The endemic herpetofauna of Central America: a casualty of anthropocentrism Author Mata-Silva, Vicente Department of Biological Sciences, The University of Texas at El Paso, El Paso, Texas 79968 - 0500, USA vmata@utep.edu Author DeSantis, Dominic L. Department of Biological Sciences, The University of Texas at El Paso, El Paso, Texas 79968 - 0500, USA dldesantis@miners.utep.edu Author García-Padilla, Elí Oaxaca de Juárez, Oaxaca 68023, MEXICO eligarcia_18@hotmail.com Author Johnson, Jerry D. Department of Biological Sciences, The University of Texas at El Paso, El Paso, Texas 79968 - 0500, USA jjohnson@utep.edu Author Wilson, Larry David Centro Zamorano de Biodiversidad, Escuela Agrícola Panamericana Zamorano, Departamento de Francisco Morazán, HONDURAS bufodoc@aol.com text Amphibian & Reptile Conservation 2019 e 168 2019-01-20 13 1 1 64 journal article 297965 10.5281/zenodo.11391571 799461e0-d0dd-4c4b-82c4-886726d6b079 1525-9153 11391571 The Distributional Status of Dipsas viguieri Peters (1960) reviewed the taxonomic status of members of the subfamily Dipsadinae , and placed nine species in the Dipsas articulata group, collectively distributed from western and southeastern Mexico to northwestern Ecuador . Peters’ view of the species-level relationships among the members of the articulata group was impacted by the paucity of specimens of each taxon known at that time, and he noted that only minimal scale and color differences separated certain species. Peters (1960) indicated the range of D. viguieri as the Pacific coast of Panama . Pérez Santos and Moreno (1988) reported on a specimen of D. gracilis from the Pacific coast of Colombia (see discussion below), and Pérez Santos (1999) noted the occurrence of D. viguieri from both versants of Panama , including a specimen from the province of Bocas del Toro in the western part of the country. Subsequently, Köhler (2001 ; 2003 ; 2008 ) noted the range of Dipsas viguieri as eastern Panama and western Colombia but did not provide additional information. Cadle (2005) , in a paper on the systematics of the Dipsas oreas complex, tentatively referred to a specimen ( FMNH 74376) from northwestern Colombia near the Panama border as D. viguieri , which previously had been identified as D. gracilis . Nonetheless, Cadle (2005) stated that these two taxa were not distinguishable by any reported characteristics, and on p. 128 noted that, “Without additional study, I am unable to adequately differentiate Dipsas viguieri (eastern Panama and northern Chocó , Colombia ) and D. gracilis (western Ecuador and extreme northern Peru ).” Further, based on an examination of morphological characters, Cadle indicated geography as the only currently reliable means of assigning names to these species. Similarly, in discussing a number of poorly-known Dipsas from South America, Harvey (2008) commented that he was unable to distinguish D. viguieri from D. gracilis , and thus did not include D. viguieri in his key but noted that the D. articulata complex requires further study. Plate 1. Atelopus varius (Lichtenstein and Martens, 1856) . The Harlequin Frog is a priority ten species with an EVS of 11, distributed on both versants of the cordilleras of Costa Rica and western Panama ( Frost 2018 ). This individual is from one of three known surviving populations of this species, and in Costa Rica it is being surveyed near Uvita, in the province of Puntarenas. Photo by César Barrio-Amorós. Plate 2. Incilius holdridgei (Taylor, 1952) . Holdridge’s Toad is a priority one species with an EVS of 14, which is restricted in distribution to Volcán Barva, Costa Rica ( Frost 2018 ). This individual was located in Alto del Roble, in the province of Heredia. Photo by Víctor Acosta-Chaves. Plate 3. Incilius melanochlorus (Cope, 1877) . The Wet Forest Toad is a priority eight species with an EVS of 12, with a distribution on the Atlantic versant of Costa Rica and adjacent Panama, and likely in adjacent Nicaragua ( Frost 2018 ). This individual was found in Centro Soltis, San Isidro de Peñas Blancas, in the province of Alajuela, Costa Rica. Photo by Víctor Acosta-Chaves. Jaramillo et al. (2010) and Johnson et al. (2015) regarded Dipsas viguieri as endemic to Panama . Wallach et al. (2014: 235) , however, considered D. viguieri as occupying “Eastern Panama (Canal Zone, Darién , Panamá ) and Colombia (? Chocó , Piura), NSL– 60 m . ” In their D. gracilis account, however, these authors noted, “Colombian record doubtful fide Cadle (2005: 123) : possibly D. viguieri . Dipsas gracilis and D. viguieri possibly conspecific fide Harvey (2008: 429) .” Wallach et al. (2014: 232) , however, apparently confused the information provided by Cadle (2005) , as the FMNH specimen tentatively was referred to D. viguieri and not D. gracilis . Finally, Ray (2017) indicated the range of D. viguieri as eastern Panama to northwestern Colombia . The historical timeline for information on the distribution and taxonomic status of Dipsas viguieri has been unclear, as different workers have maintained that this species is endemic to Panama or occurs in both Panama and Colombia . In the absence of a definitive analysis involving morphological and molecular approaches, for the purpose of this paper we are considering D. viguieri as not endemic to Panama . Global Status of the Central American Herpetofauna As with the Mexican herpetofauna ( Johnson et al. 2017 ), the Central American herpetofauna also is highly diverse, consisting of 60 families, 214 genera, and 1,095 species ( Table 1 ), organized into six orders ( Anura , Caudata , Gymnophiona , Crocodylia , Squamata , and Testudines ). The level of herpetofaunal diversity in Central America is intermediate between that found in Mexico and North America ( United States – Canada ). The number of species in the United States – Canada is the same as Johnson et al. (2017) reported, i.e., 650 (Center for North American Herpetology website; accessed 9 December 2017 ). Johnson et al. (2017) reported the number of species in Mexico as 1,292. Even though the number of herpetofaunal species occurring in Central America is intermediate between that found in the United States – Canada and Mexico , Central America contains about 8.5 times the number of taxa by area as found in Mexico , and 155.6 times the number found in the United States – Canada . Thus, the relative degree of biodiversity is significantly higher in Central America when compared to that in Mexico and the United States – Canada . If we consider Central America as a single region in our analysis (i.e., not one divided into seven countries), then its herpetofauna also is significant when compared to that of other areas in Latin America. With respect to amphibians, the 509 species occurring in Central America is the fifth largest in Latin America (amphibiaweb.com; 15 April 2018 ), and is closest to that for the country of Ecuador , at 562. The area /species ratio for Ecuador however, is 504.6, compared to 998.0 for Central America . Considering the numbers of crocodylian, squamate, and turtle species, the 586 species in Central America is comparable to that recorded for the neighboring country of Colombia , which is 611 (reptile-database.org; accessed 29 December 2017 ). Colombia , however, with an area of 1,141,748 km 2 , is 2.25 times the size of Central America , which contains an area of 507,966 km 2 (www. Oei.es/ historico/cultura2/ Colombia /03.htm; accessed 29 December 2017 ). Thus , the area/species ratio for Colombia is 1,868.7, compared to 868.0 for that of Central America. Only Brazil (799) and Colombia in South America contain more species than Central America (reptile-database.org; accessed 29 December 2017 ). Endemism within the Central American Herpetofauna The proportion of herpetofaunal endemism in Central America is slightly less than in Mexico , the other major segment of Mesoamerica. The percentage in Central America is 56.9 ( Table 2 ) compared to 61.1 in Mexico ( Johnson et al. 2017 ). This percentage in Central America is based on an endemic herpetofauna of 623 species and a total herpetofauna of 1,095 species ( Table 2 ). Both of the comparable figures for the Mexican herpetofauna are higher, i.e., 789 and 1,292 ( Johnson et al. 2017 ). As noted by Johnson et al. (2015: 26) , “Mesoamerica is one of the world’s most important biodiversity reservoirs, and Central America contains a substantial component of that region’s herpetofauna.” We illustrate the breakdown of the total and endemic components of the Central American herpetofauna in Fig. 1 . This graph shows the close correspondence between the endemic and total number of salamander species, the relatively distant correspondence between the endemic and total number of squamate species, and the intermediate correspondence between the two figures for anurans ( Fig. 1 ). Of the 60 families represented in Central America, 38 (63.3%) contain endemic species ( Table 2 ). This leaves 22 families with no endemic representation, including the anuran families Aromobatidae , Hemiphractidae , and Rhinophrynidae , the crocodylian families Alligatoridae and Crocodylidae , the squamate families Amphisbaenidae , Hoplocercidae , Polychrotidae , Xenosauridae , Boidae , Charinidae , Loxocemidae , Natricidae , Sibynophiidae , and Tropidophiidae , and the turtle families Cheloniidae , Chelydridae , Dermatemydidae , Dermochelyidae , Emydidae , Staurotypidae, and Testudinidae . In Central America these are small-content families, with species numbers ranging from one to five ( Table 2 ). The families with endemic representation have total numbers ranging from one to 166; the endemic numbers vary from one to 143 ( Table 2 ). Fig. 1. Graph comparing the endemic and total number of species for the Central American herpetofauna, arranged by order. Table 1. Diversity of the Central American herpetofauna at the familial, generic, and specific levels.

Orders	Families	Genera	Species
Anura	14	58	328
Caudata	1	7	166
Gymnophiona	2	4	15
Crocodylia	2	2	3
Squamata	32	130	560
Testudines	9	13	23
Totals	60	214	1,095

Table 2. Degree of endemism of the Central American herpetofauna at the species level, arranged by family.

Family	Total Number of Species	Number of Endemic Species	Percentage of Endemism
Aromobatidae	1	—	—
Bufonidae	40	24	60.0
Centrolenidae	15	4	26.7
Craugastoridae	102	77	75.5
Dendrobatidae	21	14	66.7
Eleutherodactylidae	16	10	62.5
Hemiphractidae	3	—	—
Hylidae	92	52	60.9
Leptodactylidae	9	1	11.1
Microhylidae	9	1	11.1
Phyllomedusidae	7	2	28.6
Pipidae	1	1	100

Table 2 (continued). Degree of endemism of the Central American herpetofauna at the species level, arranged by family.

Family	Total Number of Species	Number of Endemic Species	Percentage of Endemism
Ranidae	11	6	54.5
Rhinophrynidae	1	—	—
Subtotals	328	192	58.5
Plethodontidae	166	143	86.1
Subtotals	166	143	86.1
Caeciliidae	7	3	42.9
Dermophiidae	8	4	50.0
Subtotals	15	7	46.7
Totals	509	342	67.2
Alligatoridae	1	—	—
Crocodylidae	2	—	—
Subtotals	3	—	—
Amphisbaenidae	2	—	—
Anguidae	32	25	78.1
Corytophanidae	9	1	11.1
Dactyloidae	104	74	71.2
Eublepharidae	2	1	50.0
Gymnophthalmidae	14	3	21.4
Helodermatidae	2	1	50.0
Hoplocercidae	2	—	—
Iguanidae	9	7	77.8
Mabuyidae	5	4	80.0
Phrynosomatidae	17	2	11.8
Phyllodactylidae	5	3	60.0
Polychrotidae	1	—	—
Scincidae	3	1	33.3
Sphaerodactylidae	22	13	59.1
Sphenomorphidae	4	1	25.0
Teiidae	16	5	31.3
Xantusiidae	4	2	50.0
Xenosauridae	1	—	—
Anomalepididae	3	1	33.3
Boidae	5	—	—
Charinidae	2	—	—
Colubridae	79	30	38.0
Dipsadidae	145	77	53.1
Elapidae	19	6	31.6
Leptotyphlopidae	6	3	50.0
Loxocemidae	1	—	—
Natricidae	5	—	—
Sibynophiidae	2	—	—
Tropidophiidae	1	—	—
Typhlopidae	5	3	60.0
Viperidae	33	16	48.5
Subtotals	560	279	49.8
Cheloniidae	4	—	—

Table 2 (continued). Degree of endemism of the Central American herpetofauna at the species level, arranged by family.

