Bolitoglossa alberchi
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publication ID |
https://doi.org/ 10.5281/zenodo.13259133 |
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persistent identifier |
https://treatment.plazi.org/id/AC57066A-FF80-C615-F5E5-745AFEDDFA7B |
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treatment provided by |
Felipe |
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scientific name |
Bolitoglossa alberchi |
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Bolitoglossa alberchi View in CoL *
Eleven of the 16 single-region SLT species (68.8%) are state endemics, three (18.8%) are country endemics, A Coefficient of Biogeographic Resemblance (CBR) matrix was constructed for assessing the herpetofaunal similarity relationships ( Duellman 1990) among the four physiographic regions in Veracruz ( Table 7) and those data were used to create a UPGMA dendrogram ( Fig. 7 View Fig ; Sokal and Michener 1958). The SMO contains the greatest amount of species richness (236 species) and the SLT the least (179 species). The mean species richness value for all four regions is 206.8. The number of shared species between each regional pair ranges from a high of 190 between the SMO and TVB, to a low of 100 between the TVB and GCL. The mean value of shared species among all four regions is 123.0. The lowest number of shared species between the TVB and GCL (100 species) was expected, as these two regions are completely separated from each other by the SMO and are environmentally different on an elevational scale. The GCL, with an elevational range from sea level to about 200 m, contains tropical evergreen forest, scrub, sub-deciduous forest, and tropical dry forest (CONABIO, 2008). Conversely, with a limited geographic area within Veracruz, the TVB contains primarily coniferous and oak forest vegetation with the remainder comprised of subalpine scrub, cloud forest, xerophilous scrub, and tropical dry forest. The elevations for the TVB range from 1,000 m in sloping river valleys to 5,700 m on the highest volcanic peak. The TVB and SMO share the most No. 25. Scincella cherriei (Cope, 1893) . The Brown Forest Skink is a non-endemic species occurring from Veracruz to western Panama ( Uetz et al. 2020). This individual was found at Los Tuxtlas, in the municipality of San Andrés Tuxtla. Wilson et al. (2013b) calculated its EVS as 8, placing it in the upper portion of the low vulnerability category. Its conservation status has not been assessed by either the IUCN or SEMARNAT. Photo by Christian Berriozabal-Islas.
No. 27. Sphaerodactylus glaucus (Cope, 1866) . The Collared Dwarf Gecko is non-endemic species occurring from Veracruz, Tabasco, Oaxaca, and Chiapas in Mexico, through the Yucatan Peninsula and northern Guatemala to the interior of western Honduras, at elevations from 200 to 1,000 m ( Köhler 2008). This individual was found at Los Tuxtlas, in the municipality of San Andrés Tuxtla. Wilson et al. (2013b) determined its EVS as 12, placing it in the medium vulnerability category. This species has not been assessed by either the IUCN or SEMARNAT. Photo by Christian Berriozabal-Islas.
No. 26. Lepidophyma tuxtlae Werler and Shannon, 1957 . The Tuxtla Tropical Night Lizard is a country endemic species distributed from the Sierra de Los Tuxtlas to the El Ocote region of Chiapas ( Köhler 2008). This individual was located at Los Tuxtlas, in the municipality of San Andrés Tuxtla. Wilson et al. (2013b) determined its EVS as 11, placing it in the middle of the medium vulnerability category. Its conservation status is designated as Data Deficient by the IUCN, and as Threatened (A) by SEMARNAT. Photo by Christian Berriozabal-Islas.
No. 28. Boa imperator Daudin, 1803 . The Central American Boa Constrictor is a non-endemic species occurring in Central America (including South American populations in the Choco of Colombia and Ecuador [and probably Peru], and North American populations along the Gulf coast of Mexico (west of the Isthmus of Tehuantepec; Card et al. 2016). This individual was encountered at Los Tuxtlas, in the municipality of San Andrés Tuxtla. We calculated its EVS as 10, placing it at the lower limit of the medium vulnerability category. Its conservation status has not been determined by the IUCN or SEMARNAT. Photo by Christian Berriozabal-Islas.
