitus as part of the Sinaloan herpetofauna, in part based on Duellman’s (1961) misidentifica- tion of specimens from the supposed intermediate localities in Michoacán. An alternative possibility is that the holotype of S. lineolatus came from Mazatlán (= San Juan Mazatlán), Oaxaca, a village located about 76 km N Mesoamerican Herpetology 420 December 2015 | Volume 2 | Number 4 Wilson and Mata-Silva Tantilla clade checklist and key of Tehuantepec and 46 km WNW of Matías Romero at an approximate elevation of 400 m, lying in the lowlands of the Isthmus of Tehuantepec.” Google Earth shows this village at coordinates (17°01'38.79"N, 95°26'23.45"W) at an elevation of approximately 1,746 feet (= 532 m), 81 km NNW of Tehuantepec and 46 km WNW of Matías Romero. The locality mentioned by Hartweg and Oliver (1940) for this species, “Quiengola Mountain,” presumably refers to the hill (Cerro Quiengola) on which the Zapotec archeological site is located, 14 km N of Tehuantepec (16°23'09"N, 95°19'24"W; www.wikipedia.org). Duellman (1960) indicated the location of this hill as 15 km WNW of Tehuantepec, with “dense scrub forest” on its slopes. The Genus Scolecophis Fitzinger (1843) described the genus Scolecophis to contain the single species Calamarina atrocincta Schlegel, 1837. Schlegel (1837) reported that the holotype of this species was collected in Chile, an obvious error. Wilson and Williams (2002: 758.1) defined Scolecophis as follows: “a colubrid genus characterized by a slender body; a head only slightly distinct from the body; head shape normal; dorsal head scutellation consisting of a normal rostral followed by two internasals, two prefrontals, two supraoculars, one frontal, and two parietals; lateral head scutellation of a divided nasal, a single loreal, one preocular, two postoculars, no suboculars, 1+1+1 temporals; seven supralabials, with the 3rd and 4th entering the orbit; six or seven infralabials, with three or four touching the anterior chin shields, the fourth the largest, and the first pair in medial contact, separating the mental and ante- rior chinshields; dorsal scales are smooth and in 15 rows; ventrals 181–198; cloacal scute (= anal plate) divided; subcaudals 45–54 and paired; maximum known TL [= total length] about 470 mm (KU 125497 from Costa Rica); relative tail length 0.136–0.191; maxillary teeth 13–14, the two posterior teeth grooved laterally, the grooves broad and shallow; hemipenis simple with single sulcus spermaticus, distal two-thirds spinose, spines increasing in size proximally, terminating in several enlarged basal spines, and the proximal one-third of the organ naked; a body pat- tern of alternating black and pale rings, the black rings 1½–4 scales long, the pale rings 2–4 scales long, pale rings are cream-colored on the first 4–5 scale rows, each scale with a black tip (except in the first row), reddish orange on dorsal rows 5–11 or 6–10, black rings number 24–49 on body and 5–10 on tail; head pattern of a black head cap with a broad pale band across the prefrontals extending laterally to the lip line, a postocular pale spot, and a pale nuchal band (cream laterally, reddish orange middorsally) beginning on the posterior portion of the parietals and extending 1–2½ middorsals beyond.” Scolecophis has been recognized in the herpetological literature since its description by Fitzinger (1843), other than from 1854 to 1866 when it was relegated either to the synonymy of Homalocranion, a synonym of Tantilla (Wilson, 1982a), or placed in the genus Platycranion, erected by Jan (1863) to contain the same species by monotypy as contained in Scolecophis (i.e., Calamaria atrocincta Schlegel, 1837). European herpetologists (e.g., G. A. Boulenger, A. Günther, and G. Jan), however, essentially used the genus Homalocranion (or Homalocranium) for species placed in Tantilla by American herpetologists of the same era (e.g., E. D. Cope). In his phylogenetic analysis, Holm (2008) placed Scolecophis as the sister genus to his “expanded” Tantilla (including Geagras and Tantillita). He stated (p. 95) that “two characters support monophyly of the expanded Tantilla with respect to its sister taxon Scolecophis. These are absence of apical pits on dorsal scales and absence or near absence of the loreals.” Scolecophis exhibits a single apical pit and a single loreal separating the posterior nasal and the single preocular (McCranie, 2011a). Scolecophis atrocinctus Schlegel Calamaria atrocincta Schlegel, 1837: 47. Scolecophis atrocincta: Fitzinger, 1843: 25. Elaps zonatus Hallowell, 1855: 35. Scolecophis atrocinctus: Günther, 1895: 156. Holotype: Muséum National d'Histoire Naturelle, Paris (MNHN) 519, adult male, collected on an unknown date by Gay. Type-locality: “Chile,” in error. Mesoamerican Herpetology 421 December 2015 | Volume 2 | Number 4 Wilson and Mata-Silva Tantilla clade checklist and key Distribution: Low, moderate, and intermediate elevations (100–1,530 m) on the Pacific versant from south- eastern Guatemala to northwestern Costa Rica; the species also is found on the Atlantic versant in southwestern Honduras, western Nicaragua, and northwestern Costa Rica. References: Savage (2002); Wilson and Williams (2002); Köhler (2008); Acevedo et al. (2010); Greenbaum and Komar (2010); Sasa et al. (2010); Sunyer and Köhler (2010); Wilson and Johnson (2010); Wilson and Townsend (2010); McCranie (2011a); Wallach et al. (2014); Johnson et al. (2015b). Remarks: One of the many phylogenetic issues of interest in the systematics of members of the Tantilla clade is the relationship of the banded species to one another. In addition to S. atrocinctus, the single species allocated to the monospecific genus Scolecophis, considered by Holm (2008) to be the sister genus of his expanded concept of Tantilla (see section above), three species of Tantilla also have a banded pattern: T. semicincta, T. shawi, and T. supracincta. These three species and S. atrocinctus occupy non-overlapping ranges from northeastern Mexico (T. shawi), through largely Pacific versant portions of Central America from southeastern Guatemala to northwestern Costa Rica (S. atrocinctus), and the Atlantic versant from southeastern Nicaragua to central Panama, as well as the Pacific versant of Costa Rica, Panama, and Ecuador (T. supracincta), to the Caribbean coastal portions of Colombia and Venezuela (T. semicincta). Beyond the presence of dark bands on the dorsum of the body, the patterns of these four species show scant resemblance to one another. Tantilla shawi and T. supracincta generally show some band- ing on the anterior portion of the body, whereas T. semicincta and S. atrocinctus are banded along the length of the body. In addition to a banded pattern, some individuals of T. semicincta also are boldly striped. Given that Holm (2008) placed Scolecophis as a sister genus to Tantilla (including both Geagras and Tantillita), he did not envision a close relationship between Scolecophis and any of the banded species of Tantilla. His view of the relationship of Scolecophis to Tantilla likely will be borne out by future molecular research. ' Adult Scolecophis atrocinctus from Tilarán, Guanacaste, Costa Rica. EVS 14 (high vulnerability species). © Fabio Hidalgo Mesoamerican Herpetology 422 December 2015 | Volume 2 | Number 4 Wilson and Mata-Silva Tantilla clade checklist and key The Genus Tantilla Wilson (1999) provided the last taxonomic summary of the entire genus Tantilla, in which he listed 53 species. A number of changes have occurred in the interim, which we indicate below. Greenbaum et al. (2004) synonymized Tantilla equatoriana Wilson and Mena, 1980, with T. melanocephala (Linnaeus, 1758). The following two species have been resurrected: Tantilla armillata Cope, 1876, from the synonymy of T. melanocephala (Linnaeus, 1758) by Savage (2002). Tantilla ruficeps Cope, 1894, from the synonymy of T. melanocephala (Linnaeus, 1758) by Savage (2002). The following seven species have been described as new (country of origin in parentheses): Tantilla boipiranga Sawaya and Sazima, 2003 (Brazil) Tantilla ceboruca Canseco-Márquez, Smith, Flores-Villela, and Campbell, 2007 (Mexico) Tantilla hendersoni Stafford, 2004 (Belize) Tantilla olympia Townsend, Wilson, Medina-Flores, and Herrera-B., 2013 (Honduras) Tantilla psittaca McCranie, 2011b (Honduras) Tantilla robusta Canseco-Márquez, Mendelson, and Gutiérrez-Mayén, 2002 (Mexico) Tantilla sertula Wilson and Campbell, 2000 (Mexico) In summary, of the 53 species listed by Wilson (1999), 52 presently are regarded as valid members of the genus Tantilla; the difference is due to the synonymization of T. equatoriana with T. melanocephala. We do not recognize T. marcovani, which was described after the Wilson (1999) paper appeared, because we consider it insep- arable from T. melanocephala. Two taxa, T. armillata and T. ruficeps, subsequently were resurrected. In addition, seven new taxa were described. Thus, 61 species currently are included in the genus. 