Family	Total Number of Species	Number of Endemic Species	Percentage of Endemism
Chelydridae	2	—	—
Dermatemydidae	1	—	—
Dermochelyidae	1	—	—
Emydidae	2	—	—
Geoemydidae	5	1	20.0
Kinosternidae	4	1	25.0
Staurotypidae	3	—	—
Testudinidae	1	—	—
Subtotals	23	2	8.7
Totals	586	281	48.0
Sum Totals	1,095	623	56.9

Of the 14 anuran families with representatives in Central America, 11 contain endemic species, which include 192 (58.5%) of the total of 328 species ( Table 2 ). Of these 11 families, the largest numbers of endemics are 77 in the Craugastoridae and 52 in the Hylidae . Other than in the Pipidae , with one total and one endemic species (100%), as might be expected, the percentage of endemism is next highest in the Craugastoridae (75.5%), but the third highest is in the Dendrobatidae (66.7%), and not the Hylidae (60.9%). The value for the Eleutherodactylidae (62.5%) also is higher than that for the Hylidae . The remaining families contain from one to 24 endemic species ( Table 2 ). A single family of salamanders, the Plethodontidae , occurs in Central America. The percentage of endemism (86.1%) is amazingly high and is the highest in all the 38 families represented ( Table 2 ). The endemic species of caecilians (seven) make up less than one-half (46.7%) of the total number of 15 in Central America. Three of the endemics are caeciliids and four are dermophiids. None of the three species of crocodylians in Central America is endemic. Crocodylus acutus and Caiman crocodilus rather are among the naturally most broadly distributed herpetofaunal species in the Western Hemisphere. The squamates are the most speciose group of herpetofaunal organisms in Central America, with 560 species distributed among 32 families ( Table 2 ). Only the endemic proportions of turtles (8.7%) and crocodylians (0.0%) are lower than those of squamates (49.8%). The endemic squamates are more or less evenly divided between the lizards (143) and snakes (136). Of the 19 families of lizards with representatives in Central America, 15 contain endemic species (78.9%). The largest numbers of endemic squamate species are found within the families Dactyloidae (74) and Dipsadidae (77). The next largest number of endemic lizards (25) is allocated to the family Anguidae . The remaining 12 lizard families contain only one to 13 endemic species. The percentage of endemism among the lizard families ranges from 11.1% in the Corytophanidae to 80.0% in the Mabuyidae ( Table 2 ). Thirteen families of snakes are represented in Central America, of which seven contain endemic species (53.8%). The greatest numbers of endemic species are in the families Dipsadidae (77) and Colubridae (30). The next largest number (16) lies within the family Viperidae ; the remaining four snake families contain from one to six endemic species ( Table 2 ). The percentage of endemism among the snake families ranges from 31.6% in the Elapidae to 60.0% in the Typhlopidae ( Table 2 ). The percentage of endemism among turtles is very low, with only two such species among a total of 23 (8.7%), one each in the families Geoemydidae and Kinosternidae ( Table 2 ). At the ordinal level, the percentage of endemism is highest among the salamanders at 86.1%, the same figure as for the family Plethodontidae , since it is the only family of salamanders found in Central America ( Table 2 ). The percentage is lowest among the crocodylians at 0.0%. Intermediate values are evident for the anurans (58.5%), squamates (49.8%), and caecilians (46.7%). The level of herpetofaunal endemism in Central America is comparable to that found in the other portions of the North American continent, i.e., Mexico , as well as the United States – Canada ( Table 3 ). The overall level for Central America, however, is slightly lower (56.9%) than for either Mexico (61.5%) or the United States – Canada (61.2%). Although, as expected, the total number of herpetofaunal species is lower in the United States – Canada than to the south in Mesoamerica ( Table 3 ); even with the greater area of the two northern nations, the level of endemism still is impressive. Of the total of 650 species in the United States – Canada , 398 are endemic, for a percentage of endemism of 61.2%. Notably, this level of endemism is based heavily on the amphibians, especially the salamanders. The proportion of amphibian endemism is more than 10 points higher in the United States – Canada (78.6%) than in Mexico (68.3%) or Central America (67.2%). The percentage of endemism is higher for both anurans and salamanders in the United States – Canada (64.4% and 86.4%, respectively) than for these two groups in either Mexico (60.0% and 82.8%, respectively) or Central America (58.5% and 86.1%, respectively). Significantly, the level of amphibian endemism in all three regions heavily depends on the salamanders, i.e., over 80%. Curiously enough, such high incidences of salamander species-level endemism are not correlated with the incidence of family-level endemism, which decreases markedly from the United States – Canada through Mexico to Central America. The 191 salamander species in the United States – Canada are organized within eight families, but with the majority allocated to the family Plethodontidae (Center for North American Herpetology website; accessed 2 January 2018 ). Four of these eight families, the Amphiumidae , Cryptobranchidae , Proteidae , and Rhyacotritonidae occur no farther south than the United States . Four families, the Ambystomatidae , Plethodontidae , Salamandridae , and Sirenidae are represented in Mexico , but with the greatest number of species in the Plethodontidae , as in the United States – Canada . Only a single family, the Plethodontidae , is found in Central America. In the United States – Canada , 145 of the 191 species of salamanders are in the family Plethodontidae (75.9%). Of the 151 species found in Mexico 130 (86.1%) are in the Plethodontidae ( Wilson et al., 2017 ) ; the remaining 21 species are allocated to three families (see above). Finally, of the 166 species of salamanders found in Central America, all are in the Plethodontidae ; obviously the percentage of occupancy is 100%. Plate 4. Craugastor sandersoni (Schmidt, 1941) . Sanderson’s Rainfrog is a priority two species with an EVS of 19, which is distributed on the “Caribbean slopes of the Maya Mountains in east-central Belize southward to the Montañas del Mico in eastern Guatemala and westward into the Sierra de Santa Cruz, the eastern portion of the Sierra de las Minas, and the foothills of the northern Alta Verapaz” ( Frost 2018 ).This individual was found in Montañas del Mico, Guatemala. Photo by Sean Michael Rovito. Plate 5. Oophaga granulifera (Taylor, 1958) .The Granular Poison Frog is a priority two species with an EVS of 17, distributed in the Golfo Dulce region of Pacific coastal Costa Rica and presumably in adjacent Panama ( Frost 2018 ). This individual was encountered in Ciudad Cortéz de Osa, in the province of Puntarenas, Costa Rica. Photo by Víctor Acosta-Chaves. Plate 6. Oophaga pumilio (Schmidt, 1857) . The Strawberry Poison Frog is a priority one species with an EVS of 16, and its distribution extends along the Atlantic versant from “eastern Nicaragua…south through the lowlands of Costa Rica and northwestern Panama (Savage 2002: 388). This individual came from the mainland in the province of Bocas del Toro, Panama. Photo by Abel Batista. Plate 7. Oophaga vicentei (Jungfer, Weyoldt, and Juraska, 1996) . This dendrobatid frog is a priority two species with an EVS of 16, distributed on the Atlantic versant “of the provinces of Veraguas and Coclé and the upper reaches of Pacific versant in Coclé, central Panama” ( Frost 2018 ). This individual is from Santa Fé National Park, in the province of Veraguas, Panama. Photo by Abel Batista. Plate 8. Phyllobates lugubris (Schmidt, 1857) . The Lovely Poison Frog is a priority one species with an EVS of 17, which ranges along the Atlantic versant from “extreme southeastern Nicaragua to northwestern Panama; a single record from just west of the Panama Canal” (Savage 2002:390).This individual was found in the Donoso region, in the province of Colón, Panama. Photo by Abel Batista. Plate 9. Diasporus hylaeformis (Cope, 1876) . The Pico Blanco Robber Frog is a priority one species with an EVS of 17, with a distribution in the highlands of the cordilleras of Costa Rica and Panama ( Frost 2018 ). This individual came from Alto del Roble, in the province of Heredia, Costa Rica. Photo by Víctor Acosta-Chaves. Table 3. Total number of species, endemic species, and relative endemism within the herpetofaunal groups in Central America, North America (United States–Canada), and Mexico. Data for Central America from this paper, for North America from CNAH (www.cnah.org; accessed 9 January 2018), and for Mexico from updated figures in Johnson et al. (2017) .

Herpetofaunal Groups	Total Species in Central America	Endemic Species in Central America	Relative Endemism in Central America (%)	Total Species in North America	Endemic Species in North America	Relative Endemism in North America (%)	Total Species in Mexico	Endemic Species in Mexico	Relative Endemism in Mexico (%)
Anurans	328	192	58.5	104	67	64.4	247	148	60.0
Salamanders	166	143	86.1	191	165	86.4	151	125	82.8
Caecilians	15	7	46.7	—	—	—	3	1	33.3
Subtotals	509	342	67.2	295	232	78.6	401	274	68.3
Crocodylians	3	—	—	2	1	50.0	3	—	—
Squamates	560	279	49.8	287	117	40.8	863	517	60.0
Turtles	23	2	8.3	66	48	72.7	51	20	39.2
Subtotals	586	281	48.0	355	166	46.8	917	537	58.6
Totals	1,095	623	56.9	650	398	61.2	1,318	811	61.5

Unlike the situation among the amphibians, the level of endemism among the crocodylians, squamates, and turtles is significantly lower in all three regions dealt with in Table 3 . The level of endemism among these taxa is lowest in the United States – Canada (46.8%), next lowest in Central America (48.0%), and highest in Mexico (58.6%). Given that squamates constitute the largest group, compared to the other two, the same pattern would be expected for them as for the entire group. Thus, the level of squamate endemism is lowest in the United States – Canada (40.8%), intermediate in Central America (49.8%), and highest in Mexico (60.0%). The differential between the percentages of endemism for amphibians versus the remainder of the herpetofauna ( Table 3 ) increases from that seen in Mexico (9.7%), through Central America (19.2%), to the United States – Canada (31.8%). Thus, in all three regions amphibians contribute more to the degree of endemicity than the remainder of the herpetofauna ( Table 3 ). Physiographic Distribution of the Endemic Central American Herpetofauna Given the considerable global significance of the diversity and endemicity of the Central American herpetofauna, as documented above, it is of paramount importance to protect its elements.As an initial step to determine the distributional patterns of these organisms in Central America, we collated the available information on the occurrence of the members of the herpetofauna among the 10 physiographic regions traditionally recognized in this portion of Mesoamerica ( Campbell, 1999 ; Wilson and Johnson, 2010 ; Fig. 2 ). Six of these regions occupy the lowlands of Central America, including the Yucatan Platform, the Caribbean lowlands of eastern Guatemala and northern Honduras , the Caribbean lowlands from Nicaragua to Panama , the Pacific lowlands from eastern Chiapas to south-central Guatemala , the Pacific lowlands from southeastern Guatemala to northwestern Costa Rica , and the Pacific lowlands from central Costa Rica through Panama ( Table 4 ). Four regions are found in the highlands of Central America, including the western nuclear Central American highlands, the eastern nuclear Central American highlands, the Isthmian Central American highlands, and the highlands of eastern Panama ( Table 4 ). We document the distribution of the 623 endemic members of the Central American herpetofauna among the 10 physiographic regions in Table 4 , and summarize these data in Table 5 and Fig. 3 . Thetotalnumberoftheendemicspeciesdistributedwithin the 10 physiographic regions ranges from a low of six in the Yucatan Platform to 254 in the Isthmian Central American highlands ( Table 5 ). The mean regional occupancy figure is 94.3. Four of the regional values lie above or close to the mean figure, as follows: western nuclear Central American highlands ( CG ; 117), eastern nuclear Central American highlands (HN; 178), Isthmian Central American highlands ( CRP ; 254), Caribbean lowlands of eastern Guatemala and northern Honduras (GH; 94), and Caribbean lowlands from Nicaragua to Panama ( NP ; 123). Given these species numbers, the five regions with values above or close to the mean are the most significant for conservation remediation. Fig. 2. Physiographic regions of Central America, after Campbell (1999) . Abbreviations are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform. Table 4. Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions.Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform.? = species known from indeterminate locality (see text).

Physiographic Regions of Central America										Totals
CG	HN	CRP	EP	YP	GH	NP	CGU	GCR	CP
Anura (192 species)
Bufonidae (24 species)
Atelopus certus	+	+	2
Atelopus chiriquiensis	+	1
Atelopus chirripoensis	+	1
Atelopus limosus	+	1
Atelopus senex	+	1
Atelopus varius	+	+	+	3
Atelopus zeteki	+	+	2

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform.? = species known from indeterminate locality (see text).

Physiographic Regions of Central America										Totals
CG	HN	CRP	EP	YP	GH	NP	CGU	GCR	CP
Incilius aucoinae	+	1
Incilius chompipe	+	1
Incilius epioticus	+	1
Incilius fastidiosus	+	1
Incilius guanacaste	+	1
Incilius holdridgei	+	1
Incilius ibarrai	+	+	2
Incilius karenlipsae	+	1
Incilius leucomyos	+	+	2
Incilius majordomus	+	1
Incilius melanochlorus	+	1
Incilius periglenes	+	1
Incilius peripatetes	+	1
Incilius porteri	+	1
Incilius signifer	+	+	2
Rhinella centralis	+	+	2
Rhinella chrysophora	+	1
Centrolenidae (4 species)
Cochranella granulosa	+	+	+	+	+	5
Hyalinobatrachium dianae	+	+	2
Hyalinobatrachium talamancae	+	1
Hyalinobatrachium vireovittatum	+	1
Craugastoridae (77 species)
Craugastor adamastus	+	1
Craugastor anciano	+	1
Craugastor andi	+	1
Craugastor angelicus	+	1
Craugastor aphanus	+	1
Craugastor aurilegulus	+	+	2
Craugastor azueroensis	+	1
Craugastor bocourti	+	1
Craugastor bransfordii	+	+	2
Craugastor campbelli	+	+	2
Craugastor catalinae	+	1
Craugastor chac	+	+	+	3
Craugastor charadra	+	+	2
Craugastor chingopetaca	+	1
Craugastor chrysozetetes	+	1
Craugastor coffeus	+	1
Craugastor cruzi	+	1
Craugastor cuaquero	+	1

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform.? = species known from indeterminate locality (see text).