species (190), which also was expected because they are directly adjacent to each other in Veracruz and share many of the same montane environments in their limited geographic ranges within the state. The SLT and GCL share the second largest number of species (137). The SLT is an isolated mountainous region with a maximum elevation of 1,700 m, which is nearly surrounded by lowland habitats, some of which ascend upward to an elevation of 200 m into lower montane areas on volcanic slopes. Pairwise comparisons of the aligned regions in order from highest to lowest species richness (underlined values) and their corresponding numbers of shared species (in parentheses) are as follows:
SMO 236: TVB (190), SLT (104), GCL (105) TVB 222: SMO (190), SLT (102), GCL (100) GCL 190: SLT (137), SMO (105), TVB (100) SLT 179: GCL (137), SMO (104), TVB (102)
In general, the pattern indicates how species richness values within each of the four biogeographic regions of Veracruz equate to numbers of shared species among the other three regions. There is a higher correlation of species richness values to the numbers of shared species between regions that are in contact with each other, but also a correlation between regions that share ecological parameters. Interestingly, the two regions that share the second highest number of species (137) are a highland region (SLT) and a lowland region (GCL), which is probably due to the GCL containing many generalist species that tolerate both montane and non-montane environments at low to moderate elevations. The fact that the GCL shares fewer species with the SMO and TVB gives credibility to the premise that regions separated by ecological barriers will share fewer species than they will with regions in direct contact.
The following data show the ranges and mean numbers of shared species (bold in parentheses) for each of the four regions, and are arranged according to increasing species richness (underlined values) in each region:
Sierra Madre Oriental, SMO (236): 104–190 (133)
Transmexican Volcanic Belt, TVB (222): 100–190 (130.7)
Gulf Coastal Lowlands, GCL (190): 100–137 (114)
Sierra de Los Tuxtlas, SLT (179): 102–137 (114.3)
The mean numbers of shared species compared to the species richness in the four regions indicate that higher species richness in a pairwise comparison tends, with one exception, to translate into higher reciprocal numbers when all the regional pairs are totaled. The comparison between the SMO and TVB are 1 st and 2 nd in species richness and 1 st and 2 nd in the average value of shared species, respectively. The minor exception is that the GCL, a lowland region, is 3 rd in species richness but last (4 th) in the mean number of shared species, whereas the SLT is lower in species richness but slightly higher in the mean numbers of shared species. Apparently, the size of the region has an important effect on species richness, and the ecological variability (highlands vs. lowlands) has an important effect on the average number of shared species in Veracruz.
Regarding area, the GCL in Veracruz is by far larger than all three mountainous areas combined (SMO, TVB, and SLT), but is 3 rd in species richness; it has 11 more species than the SLT. The total area of the SLT is much smaller compared to the other three regions and it is located only within a small portion of southeastern Veracruz, whereas the other three regions have much larger ranges outside of the state.
Based on the data in Table 7, a UPGMA dendrogram ( Fig. 7 View Fig ) depicts herpetofaunal similarity resemblance patterns in a hierarchical fashion among the four physiographic regions of Veracruz ( Fig. 1 View Fig ). The dendrogram is composed of two distinct clusters: one comprising two montane regions (SMO and TVB) at the 0.83 level, and the other containing one montane region (SLT) and one lowland region (GCL) at the 0.74 level. The two clusters connect with each other at the 0.50 level. Regions in both clusters have portions adjacent to each other somewhere in their total ranges, and the GCL surrounds the SLT to a varying degree only in southeastern Veracruz. Many of the shared species with the GCL contain wide-ranging generalist species that occur all along the Gulf lowlands from Tamaulipas and the adjacent USA into the Yucatan Peninsula of Mexico and northern Central America, and a few enter northern South America ( Wilson and Johnson 2010). In our opinion, those lowland generalist species are the main reason why the SLT clusters with the GLC instead of with the SMO, even though the GLC and SMO share borders along the northern half of Veracruz.
Distribution Status Categorizations
This analysis of the distributional status of the members of the Veracruz herpetofauna utilizes the same system employed by Alvarado-Díaz et al. (2013) and all the other entries in the MCS (see above). The four categories in this system are non-endemic, country endemic, state endemic, and non-native. The data are presented based on these categories in Table 8, and summarized in Table 9.
In descending order of size, the numbers of species in each of these categories are non-endemics: 169 (47.1%), country endemics: 138 (38.4%), state endemics: 44 (12.3%), and non-natives: 8 (2.2%). The herpetofauna of Veracruz, therefore, resembles several other state herpetofaunas covered previously in the MCS in that the largest number of species fall into the non-endemic category, i.e., Oaxaca ( Mata-Silva et al. 2015); Chiapas ( Johnson et al. 2015a); Tamaulipas ( Terán-Juárez et al. 2016); Nuevo León (Nevárez-de los Reyes et al. 2016); the Mexican Yucatan Peninsula ( González-Sánchez et al. 2017); and Coahuila ( Lazcano et al. 2019). In other No. 29. Spilotes pullatus Linnaeus, 1758 . The Tropical Tree Snake is a non-endemic species found from Tamaulipas southward through Central America and South America to Argentina on the Atlantic versant, and from the Isthmus of Tehuantepec to Ecuador on the Pacific versant ( Lemos-Espinal and Dixon 2013). This individual was found at Los Tuxtlas, in the municipality of San Andrés Tuxtla. Wilson et al. (2013b) determined its EVS as 6, placing it in the middle of the low vulnerability category. Its conservation status has not been determined by either the IUCN or SEMARNAT. Photo by Christian Berriozabal-Islas.