1. Tantilla albiceps Barbour Tantilla albiceps Barbour, 1925: 156. Holotype: Museum of Comparative Zoology, Harvard University (MCZ) 20600, adult (?) male, collected in February of 1925 by Thomas Barbour. Type-locality: Barro Colorado Island, elevation 60 m, Gatún Lake, Atlantic versant of the Canal Zone, Panama. Distribution: Known only from the type locality, an island created with the construction of the Panama Canal, at a low elevation on the Atlantic versant of Panama. Species group: Tantilla albiceps is one of 14 species that has not been placed in any of the phenetic groups recognized in the genus (Wilson and Mata-Silva, 2014). This situation likely will not improve because this species is known only from the holotype and has not been seen again for 90 years. Holm (2008) indicated T. albiceps as the sister species of T. nigra. Interestingly, each of these taxa is known from a single specimen. References: Wilson (1982b, 1985a, 1999); Joseph-Ouni (2007); Köhler (2008); Jaramillo et al. (2010); Wallach et al. (2014); Johnson et al. (2015b); Ray and Ruback (2015). Remarks: Jaramillo et al. (2010) opined that, “this snake possibly is extinct, especially since the herpeto- fauna of this island [Barro Colorado] has been well sampled (Myers and Rand, 1969; Rand and Meyers, 1990).” Apparently, it has not been found since its discovery when “an Indian boy engaged in digging out a stump near the laboratory building came upon [the] small snake when the author was visiting the research station (Barbour 1925: 156). Joseph-Ouni (2007) considered this species extinct, indicating that “the reason for its extinction is not certain, but probably tied to the industrial and commercial development of the region.” Also see Remarks section of T. moesta account. Tantilla albiceps is one of several species in the genus that remains known only from the holotype. This group of species includes (with the number of years past the original description indicated in parentheses): T. albiceps (90); T. andinista (35); T. briggsi (44); T. hendersoni (11); T. johnsoni (16); T. miyatai (28); T. nigra (101); T. olympia (2); T. petersi (36); T. robusta (13); T. tecta (18); and T. trilineata (135). These 12 species comprise 19.7% of the 61 species of Tantilla. Mesoamerican Herpetology 423 December 2015 | Volume 2 | Number 4 Wilson and Mata-Silva Tantilla clade checklist and key 2. Tantilla alticola (Boulenger) Homalocranium alticola Boulenger, 1903: 353. Homalocranium coralliventre Boulenger, 1913: 1,035. Tantilla costaricensis Taylor, 1954: 766. Tantilla alticola: Amaral, “1929” (1930): 219. Syntypes: British Museum of Natural History (BMNH) 1946.1.8.63–65 (formerly BMNH 98.10.27.7–9), first two males, third apparently a female, collected by A. E. Pratt (date of collection unknown; accessioned 27 October 1898). Type-locality: Santa Rita, north of Medellín, 9,000 ft (= 2,743 m), Departamento de Antioquia, Colombia. Distribution: Low to intermediate elevations (91–2,743 m) on the Atlantic versant of Nicaragua and Costa Rica, and on the Pacific versant of southwestern Costa Rica and western Panama, as well as western Colombia. Species group: This species is one of 14 that remain unallocated to a phenetic group (Wilson and Mata-Silva, 2014). References: Wilson et al. (1977); Wilson (1982b, 1986, 1987a, 1999); Savage (2002); Castaño-M. et al. (2004); Mueses-Cisneros and Cepeda-Quilindo (2006); Köhler (2008); Jaramillo et al. (2010); Sasa et al. (2010); Sunyer and Köhler (2010); Wilson and Johnson (2010); Myers et al. (2013); Sunyer (2014); Wallach et al. (2014); Johnson et al. (2015b); Vanegas Guerrero et al. (2015); Vera–Pérez et al. (2015). Remarks: Vanegas Guerrero et al. (2015) recently reported a specimen of T. alticola that filled a distribution gap of 455 km in western Colombia. In doing so, they reviewed and mapped the known distribution of this species, which extends from west-central Nicaragua southward through Costa Rica and Panama to southwestern Colombia near the border with Ecuador. These authors, however, indicated that T. alticola can be distinguished from T. mela- nocephala, a species widely distributed in Colombia, on the basis of differences in the number of infralabial scales, in addition to color pattern. They indicated the number of infralabials as six in T. alticola and seven in T. melano- cephala, and cited Wilson and Mena (1980) to support the latter assertion. Wilson and Mena (1980: 15), however, noted that, “the majority of the specimens [of T. melanocephala] examined (92.8%) have 6 infralabials,” the typical condition for members of the genus Tantilla (Wilson, 1982a). Vanegas et al. (2015) also maintained that T. alticola has the broadest elevational range (i.e., 91–2,743 m), based on information provided by Uetz and Hošek (2014). Wilson and Mena (1980), however, documented that T. melanocephala, as they envisioned this taxon, is found from sea level to 2,851 m in the intermontane basin of Quito in Ecuador. This range is the same as presently understood, Tantilla alticola from the Municipio de Dagua, Departamento de Valle del Cauca, Colombia. EVS 11 (medium vulnerability species). ' © Fernando Vargas-Salinas Mesoamerican Herpetology 424 December 2015 | Volume 2 | Number 4 Wilson and Mata-Silva Tantilla clade checklist and key even given that T. armillata and T. ruficeps were recovered from synonymy of T. melanocephala by Savage (2002) and T. equatoriana was placed in synonymy by Greenbaum et al. (2004). Also see T. jani account concerning the status of the paralectotype of Homalocranium jani Günther (1895) from Matagalpa, Nicaragua. 