Physiographic Regions of Central America										Totals
CG	HN	CRP	EP	YP	GH	NP	CGU	GCR	CP
Craugastor cyanochthebius	+	1
Craugastor daryi	+	1
Craugastor emcelae	+	1
Craugastor emleni	+	1
Craugastor epochthidius	+	+	2
Craugastor escoces	+	1
Craugastor evanesco	+	+	2
Craugastor fecundus	+	+	2
Craugastor fleischmanni	+	1
Craugastor gabbi	+	1
Craugastor gollmeri	+	+	+	3
Craugastor gulosus	+	1
Craugastor inachus	+	1
Craugastor jota	+	1
Craugastor laevissimus	+	+	+	3
Craugastor lauraster	+	+	2
Craugastor megacephalus	+	+	+	+	4
Craugastor melanostictus	+	1
Craugastor merendonensis	+	1
Craugastor milesi	+	1
Craugastor mimus	+	+	+	+	4
Craugastor monnichorum	+	1
Craugastor myllomyllon	+	1
Craugastor nefrens	+	1
Craugastor noblei	+	+	+	+	+	5
Craugastor obesus	+	+	2
Craugastor olanchano	+	1
Craugastor omoaensis	+	1
Craugastor pechorum	+	+	2
Craugastor persimilis	+	+	2
Craugastor phasma	+	1
Craugastor podiciferus	+	1
Craugastor polyptychus	+	1
Craugastor punctariolus	+	1
Craugastor ranoides	+	+	+	+	4
Craugastor rayo	+	1
Craugastor rhyacobatrachus	+	1
Craugastor rivulus	+	1
Craugastor rostralis	+	+	2
Craugastor rugosus	+	+	2
Craugastor sabrinus	+	+	2

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform.? = species known from indeterminate locality (see text).

Physiographic Regions of Central America										Totals
CG	HN	CRP	EP	YP	GH	NP	CGU	GCR	CP
Craugastor saltuarius	+	1
Craugastor sandersoni	+	+	2
Craugastor stadelmani	+	1
Craugastor stejnegerianus	+	+	+	3
Craugastor tabasarae	+	1
Craugastor talamancae	+	1
Craugastor taurus	+	1
Craugastor trachydermus	+	1
Craugastor underwoodi	+	1
Craugastor xucanebi	+	1
Pristimantis adnus	+	1
Pristimantis altae	+	+	2
Pristimantis caryophyllaceus	+	+	2
Pristimantis cerasinus	+	+	+	3
Pristimantis museosus	+	1
Pristimantis pardalis	+	+	+	3
Pristimantis pirrensis	+	1
Strabomantis laticorpus	+	1
Dendrobatidae (14 species)
Ameerega maculata ?	?
Andinobates claudiae	+	1
Andinobates geminisae	+	1
Colostethus latinasus	+	1
Ectopoglossus astralogaster	+	1
Ectopoglossus isthminus	+	1
Oophaga arborea	+	1
Oophaga granulifera	+	+	2
Oophaga pumilio	+	1
Oophaga speciosa	+	1
Oophaga vicentei	+	+	2
Phyllobates lugubris	+	1
Phyllobates vittatus	+	1
Silverstoneia flotator	+	+	+	3
Eleutherodactylidae (10 species)
Diasporus citrinobapheus	+	1
Diasporus darienensis	+	1
Diasporus diastema	+	+	+	+	4
Diasporus hylaeformis	+	1
Diasporus igneus	+	1
Diasporus majeensis	+	1
Diasporus pequeno	+	1
Diasporus sapo	+	1

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform.? = species known from indeterminate locality (see text).

Physiographic Regions of Central America										Totals
CG	HN	CRP	EP	YP	GH	NP	CGU	GCR	CP
Diasporus tigrillo	+	1
Diasporus ventrimaculatus	+	1
Hylidae (52 species)
Atlantihyla panchoi	+	+	2
Atlantihyla spinipollex	+	+	2
Bromeliohyla melacaena	+	1
Dryophytes bocourti	+	1
Duellmanohyla legleri	+	1
Duellmanohyla lythrodes	+	1
Duellmanohyla rufioculis	+	1
Duellmanohyla salvadorensis	+	+	2
Duellmanohyla salvavida	+	+	2
Duellmanohyla soralia	+	+	2
Duellmanohyla uranochroa	+	+	2
Ecnomiohyla bailarina	+	+	2
Ecnomiohyla fimbrimembra	+	+	2
Ecnomiohyla minera	+	1
Ecnomiohyla rabborum	+	1
Ecnomiohyla salvaje	+	1
Ecnomiohyla sukia	+	+	2
Ecnomiohyla thysanota	+	1
Ecnomiohyla veraguensis	+	1
Exerodonta catracha	+	1
Exerodonta perkinsi	+	1
Hyloscirtus colymba	+	+	2
Isthmohyla angustilineata	+	1
Isthmohyla calypso	+	1
Isthmohyla debilis	+	1
Isthmohyla graceae	+	1
Isthmohyla infucata	+	1
Isthmohyla insolita	+	1
Isthmohyla lancasteri	+	+	2
Isthmohyla picadoi	+	1
Isthmohyla pictipes	+	1
Isthmohyla pseudopuma	+	1
Isthmohyla rivularis	+	1
Isthmohyla tica	+	1
Isthmohyla xanthosticta	+	1
Isthmohyla zeteki	+	1
Plectrohyla calvata	+	1
Plectrohyla chrysopleura	+	1

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform.? = species known from indeterminate locality (see text).

Physiographic Regions of Central America										Totals
CG	HN	CRP	EP	YP	GH	NP	CGU	GCR	CP
Plectrohyla dasypus	+	1
Plectrohyla exquisite	+	1
Plectrohyla glandulosa	+	1
Plectrohyla pokomchi	+	1
Plectrohyla psiloderma	+	1
Plectrohyla quecchi	+	1
Plectrohyla tecunumani	+	1
Plectrohyla teuchestes	+	1
Ptychohyla dendrophasma	+	1
Ptychohyla hypomykter	+	+	+	3
Quilticohyla sanctaecrucis	+	1
Scinax altae	+	1
Smilisca manisorum	+	1
Smilisca puma	+	1
Leptodactylidae (1 species)
Leptodactylus silvanimbus	+	1
Microhylidae (1 species)
Hypopachus pictiventris	+	1
Phyllomedusidae (2 species)
Agalychnis annae	+	+	2
Agalychnis saltator	+	+	+	3
Pipidae (1 species)
Pipa myersi	+	1
Ranidae (6 species)
Lithobates juliani	+	1
Lithobates lenca	+	1
Lithobates miadis	+	1
Lithobates taylori	+	1
Lithobates vibicarius	+	1
Lithobates warszewitschii	+	+	+	+	4
Caudata (143 species)
Plethodontidae (143 species)
Bolitoglossa alvaradoi	+	+	2
Bolitoglossa anthracina	+	1
Bolitoglossa aurae	+	1
Bolitoglossa aureogularis	+	1
Bolitoglossa bramei	+	1
Bolitoglossa carri	+	1
Bolitoglossa cataguana	+	1
Bolitoglossa celaque	+	1
Bolitoglossa centenorum	+	1

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform.? = species known from indeterminate locality (see text).

Physiographic Regions of Central America										Totals
CG	HN	CRP	EP	YP	GH	NP	CGU	GCR	CP
Bolitoglossa cerroensis	+	1
Bolitoglossa chucantiensis	+	1
Bolitoglossa colonnea	+	+	+	+	4
Bolitoglossa compacta	+	1
Bolitoglossa conanti	+	1
Bolitoglossa copia	+	1
Bolitoglossa cuchumatana	+	1
Bolitoglossa cuna	+	1
Bolitoglossa daryorum	+	1
Bolitoglossa decora	+	1
Bolitoglossa diaphora	+	1
Bolitoglossa diminuta	+	1
Bolitoglossa dofleini	+	+	+	+	4
Bolitoglossa dunni	+	1
Bolitoglossa epimela	+	1
Bolitoglossa eremia	+	1
Bolitoglossa gomezi	+	1
Bolitoglossa gracilis	+	1
Bolitoglossa heiroreias	+	1
Bolitoglossa helmrichi	+	1
Bolitoglossa huehuetenanguensis	+	1
Bolitoglossa indio	+	1
Bolitoglossa insularis	+	1
Bolitoglossa jacksoni	+	1
Bolitoglossa jugivagans	+	1
Bolitoglossa kamuk	+	1
Bolitoglossa kaqchikelorum	+	1
Bolitoglossa la	+	1
Bolitoglossa lignicolor	+	+	2
Bolitoglossa longissima	+	1
Bolitoglossa magnifica	+	1
Bolitoglossa marmorea	+	1
Bolitoglossa meliana	+	1
Bolitoglossa minutula	+	1
Bolitoglossa mombachoensis	+	1
Bolitoglossa morio	+	1
Bolitoglossa nigrescens	+	1
Bolitoglossa ninadormida	+	1
Bolitoglossa nussbaumi	+	1
Bolitoglossa nympha	+	+	+	3
Bolitoglossa obscura	+	1

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform.? = species known from indeterminate locality (see text).

Physiographic Regions of Central America										Totals
CG	HN	CRP	EP	YP	GH	NP	CGU	GCR	CP
Bolitoglossa odonnelli	+	+	2
Bolitoglossa omniumsanctorum	+	1
Bolitoglossa oresbia	+	1
Bolitoglossa pacaya	+	1
Bolitoglossa pesrubra	+	1
Bolitoglossa porrasorum	+	1
Bolitoglossa psephena	+	1
Bolitoglossa pygmaea	+	1
Bolitoglossa robinsoni	+	1
Bolitoglossa robusta	+	1
Bolitoglossa salvinii	+	+	2
Bolitoglossa schizodactyla	+	+	+	3
Bolitoglossa sombra	+	1
Bolitoglossa sooyorum	+	1
Bolitoglossa splendida	+	1
Bolitoglossa striatula	+	+	+	+	+	5
Bolitoglossa subpalmata	+	1
Bolitoglossa suchitanensis	+	1
Bolitoglossa synoria	+	1
Bolitoglossa taylori	+	1
Bolitoglossa tenebrosa	+	1
Bolitoglossa tica	+	1
Bolitoglossa tzultacaj	+	1
Bolitoglossa xibalba	+	1
Bolitoglossa zacapensis	+	1
Cryptotriton monzoni	+	1
Cryptotriton nasalis	+	+	2
Cryptotriton necopinus	+	1
Cryptotriton sierraminensis	+	1
Cryptotriton veraepacis	+	1
Cryptotriton xucaneborum	+	1
Dendrotriton bromeliacius	+	1
Dendrotriton chujorum	+	1
Dendrotriton cuchumatanus	+	1
Dendrotriton kekchiorum	+	1
Dendrotriton rabbi	+	1
Dendrotriton sanctibarbarus	+	1
Nototriton abscondens	+	1
Nototriton barbouri	+	1
Nototriton brodiei	+	1
Nototriton costaricense	+	1

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform.? = species known from indeterminate locality (see text).

Physiographic Regions of Central America										Totals
CG	HN	CRP	EP	YP	GH	NP	CGU	GCR	CP
Nototriton gamezi	+	1
Nototriton guanacaste	+	1
Nototriton lignicola	+	1
Nototriton limnospectator	+	1
Nototriton major	+	1
Nototriton matama	+	1
Nototriton mime	+	1
Nototriton nelsoni	+	1
Nototriton oreadorum	+	1
Nototriton picadoi	+	1
Nototriton picucha	+	1
Nototriton richardi	+	1
Nototriton saslaya	+	1
Nototriton stuarti	+	1
Nototriton tapanti	+	1
Nototriton tomamorum	+	1
Oedipina alfaroi	+	+	2
Oedipina alleni	+	+	2
Oedipina altura	+	1
Oedipina berlini	+	1
Oedipina capitalina	+	1
Oedipina carablanca	+	1
Oedipina chortiorum	+	1
Oedipina collaris	+	1
Oedipina cyclocauda	+	1
Oedipina fortunensis	+	1
Oedipina gephyra	+	1
Oedipina gracilis	+	1
Oedipina grandis	+	1
Oedipina ignea	+	+	2
Oedipina kasios	+	1
Oedipina koehleri	+	1
Oedipina leptopoda	+	1
Oedipina maritima	+	1
Oedipina motaguae	+	1
Oedipina nica	+	1
Oedipina nimaso	+	1
Oedipina pacificensis	+	1
Oedipina paucidentata	+	1
Oedipina petiola	+	1
Oedipina poelzi	+	1

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform.? = species known from indeterminate locality (see text).