No. 31. Clelia scytalina (Cope, 1867) . The Mexican Snake Eater is a non-endemic species ranging from southern Mexico to Guatemala and Belize ( Köhler 2008). This individual was found at Los Tuxtlas, in the municipality of San Andrés Tuxtla. Wilson et al. (2013b) determined its EVS as 13, placing it at the upper limit of the medium vulnerability category. Its conservation status has not been determined by either the IUCN or SEMARNAT. Photo by Christian Berriozabal-Islas. No. 30. Adelphicos quadrivirgatum Jan, 1862 . The Mesoamerican Earth Snake is a non-endemic species distributed from Tamaulipas to Honduras on the Atlantic versant, and from Oaxaca to Guatemala on the Pacific versant ( Lemos-Espinal and Dixon 2013). This individual was located at Los Tuxtlas, in the municipality of San Andrés Tuxtla. Wilson et al. (2013b) ascertained its EVS as 10, placing it at the lower limit of the medium vulnerability category. Its conservation status has been calculated as Least Concern by the IUCN, and as Special Protection (Pr) by SEMARNAT. Photo by Christian Berriozabal-Islas.
No. 32. Imantodes cenchoa (Linnaeus, 1758) . The Bluntheaded Treesnake is a non-endemic species occurring at low and moderate elevations (up to 1,600 m) on the Atlantic versant from southern Tamaulipas, southward through Central and South America to Argentina. It also occurs along the Pacific lowlands and premontane slopes from Chiapas to Guatemala. In the Yucatán Peninsula, it is known from southern Campeche and Quintana Roo, but apparently is absent from the arid north-western region of the peninsula ( Heimes 2016). This individual was found at Los Tuxtlas, in the municipality of San Andrés Tuxtla. Wilson et al. (2013b) determined its EVS as 6, placing it in the low vulnerability category. Its conservation status has not been determined by either the IUCN or SEMARNAT. Photo by Elí García-Padilla. state herpetofaunas, the largest number of species lies in the country endemic category, including Michoacán ( Alvarado-Díaz et al. 2013), Nayarit ( Woolrich-Piña et al. 2016), Jalisco ( Cruz-Sáenz et al. 2017), Puebla ( Woolrich-Piña et al. 2017), and Hidalgo ( Ramírez-Bautista et al. 2020).
In the 11 previous MCS entries, the numbers of state endemic species vary significantly from one in Nayarit and Nuevo León ( Woolrich-Piña et al. 2016; Nevárez- de los Reyes et al. 2016, respectively) to 93 in Oaxaca ( Mata-Silva et al. 2015). The number of state endemics in Veracruz lies below the mid-point in this range at 44 ( Table 9). Half of these 44 species (22) are plethodontid salamanders of the genera Aquiloeurycea (one species), Chiropterotriton (six), Isthmura (two), Parvimolge (one), Pseudoeurycea (four), and Thorius (eight). The remaining species are anurans (seven species) and squamates (15).
As noted above, in some cases in the MCS the number of non-endemic species exceeds that of the country endemics, whereas in other cases the situation is reversed. So, the ratio of non-endemics to country endemics varies considerably. The ratios in which the number of non-endemic species exceeds that of the country endemic species range from 1.12 in the case of Oaxaca ( Mata-Silva et al. 2015) to 127.0 in the Yucatan Peninsula ( González-Sánchez et al. 2017). The ratios in which the number of country endemic species supersedes that of the non-endemics vary from 0.53 in the case of Jalisco to 0.88 in Hidalgo. As expected, the nature of this ratio depends on the proximity of the given Mexican state or region to either the USA or to Central America. In the case of the three MCS states that border the USA, the ratios are 3.22 (100/ 31 in Coahuila; Lazcano et al. 2019), 2.44 (95/ 39 in Nuevo León; Nevárez-de los Reyes et al. 2016), and 2.43 (119/ 49 in Tamaulipas; Terán- No. 33. Leptodeira maculata (Hallowell, 1861) . The Banded Cat-eyed Snake is a non-endemic species distributed from southern Tamaulipas along the Atlantic versant and from southern Sinaloa on the Pacific versant southward to Chiapas and western Guatemala ( Daza et al. 2009). This individual was located at Jonotal, in the municipality of Las Vigas de Ramírez. Wilson et al. (2013b) calculated its EVS as 7, placing it in the middle of the low vulnerability category. Its conservation status has been assessed as Least Concern by the IUCN, and as Special Protection (Pr) by SEMARNAT. Photo by Christian Berriozabal-Islas.