3. Tantilla andinista Wilson and Mena Tantilla andinista Wilson and Mena, 1980: 21. Holotype: University of Kansas Museum of Natural History (KU) 135209, adult female, collected on 9 June 1970 by Thomas H. Fritts. Type-locality: 5 km E of Alausí, elevation 2,600–2,750 m, Provincia Chimborazo, Ecuador. Distribution: Known only from the type locality, at intermediate and high elevations on the continental divide in Ecuador. Species group: melanocephala (Wilson and Mena, 1980). References: Wilson and Mena (1980); Wilson (1985b, c, 1987a, 1999); Wallach et al. (2014). Remarks: This species was described 35 years ago, but remains known only from the holotype. See Remarks section of T. albiceps account. 4. Tantilla armillata Cope Tantilla armillata Cope, “1875” (1876): 143. Holotype: Academy of Natural Sciences of Philadelphia 336, collected within period of 1865–1875 by C. H. Van Patten. Type-locality: “Middle Costa Rica.” Distribution: Low to moderate elevations (0–1,435 m) on the Pacific versant from Guatemala to northwestern Costa Rica and perhaps western Panama (Knight et al., 2012); also occurs on the Atlantic versant in central and northeastern Honduras, southwestern Nicaragua, and central Costa Rica. Species group: melanocephala (Wilson and Mena, 1980; Savage, 2002). Tantilla armillata from Parque Nacional Volcán Masaya, Nicaragaua. EVS 11 (medium vulnerability species). ' © José Gabriel Martínez-Fonseca Mesoamerican Herpetology 425 December 2015 | Volume 2 | Number 4 Wilson and Mata-Silva Tantilla clade checklist and key References: Hardy and Cole (1967); Wilson and Villa (1973); Wilson and Mena (1980); Wilson (1984 [as T. melanocephala], 1999); Savage (2002); Köhler (2008); Acevedo et al. (2010); Greenbaum and Komar (2010); Sasa et al. (2010); Sunyer and Köhler (2010, as T. melanocephala); Townsend and Wilson (2010); Wilson and Johnson (2010); McCranie (2011a); Solís et al. (2014); Sunyer (2014); Sunyer et al. (2014); Townsend (2014); Wallach et al. (2014); Acosta-Chaves and Villalobos-Chaves (2015); Johnson et al. (2015b); McCranie (2015); Ray and Ruback (2015). Remarks: Previously this taxon was recognized as a distinct species, but Wilson and Mena (1980) syn- onymized it with the widespread T. melanocephala. Savage (2002) argued, however, that it and T. ruficeps, also synonymized with T. melanocephala by Wilson and Mena (1980), should be resurrected from the synonymy of this taxon and recognized as distinct species, and we concur with this decision. Knight et al. (2012) noted the presence of two eggs in an individual allocated to this species from western Panama. Acosta-Chaves and Villalobos-Chaves (2015) reported the Central American Bark Scorpion (Centruroides edwardsii) preying on T. armillata. Sunyer et al. (2014) reported the first individuals of this species from within protected areas in Nicaragua. Tantilla armillata is the northernmost member of the melanocephala group. 5. Tantilla atriceps (Günther) Homalocranium atriceps Günther, 1895: 146. Tantilla atriceps: Amaral, 1930: 219. Syntypes: British Museum of Natural History (BMNH) 1946.1.8.81–82 (original numbers 89.7.3.36–37, respectively), both males, obtained in 1889 from W. Taylor. Type-locality: “Nuevo León, Mexico.” Distribution: “Low, moderate, and intermediate elevations [0–2,134 m] in southeastern Chihuahua, Coahuila, northwestern and southern Nuevo León, western San Luis Potosí, southern Durango, and northeastern Zacatecas, Mexico, with apparently disjunct populations in Tamaulipas, Mexico, and southern Texas, United States” (Wilson and Mata-Silva, 2014). Species group: planiceps (Cole and Hardy, 1981). References: Cole and Hardy (1981, 1983); Wilson (1999); Gotte and Wilson (2005); Lavín and Lazcano (2010); Wilson et al. (2013a); Wallach et al. (2014); Wilson and Mata-Silva (2014). ' Tantilla atriceps from Aramberri, Nuevo León, Mexico. EVS 11 (medium vulnerability species). © Timothy Burkhardt Mesoamerican Herpetology 426 December 2015 | Volume 2 | Number 4