Physiographic Regions of Central America										Totals
CG	HN	CRP	EP	YP	GH	NP	CGU	GCR	CP
Oedipina pseudouniformis	+	+	+	3
Oedipina quadra	+	1
Oedipina salvadorensis	+	1
Oedipina savagei	+	1
Oedipina stenopodia	+	1
Oedipina stuarti	+	+	2
Oedipina taylori	+	1
Oedipina tomasi	+	1
Oedipina tzutujilorum	+	1
Oedipina uniformis	+	1
Pseudoeurycea exspectata	+	1
Gymnophiona (7 species)
Caecilidae (3 species)
Caecilia volcani	+	1
Oscaecilia elongata	+	1
Oscaecilia osae	+	1
Dermophiidae (4 species)
Dermophis costaricensis	+	1
Dermophis gracilior	+	1
Dermophis occidentalis	+	+	2
Gymnopis multiplicata	+	+	+	+	+	5
Squamata (279 Species)
Anguidae (25 species)
Abronia anzuetoi	+	1
Abronia aurita	+	1
Abronia campbelli	+	1
Abronia fimbriata	+	1
Abronia frosti	+	1
Abronia gaiophantasma	+	1
Abronia meledona	+	1
Abronia montecristoi	+	1
Abronia salvadorensis	+	1
Abronia vasconcelosii	+	1
Celestus adercus	+	1
Celestus atitlanensis	+	+	2
Celestus bivittatus	+	1
Celestus cyanochloris	+	1
Celestus hylaius	+	1
Celestus laf	+	1
Celestus montanus	+	1
Celestus orobius	+	1

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform.? = species known from indeterminate locality (see text).

Physiographic Regions of Central America										Totals
CG	HN	CRP	EP	YP	GH	NP	CGU	GCR	CP
Celestus scansorius	+	1
Coloptychon rhombifer	+	1
Diploglossus bilobatus	+	+	+	3
Diploglossus montisilvestris	+	1
Mesaspis cuchumatanus	+	1
Mesaspis monticola	+	1
Mesaspis salvadorensis	+	1
Corytophanidae (1 species)
Basiliscus plumifrons	+	+	+	+	+	5
Dactyloidae (74 species)
Dactyloa brooksi	+	+	+	+	4
Dactyloa casildae	+	1
Dactyloa ginaelisae	+	1
Dactyloa ibanezi	+	+	2
Dactyloa insignis	+	+	+	3
Dactyloa kathydayae	+	1
Dactyloa kunayalae	+	+	2
Dactyloa maia	+	1
Dactyloa microtus	+	1
Dactyloa savagei	+	+	2
Norops alocomyos	+	1
Norops altae	+	1
Norops amplisquamosus	+	1
Norops apletophallus	+	+	+	+	4
Norops aquaticus	+	+	2
Norops benedikti	+	1
Norops bicaorum	+	1
Norops campbelli	+	1
Norops carpenteri	+	+	2
Norops charlesmyersi	+	+	2
Norops cobanensis	+	1
Norops cryptolimifrons	+	+	2
Norops cupreus	+	+	+	+	+	+	6
Norops cusuco	+	1
Norops datzorum	+	1
Norops elcopeensis	+	+	2
Norops fortunensis	+	1
Norops fungosus	+	1
Norops gruuo	+	1
Norops haguei	+	1
Norops heteropholidotus	+	1

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform.? = species known from indeterminate locality (see text).

Physiographic Regions of Central America										Totals
CG	HN	CRP	EP	YP	GH	NP	CGU	GCR	CP
Norops humilis	+	+	+	+	4
Norops intermedius	+	1
Norops johnmeyeri	+	1
Norops kemptoni	+	1
Norops kreutzi	+	1
Norops leditzigorum	+	1
Norops limifrons	+	+	+	+	+	+	6
Norops lionotus	+	+	+	3
Norops loveridgei	+	+	2
Norops macrophallus	+	+	+	3
Norops magnaphallus	+	1
Norops marsupialis	+	+	2
Norops mccraniei	+	+	2
Norops monteverde	+	1
Norops morazani	+	1
Norops muralla	+	1
Norops ocelloscapularis	+	1
Norops osa	+	1
Norops oxylophus	+	+	+	+	4
Norops pachypus	+	1
Norops pijolensis	+	1
Norops polylepis	+	+	2
Norops pseudokemptoni	+	1
Norops pseudopachypus	+	1
Norops purpurgularis	+	1
Norops quaggulus	+	+	+	3
Norops roatanensis	+	1
Norops rubribarbaris	+	1
Norops salvini	+	1
Norops sminthus	+	1
Norops tenorioensis	+	1
Norops townsendi	+	1
Norops triumphalis	+	1
Norops tropidolepis	+	1
Norops utilensis	+	1
Norops villai	+	1
Norops wampuensis	+	1
Norops wellbornae	+	+	+	+	4
Norops wermuthi	+	1
Norops wilsoni	+	+	2
Norops woodi	+	1

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform.? = species known from indeterminate locality (see text).

Physiographic Regions of Central America										Totals
CG	HN	CRP	EP	YP	GH	NP	CGU	GCR	CP
Norops yoroensis	+	1
Norops zeus	+	+	2
Eublepharidae (1 species)
Coleonyx mitratus	+	+	+	+	+	5
Gymnophthalmidae (3 species)
Bachia blairi	+	1
Echinosaura panamensis	+	+	+	3
Echinosaura apodema	+	+	2
Helodermatidae (1 species)
Heloderma charlesbogerti	+	+	2
Iguanidae (7 species)
Ctenosaura bakeri	+	1
Ctenosaura flavidorsalis	+	+	+	3
Ctenosaura melanosterna	+	+	2
Ctenosaura oedirhina	+	1
Ctenosaura palearis	+	1
Ctenosura praeocularis	+	+	2
Ctenosaura quinquecarinata	+	+	2
Mabuyidae (4 species)
Marisora alliacea	+	1
Marisora magnacornae	+	1
Marisora roatanae	+	1
Marisora unimarginata	+	+	+	+	+	5
Phrynosomatidae (2 species)
Sceloporus lunaei	+	+	2
Sceloporus malachiticus	+	+	+	3
Phyllodactylidae (3 species)
Phyllodactylus insularis	+	1
Phyllodactylus palmeus	+	1
Phyllodactylus paralepis	+	1
Scincidae (1 species)
Mesoscincus managuae	+	+	2
Sphaerodactylidae (13 species)
Lepidoblepharis emberawoundule	+	1
Lepidoblepharis rufigularis	+	1
Lepidoblepharis victormartinezi	+	+	2
Sphaerodactylus alphus	+	1
Sphaerodactylus dunni	+	1
Sphaerodactylus graptolaemus	+	1
Sphaerodactylus guanaje	+	1
Sphaerodactylus homolepis	+	1

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform.? = species known from indeterminate locality (see text).

Physiographic Regions of Central America										Totals
CG	HN	CRP	EP	YP	GH	NP	CGU	GCR	CP
Sphaerodactylus leonardovaldesi	+	1
Sphaerodactylus millepunctatus	+	+	+	+	+	+	6
Sphaerodactylus pacificus	+	1
Sphaerodactylus poindexteri	+	1
Sphaerodactylus rosaurae	+	1
Sphenomorphidae (1 species)
Scincella rara	+	1
Teiidae (5 species)
Cnemidophorus duellmani	+	1
Cnemidophorus ruatanus	+	+	2
Holcosus leptophrys	+	+	+	+	4
Holcosus miadis	+	1
Holcosus quadrilineatus	+	+	+	+	4
Xantusiidae (2 species)
Lepidophyma mayae	+	+	2
Lepidophyma reticulatum	+	+	2
Anomalepididae (1 species)
Helminthophis frontalis	+	+	2
Colubridae (30 species)
Dendrophidion apharocybe	+	+	+	+	4
Dendrophidion crybelum	+	1
Dendrophidion paucicarinatum	+	1
Dendrophidion rufiterminorum	+	+	+	+	+	5
Drymobius melanotropis	+	+	+	3
Leptodrymus pulcherrimus	+	+	+	+	4
Leptophis nebulosus	+	+	+	+	4
Mastigodryas alternatus	+	+	+	+	+	+	6
Mastigodryas dorsalis	+	+	2
Oxybelis wilsoni	+	1
Scolecophis atrocinctus	+	+	+	3
Tantilla albiceps	+	1
Tantilla armillata	+	+	+	+	4
Tantilla bairdi	+	1
Tantilla berguidoi	+	1
Tantilla brevicauda	+	+	2
Tantilla excelsa	+	+	2
Tantilla gottei	+	1
Tantilla hendersoni	+	1
Tantilla jani	+	+	2
Tantilla lempira	+	1
Tantilla olympia	+	1

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform.? = species known from indeterminate locality (see text).

Physiographic Regions of Central America										Totals
CG	HN	CRP	EP	YP	GH	NP	CGU	GCR	CP
Tantilla psittaca	+	1
Tantilla ruficeps	+	+	+	3
Tantilla stenigrammi	+	1
Tantilla taeniata	+	+	2
Tantilla tecta	+	1
Tantilla tritaeniata	+	1
Tantilla vermiformis	+	1
Trimorphodon quadruplex	+	+	+	+	+	5
Dipsadidae (77 species)
Adelphicos daryi	+	1
Adelphicos ibarrorum	+	1
Adelphicos veraepacis	+	1
Atractus darienensis	+	1
Atractus depressiocellus	+	1
Atractus hostilitractus	+	1
Atractus imperfectus	+	1
Chapinophis xanthocheilus	+	1
Coniophanes joanae	+	1
Crisantophis nevermanni	+	+	+	+	4
Cubophis brooksi	+	1
Dipsas articulata	+	+	+	3
Dipsas bicolor	+	+	+	3
Dipsas nicholsi	+	1
Dipsas tenuissima	+	1
Enulius bifoveatus	+	1
Enulius roatanensis	+	1
Geophis bellus	+	1
Geophis brachycephalus	+	+	+	3
Geophis championi	+	1
Geophis damiani	+	1
Geophis downsi	+	1
Geophis dunni	+	1
Geophis fulvoguttatus	+	1
Geophis godmani	+	1
Geophis hoffmanni	+	+	+	+	+	+	6
Geophis nephodrymus	+	1
Geophis ruthveni	+	+	2
Geophis talamancae	+	1
Geophis tectus	+	+	2
Geophis zeledoni	+	1
Hydromorphus concolor	+	+	+	+	+	+	+	7

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform.? = species known from indeterminate locality (see text).

Physiographic Regions of Central America										Totals
CG	HN	CRP	EP	YP	GH	NP	CGU	GCR	CP
Hydromorphus dunni	+	1
Imantodes phantasma	+	1
Leptodeira rhombifera	+	+	+	+	+	+	+	+	8
Leptodeira rubricata	+	1
Ninia celata	+	1
Ninia espinali	+	1
Ninia maculata	+	+	+	+	+	5
Ninia pavimentata	+	+	2
Ninia psephota	+	1
Omoadiphas aurula	+	1
Omoadiphas cannula	+	1
Omoadiphas texiguatensis	+	1
Rhadinaea calligaster	+	1
Rhadinaea pulveriventris	+	1
Rhadinaea sargenti	+	+	2
Rhadinaea stadelmani	+	1
Rhadinaea vermiculaticeps	+	+	+	3
Rhadinella anachoreta	+	+	+	+	4
Rhadinella hempsteadae	+	1
Rhadinella lisyae	+	1
Rhadinella montecristi	+	+	2
Rhadinella pegosalyta	+	1
Rhadinella pilonaorum	+	+	2
Rhadinella rogerromani	+	1
Rhadinella serperaster	+	1
Rhadinella tolpanorum	+	1
Sibon anthracops	+	+	+	3
Sibon argus	+	+	+	+	4
Sibon carri	+	+	+	3
Sibon lamari	+	1
Sibon longifrenis	+	+	+	+	4
Sibon manzanaresi	+	1
Sibon merendonensis	+	1
Sibon miskitus	+	1
Sibon noalamina	+	1
Sibon perissostichon	+	1
Trimetopon barbouri	+	+	+	3
Trimetopon gracile	+	1
Trimetopon pliolepis	+	+	2
Trimetopon simile	+	+	2
Trimetopon slevini	+	1

Table 4 (continued). Distribution of the 623 endemic herpetofaunal species in Central America among the 10 physiographic regions. Abbreviations for the regions are as follows: CG = western nuclear Central American highlands; CGU = Pacific lowlands from eastern Chiapas to south-central Guatemala; CP = Pacific lowlands from central Costa Rica through Panama (area includes associated Pacific islands); CRP = Isthmian Central American highlands; EP = highlands of eastern Panama; GCR = Pacific lowlands from southeastern Guatemala to northwestern Costa Rica; GH = Caribbean lowlands of eastern Guatemala and northern Honduras (area includes associated Caribbean islands); HN = eastern nuclear Central American highlands; NP = Caribbean lowlands from Nicaragua to Panama (area includes associated Caribbean islands); and YP = Yucatan Platform.? = species known from indeterminate locality (see text).