No. 35. Bothrops asper (Garman, 1883) . The Terciopelo is a non-endemic snake ranging from southwestern Tamaulipas to coastal Venezuela on the Atlantic versant, and from Costa Rica to southern Ecuador on the Pacific versant, with a disjunct population occurring in southern Chiapas and adjacent Guatemala ( Lemos-Espinal and Dixon 2013). This individual was found at Los Tuxtlas, in the municipality of San Andrés Tuxtla. Wilson et al. (2013b) determined its EVS as 12, placing it in the upper portion of the medium vulnerability category. Its conservation status has not been determined by either the IUCN or SEMARNAT. Photo by Christian Berriozabal-Islas. No. 34. Pliocercus elapoides (Cope, 1860) . The Variegated False Coralsnake is a non-endemic species occurring at low and moderate elevations (up to about 2,000 m) on the Atlantic slope from southern Tamaulipas, and on the Pacific from western Oaxaca, southward to western Honduras and El Salvador. In the Yucatán Peninsula, the species is known only from the base, in the south, and from the north ( Heimes 2016). This individual was encountered at Los Tuxtlas, in the municipality of San Andrés Tuxtla. Wilson et al. (2013b) determined its EVS as 10, placing it in the lower portion of the medium vulnerability category. Its conservation status has been assessed as Least Concern by the IUCN, and this species is not listed by SEMARNAT. Photo by Elí García-Padilla.
No. 36. Cerrophidion petlalcalensis Lopez-Luna, Vogt, and Torre-Loranca, 1999 . The Cerro Petlalcala Montane Pitviper is a country endemic species ranging at the type locality in the Cerro Petlalcala, in west-central Veracruz at elevations from 2,100 to 2,300 m ( López-Luna et al. 1999). This species also is known from the municipalities of Atlahuilco, Los Reyes, and Tequila in Veracruz, and in Oaxaca from the Sierra Mazateca ( De La Torre-Loranca et al. 2019) . This individual was found at San Andrés Tenejapan, in the municipality of the same name. Wilson et al. (2013b) calculated its EVS at 18, placing it in the upper portion of the high vulnerability category. Its conservation status has been assessed as Data Deficient by the IUCN, and this species is not listed by SEMARNAT. Photo by René Ávalos-Vela.
Juárez et al. 2016). In the case of the states or the region bordering Central America, the ratios are 8.38 (268/ 32 in Chiapas; Johnson et al. 2015a) and 127.00 (127/ 1 in the Yucatan Peninsula; González-Sánchez et al. 2017). The extreme figure for the Yucatan Peninsula is due, in part, to this region of Mexico being confluent with the portion of the peninsula that lies in Central America, principally northern Guatemala. This MCS area is the only one in which the number of country endemics is overwhelmed by the number of regional endemics, i.e., one compared to 11 ( González-Sánchez et al. 2017). As indicated above, Veracruz is a state in which the number of non-endemic species is greater than that of the country endemics, so the ratio of the former to the latter is 169/138, or 1.22.
Eight non-native species currently reside in Veracruz: Eleutherodactylus planirostris , Lithobates catesbeianus , Norops sagrei , Hemidactylus frenatus , H. mabouia , H. turcicus , Virgotyphlops braminus , and Trachemys scipta . Two of these species ( H. frenatus and I. braminus ) are the most widespread of the non-native species reported in the previous 11 MCS entries, as they have been reported in 11 and 12 states, respectively ( González-Sánchez et al. 2017, which covers the three Mexican states of the Yucatan Peninsula).
Wilson et al. (2017) designed a system for categorizing the distribution of the herpetofauna of Mesoamerica, and it was applied to the pertinent categories in this study, with the data summarized in Table 10. As mentioned above, of the 359 total species in Veracruz, 169 are non-endemic, and these 169 taxa were placed in six of the nine categories recognized by Wilson et al. (2017), including MXUS, MXCA, MXSA, USCA, USSA, and OCEA. Given the proximity of Veracruz to Guatemala, in Central America, and the location of the state south of the Tropic of Cancer, understandably the largest number of species is in the MXCA category (89; 52.7%). The next largest number is allocated to the MXSA category (30; 17.8%), followed closely by the number in the MXUS category (29; 17.2%). The remaining 20 species are placed in the USCA (11; 6.5%), USSA (four; 2.4%), and OCEA (five; 3.0%) categories.
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