Physiographic Regions of Central America										Totals
CG	HN	CRP	EP	YP	GH	NP	CGU	GCR	CP
Trimetopon viquezi	+	1
Urotheca guentheri	+	+	+	+	+	5
Urotheca myersi	+	1
Urotheca pachyura	+	+	2
Elapidae (6 species)
Micrurus alleni	+	+	+	+	4
Micrurus hippocrepis	+	+	2
Micrurus mosquitensis	+	1
Micrurus ruatanus	+	1
Micrurus stewarti	+	+	+	3
Micrurus stuarti	+	1
Leptotyphlopidae (3 species)
Epictia ater	+	+	+	+	+	5
Epictia martinezi	+	1
Epictia pauldwyeri	+	1
Typhlopidae (3 species)
Amerotyphlops costaricensis	+	+	+	+	4
Amerotyphlops stadelmani	+	+	2
Typhlops tycherus	+	1
Viperidae (16 species)
Agkistrodon howardgloydi	+	+	+	+	4
Atropoides indomitus	+	1
Atropoides picadoi	+	+	+	3
Bothriechis guifarroi	+	1
Bothriechis lateralis	+	1
Bothriechis marchi	+	1
Bothriechis nigroviridis	+	1
Bothriechis nubestris	+	1
Bothriechis supraciliaris	+	+	2
Bothriechis thalassinus	+	1
Cerrophidion sasai	+	1
Cerrophidion wilsoni	+	1
Lachesis melanocephala	+	+	2
Lachesis stenophrys	+	+	2
Porthidium porrasi	+	1
Porthidium volcanicum	+	1
Testudines (2 species)
Geoemydidae (1 species)
Rhinoclemmys funerea	+	+	2
Kinosternidae (1 species)
Kinosternon angustipons	+	1

Fig. 3. Graph indicating the number and percentage of Central American endemic species in each of the 10 physiographic regions recognized. Table 5. Distributional summary of herpetofaunal families containing priority level one species in Central America, among the 10 physiographic regions. See Table 4 for explanation of abbreviations.

Families	Number of Species	CG	HN	CRP	Physiographic Regions EP YP GH NP				CGU	GCR	CP
Bufonidae	12	—	1	9	—	—	—	1	—	—	1
Centrolenidae	2	—	—	2	—	—	—	—	—	—	—
Craugastoridae	50	9	12	20	3	—	1	3	—	—	2
Dendrobatidae	10	—	—	2	3	—	—	4	—	—	1
Eleutherodactylidae	9	—	—	4	4	—	—	1	—	—	—
Hylidae	31	7	8	11	1	—	—	3	—	—	1
Leptodactylidae	1	—	1	—	—	—	—	—	—	—	—
Microhylidae	1	—	—	—	—	—	—	1	—	—	—
Pipidae	1	—	—	—	—	—	—	—	—	—	1
Ranidae	3	—	1	1	—	—	—	1	—	—	—
Subtotals	120	16	23	49	11	—	1	14	—	—	6
Plethodontidae	127	34	35	45	2	—	2	7	—	1	1
Subtotals	127	34	35	45	2	—	2	7	—	1	1
Caecilidae	3	—	—	—	—	—	—	—	—	—	3
Dermophiidae	2	—	—	2	—	—	—	—	—	—	—
Subtotals	5	—	—	2	—	—	—	—	—	—	3
Totals	252	50	58	96	13	—	3	21	—	1	10
Anguidae	23	9	6	5	1	—	—	1	—	—	1
Dactyloidae	46	2	14	22	—	—	4	1	—	—	3
Gymnophthalmidae	1	—	—	—	—	—	—	—	—	—	1

Table 5 (continued). Distributional summary of herpetofaunal families containing priority level one species in Central America, among the 10 physiographic regions. See Table 4 for explanation of abbreviations.

Families	Number of Species	CG	HN	CRP	Physiographic Regions EP YP GH NP				CGU	GCR	CP
Iguanidae	3	1	—	—	—	—	2	—	—	—	—
Mabuyidae	3	—	—	—	—	—	1	2	—	—	—
Phyllodactylidae	3	—	—	—	—	—	3	—	—	—	—
Sphaerodactylidae	11	—	—	—	2	—	6	1	—	—	2
Sphenomorphidae	1	—	—	—	—	—	—	1	—	—	—
Teiidae	2	—	—	—	—	—	—	1	—	—	1
Colubridae	15	2	4	2	1	1	3	1	—	1	—
Dipsadidae	48	4	13	14	4	—	5	3	—	—	5
Elapidae	3	1	—	—	—	—	1	1	—	—	—
Leptotyphlopidae	2	—	1	—	—	—	—	—	—	—	1
Typhlopidae	1	—	1	—	—	—	—	—	—	—	—
Viperidae	11	—	5	4	—	—	—	—	—	1	1
Subtotals	173	19	44	47	8	1	25	12	—	2	15
Kinosternidae	1	—	—	—	—	—	—	1	—	—	—
Subtotals	1	—	—	—	—	—	—	1	—	—	—
Totals	174	19	44	47	8	1	25	13	2	15
Sum Totals	426	69	102	143	21	1	28	34	—	3	25

The other five of the regional values lie below the mean figure, as follows: highlands of eastern Panama ( EP ; 37), Yucatan Platform ( YP ; 6), Pacific lowlands from eastern Chiapas to south-central Guatemala ( CGU ; 9), Pacific lowlands from southeastern Guatemala to northwestern Costa Rica ( GCR ; 39), and Pacific lowlands from Central Costa Rica through Panama (CP; 86). Even though these values are relatively low, collectively they amount to 177 species, 28.4% of the total of 623 endemic species; thus, they also are of considerable importance. The five regions containing the highest numbers of endemic species include three in highland and two in lowland areas. The five regions with the lowest numbers include one in highland and four in lowland areas. The numbers in the four highland regions range from 37 to 254, and in the six lowland regions from six to 86. Obviously, the 623 Central American endemic species are distributed unevenly throughout the 10 physiographic regions we recognize. In order to examine their distribution, we constructed a table indicating the total number of regions inhabited by the component species ( Table 6 ). The regions, listed in order of their total number of constituent species, range from six in the Yucatan Platform to 254 in the Isthmian Central American highlands. The number of physiographic regions occupied by these species ranges from one to eight, and their corresponding number of species also decreases markedly ( Table 6 ). Thus, 450 species occupy a single region, with the numbers ranging from one in the Yucatan Platform to 154 in the Isthmian Central American highlands; no single-region species are present in the Pacific lowlands from eastern Chiapas to south-central Guatemala . At the opposite extreme, a single species ( Leptodeira rhombifera ) occupies eight regions, and one species ( Hydromorphus concolor ) inhabits seven regions. The single-region species comprise the most speciose categories for seven of the 10 physiographic regions ( Table 6 ). The three exceptions are subhumid regions on the Atlantic (Yucatan Platform) and Pacific versants (Pacific lowlands from eastern Chiapas to south-central Guatemala and Pacific lowlands from southeastern Guatemala to northwestern Costa Rica ). The 450 single-region species comprise 72.2% of the 623 Central American endemic species. The 95 two-region species contribute 15.2% of the total number. Together, the single-region and two-region species constitute 545 taxa, 87.5% of the total. Thus, only 78 of the remaining species occupy from three to eight regions. This feature is of tremendous conservation significance for Central America, and we review this matter in greater detail below. Conservation Status of the Endemic Central American Herpetofauna In a previous paper on the Mexican endemic herpetofauna ( Johnson et al. 2017 ), we utilized the Environmental Vulnerability Score (EVS) system of conservation assessment. Along with various other authors, we have been involved with a series of papers published on the Mesoamerican herpetofauna since 2013 (see Johnson et al. 2017 , for a listing) including a recent paper on the herpetofauna of the Mexican state of Puebla ( Woolrich-Piña et al. 2017 ). Herein, we use the same system to evaluate the conservation status of the 623 species comprising the Central American herpetofauna. In calculating the EVS for these species, we used the scores included in Johnson et al. (2015) , supplemented by the scores we determined for the 43 species described since this paper was published. We placed these scores in Table 7 , incorporated them into those for the entire Central American endemic herpetofauna in Table 8 , and provide a graph of the data in Fig. 4 . Plate 10. Diasporus ventrimaculatus Chaves , García-Rodríguez, Mora, and Leal, 2009. This frog is a priority one species “known only from theValle del Silencio on the Caribbean versant of the Cordillera de Talamanca, Limon Province, Costa Rica” ( Frost 2018 ). This individual was observed in Valle del Silencio, Parque Internacional La Amistad, in the province of Limón, Costa Rica. Photo by Víctor Acosta-Chaves. Plate 11. Duellmanohyla rufioculis (Taylor, 1952) . This treefrog is a priority one species with an EVS of 14, which ranges on both “the Caribbean and Pacific slopes of the mountains of Costa Rica” ( Frost 2018 ). This individual came from Centro Soltis, San Isidro de Peñas Blancas, in the province of Alajuela, Costa Rica. Photo by Víctor Acosta-Chaves. Plate 12. Isthmohyla lancasteri (Barbour, 1928) . Lancaster’s Treefrog is a priority two species with an EVS of 14, which occurs in “the Cordillera de Talamanca of Costa Rica and western Panama” ( Frost 2018 ). This individual was found in Guayacán, in the province of Limón, Costa Rica. Photo by Víctor Acosta-Chaves. Plate 13. Plectrohyla pokomchi Duellman and Campbell, 1984 . The Rio Sanaja Spikethumb Frog is a priority eight species with an EVS of 13, which is distributed in “the Sierra de las Minas and the contiguous Sierra de Xucaneb in central and eastern Guatemala” ( Frost 2018 ).This individual came from Purulhá, in the department of Baja Verapaz, Guatemala. Photo by Andres Novales. Plate 14. Ptychohyla legleri (Taylor, 1958) . Legler’s Stream Frog is a priority one species with an EVS of 14, which is found on the “Pacific slopes of the Sierra de Talamanca [of] eastern Costa Rica and western Panama” ( Frost 2018 ). This individual was located in Alfombra de Pérez Zeledón, in the province of San José, Costa Rica. Photo by Víctor Acosta-Chaves. Plate 15. Smilisca puma (Cope, 1885) . The Tawny Smilisca is a priority one species with an EVS of 14, distributed on the “Caribbean lowlands of Costa Rica and adjacent Nicaragua” ( Frost 2018 ). This individual was encountered in La Selva Biological Station, in the province of Heredia, Costa Rica. Photo by Víctor Acosta-Chaves. Table 6. Number of endemic species in each of the 10 physiographic regions inhabited in Central America. See Table 4 for explanation of abbreviations.

Physiographic Regions	1	Number of endemic species in each physiographic region 2 3 4 5 6 7						8	Totals
YP	1	1	—	3	—	1	—	—	6
CGU	—	3	1	4	—	—	—	1	9
EP	23	4	1	6	1	—	1	—	36
GCR	4	8	10	6	6	5	2	1	42
CP	28	20	14	9	8	4	2	1	86
GH	28	25	12	11	11	5	2	1	95
CG	77	23	6	5	3	1	1	1	117
NP	36	35	22	13	11	5	2	1	125
HN	102	32	12	11	9	5	2	1	174
CRP	153	40	21	16	11	4	2	1	248
Totals	452	191	99	84	60	30	14	8	—

To illustrate the pattern of distribution of the EVS, we organized these scores by family in Table 9 . The data in this table indicate that the scores range from 10 to 20, out of a total theoretical range of 3 to 20. Thus, the scores occupy the entire range of medium vulnerability (10–13) and high vulnerability (14–20) in the EVS scale. None of the scores for these endemic species extend into the low vulnerability range (3–9). The highest score of 20 is found only among the anurans and, in particular, within the family Hylidae . This score is shared by six hylid species, including four species of Ecnomiohyla , one of Bromeliohyla , and one of Ptychohyla ( Table 8 ). The lowest score of 10 is seen in a broader range of herpetofaunal families ( Table 9 ), including the Hylidae (one species), Ranidae (one), Dactyloidae (one), Phrynosomatidae (one), and Leptotyphlopidae (one). The greatest number of species, i.e., 146, were assessed an EVS of 16, with species numbers decreasing more or less gradually on either side of this apex to both extremes, i.e., 10 and 20. Of the 623 total scores, 63 (10.1%) lie within the medium range and the remaining 560 (89.9%) in the high range ( Table 9 ). This large representation of high vulnerability species among the endemic species is of tremendous conservation significance, and figures prominently in the system of prioritization we present below. Priority Listing for Central American Endemic Herpetofaunal Species In prioritizing the conservation significance of the endemic herpetofaunal species in Central America, we used the same simple system developed by Johnson et al. (2017) . This system involves combining the data on physiographic distribution ( Table 4 ) and the Environmental Vulnerability Scores ( Table 8 ) for the 623 endemic species.This procedure resulted in the recognition of 14 priority levels, of which six are high vulnerability and eight are medium vulnerability groupings ( Table 10 ). We organized the high vulnerability species into six groups based on the number of physiographic regions they occupy, ranging from one to six ( Table 10 , Fig. 5 ). The numbers of species in these seven groups decrease markedly and consistently, as follows: Priority Level One (429 species); Priority Level Two (73); Priority Level Three (27); Priority Level Four (21); Priority Level Five (nine); and Priority Level Six (three). The most significant conclusion of this study is that 562 (90.2%) of the endemic species in Central America are allocated to the six high vulnerability groups. This proportion is 10 percentage points higher than the comparable figure (80.2%) for the Mexican endemic species ( Johnson et al., 2017 ). Furthermore, we believe that the difficulty of protecting these high vulnerability species increases with the fewer physiographic regions they occupy. Thus, the most critically vulnerable species are in the Priority One grouping, the 429 species that constitute 68.9% of the total number of Central American endemics. The challenge of protecting the high vulnerability species increases commensurately with the decrease in the priority level number. We arranged the medium vulnerability species into eight groups, also on the basis of the number of physiographic regions inhabited ( Table 10 , Fig. 5 ). Fewer species are included in these eight groups compared to the high vulnerability ones, as follows: Priority Level Seven (23); Priority Level Eight (21 species); Priority Level Nine (5); Priority Level Ten (four); Priority Level Eleven (four); Priority Level Twelve (two); Priority Level Thirteen (one); and Priority Level Fourteen (one). Even so, the next most important conclusion of this study is that these 61 species make up 9.8% of the total compendium of endemic species in Central America. The number of species in these eight groups also decreases sharply, as 69.8% of the 61 species fall into the first two priority levels, i.e., Seven and Eight. Table 7. Environmental Vulnerability Scores (EVS) for 43 endemic members of the Central American herpetofauna not included in Johnson et al. (2015) or requiring recalculation. Question marks indicate decisions made about reproductive mode based on phylogenetic relationships.

Environmental Vulnerability Score (EVS)
Taxa	Geographic Distribution	Ecological Distribution	Reproductive Mode/ Degree of Persecution	Total Score
Incilius majordomus	6	8	1?	15
Hyalinobatrachium dianae	5	7	3	15
Craugastor gabbi	5	8	4?	17
Diasporus dariensis	5	8	4?	17
Diasporus majeensis	6	8	4?	18
Diasporus pequeno	6	8	4?	18
Diasporus sapo	6	8	4?	18
Plectrohyla calvata	5	8	1	14
Smilisca manisorum	5	8	1?	14
Lithobates lenca	5	8	1	14
Bolitoglossa aurae	6	8	4?	18
Bolitoglossa chucutaniensis	6	8	4?	18
Cryptotriton xucaneborum	6	8	4?	18
Nototriton costaricense	6	8	4?	18
Nototriton nelson	6	7	4?	17
Nototriton oreadorum	6	8	4?	18
Oedipina berlini	5	8	4?	17
Oedipina capitalina	6	8	4?	18
Oedipina salvadorensis	5	8	4?	17
Celestus laf	6	8	3	17
Mesaspis cuchumatanus	5	7	3	15
Mesaspis salvadorensis	5	7	3	15
Dactyloa brooksi	5	7	3	15
Dactyloa kathydayae	6	8	3	17
Dactyloa maia	5	7	3	15
Dactyloa savage	5	7	3	15
Norops elcopeensis	5	7	3	15
Norops mccraniei	5	2	3	10
Norops oxylophus	5	6	3	14
Norops wilsoni	5	7	3	15
Lepidoblepharis emberawoundule	5	7	3	15
Lepidoblepharis rufigularis	6	8	3	17
Lepidoblepharis victormartinezi	5	7	3	15
Ameiva fuliginosa	5	8	3	16
Holcosus miadis	6	8	3	17
Tantilla berguidoi	6	8	2	16
Tantilla excelsa	5	6	2	13
Tantilla gottei	5	7	2	14
Tantilla stenigrammi	5	8	2	15
Rhadinella lisyae	6	7	2	15
Epictia martinezi	6	8	1	15
Epictia pauldwyeri	5	8	1	14
Bothriechis nubestris	5	7	5	17

Plate 16. Agalychnis annae (Duellman, 1963) . The Orange-eyed Treefrog is a priority two species with an EVS of 15, with a distribution in the “Northern Cordillera de Talamanca, Cordillera de Tilarán and Cordillera Central of Costa Rica” ( Frost 2018 ). This individual was found in Heredia, Costa Rica. Photo by Víctor Acosta-Chaves. Plate 17. Agalychnis saltator Taylor, 1955 . This leaf frog is a priority three species with an EVS of 14, which ranges along the “Caribbean lowlands of northeastern Honduras, Nicaragua, to east-central Costa Rica” ( Frost 2018 ). This individual was located in Centro Soltis, San Isidro de Peñas Blancas, in the province of Alajuela, Costa Rica. Photo by Víctor Acosta-Chaves. Plate 18. Lithobates taylori (Smith, 1959) . The Peralta Frog is a priority eight species with an EVS of 12, distributed “at scattered localities on the humid Atlantic lowlands from eastern Nicaragua to southeastern Costa Rica and in the humid premontane and lower montane areas of upland Costa Rica, including the Meseta Oriental and Meseta Occidental and probably the Cordillera Central” (Savage 2002: 402). This individual was found in a pond at Llano Tugrí, in the Serranía de Tabasará. Photo by Abel Batista. Table 8. Environmental Vulnerability Scores (EVS) for the endemic members of the herpetofauna of Central America.

Taxa	EVS	Taxa	EVS
Atelopus certus	14	Oedipina ignea	15
Atelopus chiriquiensis	14	Oedipina kasios	16
Atelopus chirripoensis	15	Oedipina koehleri	16
Atelopus limosus	14	Oedipina leptopoda	17
Atelopus senex	13	Oedipina maritima	18
Atelopus varius	11	Oedipina motaguae	18
Atelopus zeteki	13	Oedipina nica	17
Incilius aucoinae	14	Oedipina nimaso	18
Incilius chompipe	13	Oedipina pacificensis	16
Incilius epioticus	16	Oedipina paucidentata	18
Incilius fastidiosus	13	Oedipina petiola	18
Incilius guanacaste	17	Oedipina poelzi	16
Incilius holdridgei	14	Oedipina pseudouniformis	16
Incilius ibarrai	13	Oedipina quadra	17
Incilius karenlipsae	15	Oedipina salvadorensis	17
Incilius leucomyos	12	Oedipina savagei	18
Incilius majordomus	15	Oedipina stenopodia	17
Incilius melanochlorus	12	Oedipina stuarti	15
Incilius periglenes	15	Oedipina taylori	14
Incilius peripatetes	14	Oedipina tomasi	18
Incilius porter	14	Oedipina tzutujilorum	18
Incilius signifier	14	Oedipina uniformis	15
Rhinella centralis	14	Pseudoeurycea exspectata	18
Rhinella chrysophora	13	Caecilia volcani	17
Cochranella granulosa	15	Oscaecilia elongata	19
Hyalinobatrachium dianae	15	Oscaecilia osae	19
Hyalinobatrachium talamancae	16	Dermophis costaricensis	18
Hyalinobatrachium vireovittatum	16	Dermophis gracilior	18
Craugastor adamastus	18	Dermophis occidentalis	17
Craugastor anciano	16	Gymnopis multiplicata	14
Craugastor andi	17	Abronia anzuetoi	18
Craugastor angelicus	15	Abronia aurita	16
Craugastor aphanus	17	Abronia campbelli	18
Craugastor aurilegulus	15	Abronia fimbriata	16
Craugastor azueroensis	16	Abronia frosti	18
Craugastor bocourti	16	Abronia gaiophantasma	16
Craugastor bransfordii	13	Abronia meledona	18
Craugastor campbelli	16	Abronia montecristoi	17
Craugastor catalinae	17	Abronia salvadorensis	17
Craugastor chac	16	Abronia vasconcelosii	16
Craugastor charadra	15	Celestus adercus	17
Craugastor chingopetaca	18	Celestus atitlanensis	15
Craugastor chrysozetetes	18	Celestus bivittatus	15
Craugastor coffeus	18	Celestus cyanochloris	14
Craugastor cruzi	18	Celestus hylaius	16

Table 8 (continued). Environmental Vulnerability Scores (EVS) for the endemic members of the herpetofauna of Central America.

Taxa	EVS	Taxa	EVS
Craugastor cuaquero	18	Celestus laf	17
Craugastor cyanochthebius	18	Celestus montanus	15
Craugastor daryi	17	Celestus orobius	16
Craugastor emcelae	17	Celestus scansorius	15
Craugastor emleni	15	Coloptychon rhombifer	16
Craugastor epochthidius	16	Diploglossus bilobatus	16
Craugastor escoces	15	Diploglossus montisilvestris	18
Craugastor evanesce	17	Mesaspis cuchumatanus	15
Craugastor fecundus	16	Mesaspis monticola	14
Craugastor fleischmanni	16	Mesaspis salvadorensis	15
Craugastor gabbi	17	Basiliscus plumifrons	15
Craugastor gollmeri	16	Dactyloa brooksi	15
Craugastor gulosus	17	Dactyloa casildae	16
Craugastor inachus	17	Dactyloa ginaelisae	12
Craugastor jota	18	Dactyloa ibanezi	15
Craugastor laevissimus	12	Dactyloa insignis	14
Craugastor lauraster	16	Dactyloa kathydayae	17
Craugastor megacephalus	16	Dactyloa kunayalae	15
Craugastor melanostictus	16	Dactyloa maia	15
Craugastor merendonensis	18	Dactyloa microtus	15
Craugastor milesi	16	Dactyloa savagei	15
Craugastor mimus	16	Norops alocomyos	16
Craugastor monnichorum	16	Norops altae	15
Craugastor myllomyllon	18	Norops amplisquamosus	17
Craugastor nefrens	18	Norops apletophallus	15
Craugastor noblei	16	Norops aquaticus	15
Craugastor obesus	17	Norops benedikti	16
Craugastor olanchano	18	Norops bicaorum	17
Craugastor omoaensis	18	Norops campbelli	17
Craugastor pechorum	16	Norops carpenteri	16
Craugastor persimilis	16	Norops charlesmyersi	16
Craugastor phasma	18	Norops cobanensis	13
Craugastor podiciferus	15	Norops cryptolimifrons	16
Craugastor polyptychus	17	Norops cupreus	13
Craugastor punctariolus	16	Norops cusuco	17
Craugastor ranoides	15	Norops datzorum	15
Craugastor rayo	16	Norops elcopeensis	15
Craugastor rhyacobatrachus	16	Norops fortunensis	17
Craugastor rivulus	17	Norops fungosus	15
Craugastor rostralis	16	Norops gruuo	17
Craugastor rugosus	16	Norops haguei	17
Craugastor sabrinus	16	Norops heteropholidotus	16
Craugastor saltuarius	18	Norops humilis	14
Craugastor sandersoni	18	Norops intermedius	14
Craugastor stadelmani	16	Norops johnmeyeri	16

Table 8 (continued). Environmental Vulnerability Scores (EVS) for the endemic members of the herpetofauna of Central America.

Taxa	EVS	Taxa	EVS
Craugastor stejnegerianus	14	Norops kemptoni	15
Craugastor tabasarae	17	Norops kreutzi	17
Craugastor talamancae	17	Norops leditzigorum	15
Craugastor taurus	17	Norops limifrons	15
Craugastor trachydermus	18	Norops lionotus	14
Craugastor underwoodi	16	Norops loveridgei	14
Craugastor xucanebi	16	Norops macrophallus	15
Pristimantis adnus	18	Norops magnaphallus	17
Pristimantis altae	16	Norops marsupialis	16
Pristimantis caryophyllaceus	15	Norops mccraniei	10
Pristimantis cerasinus	16	Norops monteverde	17
Pristimantis museosus	17	Norops morazani	17
Pristimantis pardalis	17	Norops muralla	17
Pristimantis pirrensis	18	Norops ocelloscapularis	15
Strabomantis laticorpus	17	Norops osa	16
Ameerega maculate	18	Norops oxylophus	14
Andinobates claudiae	18	Norops pachypus	15
Andinobates geminisae	18	Norops pijolensis	16
Colostethus latinasus	15	Norops polylepis	15
Ectopoglossus astralogaster	18	Norops pseudokemptoni	17
Ectopoglossus isthminus	16	Norops pseudopachypus	17
Oophaga arborea	16	Norops purpurgularis	16
Oophaga granulifera	17	Norops quaggulus	15
Oophaga pumilio	16	Norops roatanensis	17
Oophaga speciosa	16	Norops rubribarbaris	17
Oophaga vicentei	16	Norops salvini	15
Phyllobates lugubris	17	Norops sminthus	15
Phyllobates vittatus	17	Norops tenorioensis	17
Silverstoneia flotator	16	Norops townsendi	17
Diasporus citrinobapheus	17	Norops triumphalis	17
Diasporus darienensis	17	Norops tropidolepis	15
Diasporus diastema	15	Norops utilensis	17
Diasporus hylaeformis	17	Norops villai	17
Diasporus igneus	18	Norops wampuensis	17
Diasporus majeensis	18	Norops wellbornae	15
Diasporus pequeno	18	Norops wermuthi	16
Diasporus sapo	18	Norops wilsoni	15
Diasporus tigrillo	18	Norops woodi	14
Diasporus ventrimaculatus	18	Norops yoroensis	15
Atlantihyla panchoi	13	Norops zeus	15
Atlantihyla spinipollex	12	Coleonyx mitratus	14
Bromeliohyla melacaena	20	Bachia blairi	15
Dryophytes bocourti	14	Echinosaura panamensis	14
Duellmanohyla legleri	14	Echinosaura apodema	15
Duellmanohyla lythrodes	14	Heloderma charlesbogerti	18
Duellmanohyla rufioculis	14	Ctenosaura bakeri	19

Table 8 (continued). Environmental Vulnerability Scores (EVS) for the endemic members of the herpetofauna of Central America.

Taxa	EVS	Taxa	EVS
Duellmanohyla salvadorensis	12	Ctenosaura flavidorsalis	18
Duellmanohyla salvavida	13	Ctenosaura melanosterna	18
Duellmanohyla soralia	12	Ctenosaura oedirhina	19
Duellmanohyla uranochroa	12	Ctenosaura palearis	19
Ecnomiohyla bailarina	20	Ctenosura praeocularis	18
Ecnomiohyla fimbrimembra	19	Ctenosaura quinquecarinata	19
Ecnomiohyla minera	18	Marisora alliacea	15
Ecnomiohyla rabborum	20	Marisora magnacornae	17
Ecnomiohyla salvaje	19	Marisora roatanae	16
Ecnomiohyla sukia	18	Marisora unimarginata	15
Ecnomiohyla thysanota	20	Sceloporus lunaei	15
Ecnomiohyla veraguensis	20	Sceloporus malachiticus	10
Exerodonta catracha	14	Phyllodactylus insularis	17
Exerodonta perkinsi	15	Phyllodactylus palmeus	16
Hyloscirtus colymba	13	Phyllodactylus paralepis	17
Isthmohyla angustilineata	13	Mesoscincus managuae	14
Isthmohyla calypso	16	Lepidoblepharis emberawoundule	15
Isthmohyla debilis	14	Lepidoblepharis rufigularis	17
Isthmohyla graceae	13	Lepidoblepharis victormartinezi	15
Isthmohyla infucata	14	Sphaerodactylus alphus	17
Isthmohyla insolita	17	Sphaerodactylus dunni	15
Isthmohyla lancasteri	14	Sphaerodactylus graptolaemus	16
Isthmohyla picadoi	19	Sphaerodactylus guanaje	17
Isthmohyla pictipes	14	Sphaerodactylus homolepis	16
Isthmohyla pseudopuma	13	Sphaerodactylus leonardovaldesi	16
Isthmohyla rivularis	13	Sphaerodactylus millepunctatus	15
Isthmohyla tica	13	Sphaerodactylus pacificus	17
Isthmohyla xanthosticta	15	Sphaerodactylus poindexteri	17
Isthmohyla zeteki	18	Sphaerodactylus rosaurae	16
Plectrohyla calvata	14	Scincella rara	17
Plectrohyla chrysopleura	13	Cnemidophorus duellmani	16
Plectrohyla dasypus	14	Cnemidophorus ruatanus	15
Plectrohyla exquisita	15	Holcosus leptophrys	16
Plectrohyla glandulosa	12	Holcosus miadis	17
Plectrohyla pokomchi	13	Holcosus quadrilineatus	16
Plectrohyla psiloderma	14	Lepidophyma mayae	13
Plectrohyla quecchi	13	Lepidophyma reticulatum	13
Plectrohyla tecunumani	14	Helminthophis frontalis	12
Plectrohyla teuchestes	15	Dendrophidion apharocybe	16
Ptychohyla dendrophasma	20	Dendrophidion crybelum	17
Ptychohyla hypomykter	10	Dendrophidion paucicarinatum	16
Quilticohyla sanctaecrucis	14	Dendrophidion rufiterminorum	16
Scinax altae	14	Drymobius melanotropis	16
Smilisca manisorum	14	Leptodrymus pulcherrimus	13
Smilisca puma	14	Leptophis nebulosus	14
Leptodactylus silvanimbus	14	Mastigodryas alternatus	12

Table 8 (continued). Environmental Vulnerability Scores (EVS) for the endemic members of the herpetofauna of Central America.

Taxa	EVS	Taxa	EVS
Hypopachus pictiventris	14	Mastigodryas dorsalis	14
Agalychnis annae	15	Oxybelis wilsoni	17
Agalychnis saltatory	14	Scolecophis atrocinctus	13
Pipa myersi	17	Tantilla albiceps	16
Lithobates juliani	12	Tantilla armillata	11
Lithobates lenca	14	Tantilla bairdi	16
Lithobates miadis	15	Tantilla berguidoi	16
Lithobates taylori	12	Tantilla brevicauda	13
Lithobates vibicarius	14	Tantilla excelsa	13
Lithobates warszewitschii	10	Tantilla gottei	14
Bolitoglossa alvaradoi	16	Tantilla hendersoni	16
Bolitoglossa anthracina	18	Tantilla jani	14
Bolitoglossa aurae	18	Tantilla lempira	14
Bolitoglossa aureogularis	18	Tantilla olympia	16
Bolitoglossa bramei	17	Tantilla psittaca	15
Bolitoglossa carri	18	Tantilla ruficeps	12
Bolitoglossa cataguana	18	Tantilla stenigrammi	15
Bolitoglossa celaque	17	Tantilla taeniata	14
Bolitoglossa centenorum	18	Tantilla tecta	16
Bolitoglossa cerroensis	16	Tantilla tritaeniata	16
Bolitoglossa chucantiensis	18	Tantilla vermiformis	14
Bolitoglossa colonnea	16	Trimorphodon quadruplex	14
Bolitoglossa compacta	17	Adelphicos daryi	16
Bolitoglossa conanti	16	Adelphicos ibarrorum	15
Bolitoglossa copia	18	Adelphicos veraepacis	14
Bolitoglossa cuchumatana	14	Atractus darienensis	16
Bolitoglossa cuna	17	Atractus depressiocellus	15
Bolitoglossa daryorum	17	Atractus hostilitractus	16
Bolitoglossa decora	18	Atractus imperfectus	16
Bolitoglossa diaphora	18	Chapinophis xanthocheilus	16
Bolitoglossa diminuta	18	Coniophanes joanae	15
Bolitoglossa dofleini	15	Crisantophis nevermanni	16
Bolitoglossa dunni	16	Cubophis brooksi	14
Bolitoglossa epimela	17	Dipsas articulata	15
Bolitoglossa eremia	18	Dipsas bicolor	17
Bolitoglossa gomezi	16	Dipsas nicholsi	15
Bolitoglossa gracilis	18	Dipsas tenuissima	14
Bolitoglossa heiroreias	17	Enulius bifoveatus	16
Bolitoglossa helmrichi	16	Enulius roatanensis	16
Bolitoglossa huehuetenanguensis	18	Geophis bellus	16
Bolitoglossa indio	17	Geophis brachycephalus	11
Bolitoglossa insularis	18	Geophis championi	16
Bolitoglossa jacksoni	18	Geophis damiani	16
Bolitoglossa jugivagans	18	Geophis downsi	16
Bolitoglossa kamuk	18	Geophis dunni	16

Table 8 (continued). Environmental Vulnerability Scores (EVS) for the endemic members of the herpetofauna of Central America.

Taxa	EVS	Taxa	EVS
Bolitoglossa kaqchikelorum	17	Geophis fulvoguttatus	14
Bolitoglossa la	17	Geophis godmani	14
Bolitoglossa lignicolor	16	Geophis hoffmanni	12
Bolitoglossa longissima	18	Geophis nephodrymus	16
Bolitoglossa magnifica	16	Geophis ruthveni	14
Bolitoglossa marmorea	17	Geophis talamancae	15
Bolitoglossa meliana	16	Geophis tectus	13
Bolitoglossa minutula	17	Geophis zeledoni	15
Bolitoglossa mombachoensis	17	Hydromorphus concolor	12
Bolitoglossa morio	13	Hydromorphus dunni	16
Bolitoglossa nigrescens	16	Imantodes phantasma	16
Bolitoglossa ninadormida	18	Leptodeira rhombifera	12
Bolitoglossa nussbaumi	18	Leptodeira rubricata	17
Bolitoglossa nympha	16	Ninia celata	15
Bolitoglossa obscura	18	Ninia espinali	14
Bolitoglossa odonnelli	16	Ninia maculata	12
Bolitoglossa omniumsanctorum	16	Ninia pavimentata	15
Bolitoglossa oresbia	17	Ninia psephota	13
Bolitoglossa pacaya	17	Omoadiphas aurula	16
Bolitoglossa pesrubra	15	Omoadiphas cannula	16
Bolitoglossa porrasorum	16	Omoadiphas texiguatensis	16
Bolitoglossa psephena	18	Rhadinaea calligaster	14
Bolitoglossa pygmaea	17	Rhadinaea pulveriventris	14
Bolitoglossa robinsoni	16	Rhadinaea sargenti	14
Bolitoglossa robusta	16	Rhadinaea stadelmani	13
Bolitoglossa salvinii	16	Rhadinaea vermiculaticeps	15
Bolitoglossa schizodactyla	15	Rhadinella anachoreta	14
Bolitoglossa sombra	16	Rhadinella hempsteadae	13
Bolitoglossa sooyorum	16	Rhadinella lisyae	15
Bolitoglossa splendida	18	Rhadinella montecristi	14
Bolitoglossa striatula	16	Rhadinella pegosalyta	16
Bolitoglossa subpalmata	15	Rhadinella pilonaorum	15
Bolitoglossa suchitanensis	18	Rhadinella rogerromani	16
Bolitoglossa synoria	17	Rhadinella serperaster	13
Bolitoglossa taylori	17	Rhadinella tolpanorum	16
Bolitoglossa tenebrosa	17	Sibon anthracops	15
Bolitoglossa tica	17	Sibon argus	16
Bolitoglossa tzultacaj	18	Sibon carri	14
Bolitoglossa xibalba	17	Sibon lamari	16
Bolitoglossa zacapensis	18	Sibon longifrenis	14
Cryptotriton monzoni	18	Sibon manzanaresi	15
Cryptotriton nasalis	18	Sibon merendonensis	16
Cryptotriton necopinus	18	Sibon miskitus	15
Cryptotriton sierraminensis	17	Sibon noalamina	15
Cryptotriton veraepacis	17	Sibon perissostichon	16

Table 8 (continued). Environmental Vulnerability Scores (EVS) for the endemic members of the herpetofauna of Central America.

Taxa	EVS	Taxa	EVS
Cryptotriton xucaneborum	18	Trimetopon barbouri	15
Dendrotriton bromeliacius	17	Trimetopon gracile	14
Dendrotriton chujorum	18	Trimetopon pliolepis	12
Dendrotriton cuchumatanus	18	Trimetopon simile	13
Dendrotriton kekchiorum	18	Trimetopon slevini	14
Dendrotriton rabbi	17	Trimetopon viquezi	15
Dendrotriton sanctibarbarus	18	Urotheca guentheri	12
Nototriton abscondens	16	Urotheca myersi	15
Nototriton barbouri	16	Urotheca pachyura	14
Nototriton brodiei	17	Micrurus alleni	16
Nototriton costaricense	18	Micrurus hippocrepis	18
Nototriton gamezi	18	Micrurus mosquitensis	17
Nototriton guanacaste	17	Micrurus ruatanus	18
Nototriton lignicola	18	Micrurus stewarti	17
Nototriton limnospectator	17	Micrurus stuarti	17
Nototriton major	18	Epictia ater	10
Nototriton matama	18	Epictia martinezi	15
Nototriton mime	18	Epictia pauldwyeri	14
Nototriton nelsoni	17	Amerotyphlops costaricensis	11
Nototriton oreadorum	18	Amerotyphlops stadelmani	12
Nototriton picadoi	16	Typhlops tycherus	14
Nototriton picucha	18	Agkistrodon howardgloydi	17
Nototriton richardi	16	Atropoides indomitus	18
Nototriton saslaya	18	Atropoides picadoi	16
Nototriton stuarti	18	Bothriechis guifarroi	19
Nototriton tapanti	18	Bothriechis lateralis	16
Nototriton tomamorum	18	Bothriechis marchi	16
Oedipina alfaroi	16	Bothriechis nigroviridis	17
Oedipina alleni	16	Bothriechis nubestris	17
Oedipina altura	18	Bothriechis supraciliaris	17
Oedipina berlini	17	Bothriechis thalassinus	17
Oedipina capitalina	18	Cerrophidion sasai	16
Oedipina carablanca	18	Cerrophidion wilsoni	15
Oedipina chortiorum	18	Lachesis melanocephala	17
Oedipina collaris	17	Lachesis stenophrys	17
Oedipina cyclocauda	15	Porthidium porrasi	18
Oedipina fortunensis	18	Porthidium volcanicum	18
Oedipina gephyra	17	Rhinoclemmys funerea	16
Oedipina gracilis	16	Kinosternon angustipons	16
Oedipina grandis	17

Fig. 4. Graph showing Central American endemic species and their corresponding Environmental Vulnerability Scores (EVS) ranging from 10 to 20. Fig. 5. Graph of Central American endemic species allocated to the 14 conservation priority groups. When we examined the 623 endemic species relative to the number of physiographic regions inhabited, the results are as follows: one region (429+23 = 452); two regions (73+21 = 94); three regions (27+5 = 32); four regions (21+4 = 25); five regions (9+4 = 13); six regions (3+2 = 5); seven regions (1); and eight regions (1). Perusal of these data supports another conclusion, i.e., that 72.6% of the total number of species occupy a single physiographic region. Based on the assumptions of this study, these 452 species can be expected to offer the major challenge in efforts to protect the endemic component of the Central American herpetofauna. The next most challenging group contains the 94 species occupying two regions. Together, the singlegroup and double-group species comprise 546 (87.6%) of the total of 623 Central American endemic species. Our analysis in this paper indicates that most of the 623 endemic CentralAmerican herpetofaunal species are judged as high vulnerability based on the EVS methodology, and are demonstrated to occupy relatively few physiographic regions (one or two). The endemic component of the Central American herpetofauna, just as with the Mexican endemic component ( Johnson et al. 2017 ), is of global significance and constitutes the most significant challenge to conservation professionals working within this segment of the Mesoamerican herpetofauna. Johnson et al. (2017) arrived at the same conclusion in their work on the Mexican endemic herpetofauna. Considered as a whole, the Mesoamerican endemic herpetofauna comprises the 789 Mexican endemic species dealt with by Johnson et al. (2017) and the 623 Central American endemic species dealt with here, as well as the 225 species restricted in distribution to Mexico and Central America (i.e., Mesoamerica; Wilson et al., 2017 ) for a total of 1,637 species. This figure represents more than three quarters of the entire Mesoamerican herpetofauna ( Wilson et al., 2017 ). We examine the parameters of the challenge facing conservation biologists working in Central America in the following section. Prognosis for the Endemic Central American Herpetofauna The same environmental issues impacting the Mexican endemic herpetofauna, as discussed by Johnson et al. (2017) , also impinge upon the Central American endemic herpetofauna. In light of this situation, we emphasize that the survival of the 623 endemic species inhabiting Central America ultimately depends on addressing the underlying issues that lead to all environmental problems, including biodiversity decline, that in turn stand in the way of designing a sustainable existence for humanity’s tenure on Earth. Johnson et al. (2017: 609) explained what we face as follows: “Fundamentally, humans have created and maintain these environmental problems because of their capacity for rational thought, i.e., their ability to connect cause to effect through the passing of time, and adopting an anthropocentric worldview that stresses the exploitation of the world’s resources to support the burgeoning human population. Such a worldview contrasts markedly with that of environmentalists, who have adopted ‘a worldview that helps us make sense of how the environment works, our place in the environment, and right and wrong environmental behaviors’ ( Raven and Berg 2004 : G-6). Obviously, the present anthropocentric worldview held by most people represents the fundamental reason why these environmental problems exist, and continued human population growth allows them to worsen over time.” The anthropocentric worldview, also known as the Western worldview, “includes human superiority and dominance over nature, the unrestricted use of natural resources, increased economic growth to manage an expanding industrial base, the inherent rights of individuals, and accumulation of wealth and unlimited consumption of goods and services to provide material comforts” ( Raven and Berg 2004: 17 ). This worldview not only creates the entire spectrum of environmental problems, but also the entire panoply of human societal issues we see played out every day in various media outlets. Ultimately, they arise from a commitment to discriminate among groups of people, i.e., on the basis of racial background, gender, religion, economic wealth, political persuasion, and so forth. Thus, not only is humanity poised against the rest of the living world, but also varying groups of humans are in conflict with one another. As the focus of humanity decreases from larger to increasingly smaller realms of interest, it can be argued that mental stability gives way to instability, and eventually gives rise to the increased incidence of the narcissistic personality disorder (NPD). This disorder is highly variable in presentation and can manifest across a broad spectrum of severity, but is generally characterized by pervasive grandiosity, an excessive need for admiration, and a lack of empathy ( Caligor et al. 2015 ). Envisioning NPD as an extreme end-point of the intensification of anthropocentrism might explain why the potential causes of this disorder remain unknown and that clinical guidelines have yet to emerge ( Caligor et al. 2015 ). Given that none of the authors of this paper possesses credentials in psychology or psychiatry, our idea about the connection between the anthropocentric worldview and the narcissistic personality disorder can be best understood as a hypothesis remaining to be tested, hopefully by a crossdiscipline team of environmental scientists, deep ecology philosophers, and biocentric psychologists/psychiatrists. Studying such a connection could lie within the realm of environmental psychology, defined as an interdisciplinary field that focuses on the interplay between environments and human cognition and behavior; considering the term “environment” broadly, including both natural and humanmade environments ( De Young 2013 ). Since its conception, research in environmental psychology has often targeted human attitudes towards the natural environment, and current trends are now shifting to a focus on sustainable living in the context of environmental issues ( De Young 2013 ). Table 9. Summary of EVS values for Central American endemic species, arranged by family. Shaded area encompasses high vulnerability scores.

Number	Environmental Vulnerability Scores
Families	of Species	10	11	12	13	14	15	16	17	18	19	20
Bufonidae	24	—	1	2	6	9	4	1	1	—	—	—
Centrolenidae	4	—	—	—	—	—	2	2	—	—	—	—
Craugastoridae	77	—	—	1	1	1	8	29	18	19	—	—
Dendrobatidae	14	—	—	—	—	—	1	6	3	4	—	—
Eleutherodactyludae	10	—	—	—	—	—	1	—	3	6	—	—
Hylidae	52	1	—	5	11	17	4	1	1	3	3	6
Leptodactylidae	1	—	—	—	—	1	—	—	—	—	—	—
Microhylidae	1	—	—	—	—	1	—	—	—	—	—	—
Phyllomedusidae	2	—	—	—	—	1	1	—	—	—	—	—
Pipidae	1	—	—	—	—	—	—	—	1	—	—	—
Ranidae	6	1	—	2	—	2	1	—	—	—	—	—
Subtotals	192	2	1	10	18	32	22	39	27	32	3	6
Plethodontidae	143	—	—	—	1	2	8	33	38	61	—	—
Subtotals	143	—	—	—	1	2	8	33	38	61	—	—
Caeciliidae	3	—	—	—	—	—	—	—	1	2	—
Dermophiidae	4	—	—	—	—	1	—	—	1	2	—	—
Subtotals	7	—	—	—	—	1	—	—	2	2	2	—
Totals	342	2	1	10	19	35	30	72	67	95	5	6
Anguidae	25	—	—	—	—	2	6	8	4	5	—	—
Corytophanidae	1	—	—	—	—	—	1	—	—	—	—	—
Dactyloidae	74	1	—	1	2	7	27	13	23	—	—	—
Eublepharidae	1	—	—	—	—	1	—	—	—	—	—	—
Gymnophthalmidae	3	—	—	—	—	1	2	—	—	—	—	—
Helodermatidae	1	—	—	—	—	—	—	—	—	1	—	—
Iguanidae	7	—	—	—	—	—	—	—	—	3	4	—
Mabuyidae	4	—	—	—	—	—	2	1	1	—	—	—
Phrynosomatidae	2	1	—	—	—	—	1	—	—	—	—	—
Phyllodactylidae	3	—	—	—	—	—	—	1	2	—	—	—
Scincidae	1	—	—	—	—	1	—	—	—	—	—	—
Sphaerodactylidae	13	—	—	—	—	—	4	4	5	—	—	—
Sphenomorphidae	1	—	—	—	—	—	—	—	1	—	—	—
Teiidae	5	—	—	—	—	—	1	3	1	—	—	—
Xantusiidae	2	—	—	—	2	—	—	—	—	—	—	—
Subtotals	143	2	—	1	4	12	44	30	37	9	4	—
Anomalepididae	1	—	—	1	—	—	—	—	—	—	—	—
Colubridae	30	—	1	2	4	8	2	11	2	—	—	—
Dipsadidae	77	—	1	6	6	17	19	26	2	—	—	—
Elapidae	6	—	—	—	—	—	—	1	3	2	—	—
Leptotyphlopidae	3	1	—	—	—	1	1	—	—	—	—	—
Typhlopidae	3	—	1	1	—	1	—	—	—	—	—	—
Viperidae	16	—	—	—	—	—	1	4	7	3	1
Subtotals	136	1	3	10	10	27	23	42	14	5	1
Geoemydidae	1	—	—	—	—	—	—	1	—	—	—	—

Table 9 (continued). Summary of EVS values for Central American endemic species, arranged by family. Shaded area encompasses high vulnerability scores.

Number	Environmental Vulnerability Scores
Families	of Species	10	11	12	13	14	15	16	17	18	19	20
Kinosternidae	1	—	—	—	—	—	—	1	—	—	—	—
Subtotals	2	—	—	—	—	—	—	2	—	—	—	—
Totals	281	3	3	11	14	39	67	74	51	14	5	—
Sum Totals	623	5	4	21	33	74	97	146	118	109	10	6
Category Totals	623	63		560

Plate 19. Lithobates warszewitschii (Schmidt, 1857) . Warszewitsch’s Frog is a priority eleven species with an EVS of 10, found on “the Atlantic versant from northeastern Honduras to central Panama, both slopes of the cordilleras of Costa Rica and western Panama, lowlands of southwestern Costa Rica, and eastern Panama and gallery forests in nonpeninsular northwestern Costa Rica” (Savage 2002: 405). This individual came from Nectandra Reserve, in the province of Alajuela, Costa Rica. Photo by Sean Michael Rovito. Plate 20. Bolitoglossa alvaradoi Taylor, 1954 . The Moravia de Chirripó Salamander is a priority two species with an EVS of 16, distributed on “the Atlantic versant of Costa Rica” ( Frost, 2018 ). This individual was found in Veragua Rainforest, in the province of Limón, Costa Rica. P hoto by Víctor Acosta-Chaves. Plate 21. Bolitoglossa centenorum Campbell, Smith, Streicher, Acevedo, and Brodie, 2010 . This salamander is a priority one species with an EVS of 18, which is known “only from the type locality in the Sierra Cuchumatanes, Huehuetenango, Guatemala” ( Frost 2018 ). This individual was encountered at the type locality, near San Mateo Ixtatán. Photo by Todd Pierson. Finding lasting solutions to environmental problems must be based on a realistic, fact-based approach that evaluates the symptoms of these problems until their causes are identified ( Wilson and McCranie 2004 ). Often, the search for an ultimate cause stops when exposed to the anthropocentric worldview. A worldview, however, is a collection of basic values that “help us to make sense of the world, understand our place in it, and determine right and wrong behaviors” ( Raven and Berg 2004 : G-17). How the values that characterize the anthropocentric worldview have arisen through the evolution of human behavior to become predominant, however, generally has not been explored. Our working assumption, i.e., our hypothesis, is that the ultimate causes are deeply engrained in the origins of human behavior and have become so pervasive as to underlie our efforts to understand our world, and our place in it. Even a discipline called environmental psychology might not expose the steps in behavioral evolution that would allow present-day humans to address the malady known as the anthropocentric worldview. In particular, this viewpoint is evident when considering that environmental psychology adopts a broad array of theories, methods, and interpretations from other disciplines as needed, and this mosaic approach can make it difficult to understand the field as a whole and the role it might play in these societal issues moving forward. An encouraging sign is the recent emergence of even more specific sub-fields, such as conservation psychology and ecopsychology, which aim to provide solutions or interventions for problems specifically related to conservation of the natural world ( Steg and Vlek 2009 ; De Young 2013 ). There is clearly a critical need to develop novel approaches for studying animal behavior and human psychology that emphasize reasons why the anthropocentric worldview has become so predominant, and what needs to be done to replace it with the environmental worldview. If, as we hypothesized, there is a psychological connection among centrist forms of thinking at larger scales (i.e., the anthropocentric worldview) and those at smaller scales (i.e., narcissism), then we are faced with an even greater challenge than commonly is envisioned. Searching for Ultimate Solutions Johnson et al. (2017:613) offered some ideas about searching for ultimate solutions to the problem of biodiversity decline, based on opinions promulgated by Wilson and Townsend (2010), Wilson (2016) , and Kopnina (2016) , and concluded as follows: “Our opinion is that humans have the rational capacity to design a sustainable world through cooperative action, but our species’ attitudes and actions will have to change. Our preparedness will have to improve as well. Such change will have to be based on realistic, fact-based appraisals of where we are now and where we want to be in the future. Biologists will have to commit to helping the rest of us understand why the protection of biodiversity is critical to enjoying a sustainable world. Cultural anthropologists also will have to assist humanity at large to understand why the maintenance of cultural diversity also is essential to living sustainably. Educational reform will have to be central to such efforts, to help people learn how to think and act critically and base decisions on the way things really are, and not how we might wish them to be by denying reality. The devotion humans have for structuring beliefs on the basis of little or no evidence, essentially reversing the benefit of rationality, will have to surrender to critical-thinking education established by top-to-bottom educational reform.” Critical-thinking educational reform, however, is much easier to conceive than to bring into reality. A fundamental question is why such reform has not been undertaken. This question is not easy to answer, but perhaps the most fundamental reason is that the educational systems currently in existence are products of the anthropocentric worldview and reflect its mindsets.These educational systems also have developed within the current economic systems responsible for the huge disparities between the rich and poor, and act to reinforce these disparities. Ultimate solutions will emerge only from a clear understanding of the evolution of human psychology, as confronted with the problems we face. If not, then the endemic herpetofauna of Central America, as well as the remainder of life on Earth, will become casualties of the biodiversity crisis that eventually will envelop all humanity.