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Townsend, Josiah H., Medina-Flores, Melissa, Reyes-Calderón, Onán, Austin, James D. (2013): A new Nototriton (Caudata: Plethodontidae) from Parque Nacional Montaña de Botaderos in northeastern Honduras. Zootaxa 3666 (3): 358-368, DOI: http://dx.doi.org/10.11646/zootaxa.3666.3.6

Nototriton mime sp. nov. Figures 3 A–D Holotype. USNM 579870 (Fig. 3 A), an adult male from Cerro Ulloa (15.3833 °N, 86.0399 °W), Parque Nacional Montaña de Botaderos, 1,705 m, Departamento de Olancho, Honduras, collected 16 April 2011 by M. Medina- Flores, O. A. Reyes-Calderón, and J. H. Townsend; original field number JHT 3400; GenBank accession numbers KC 905090 (16 S) and KC 905094 (cyt b). Paratopotypes. Three (same collectors as holotype); a female (Fig. 3 B; MVZ 269306) and a juvenile (Fig. 3 D; USNM 579872), collected 16 April 2011, 1,710 m, 15.3843 °N, 86.0396 °W; and a male (Fig. 3 C; USNM 579871), 17 April 2011, 1,720 m elevation, 15.3874 °N, 86.0467 °W. Diagnosis. A member of the genus Nototriton diagnosed by possessing 13 costal grooves (> 16 costal grooves in Oedipina), hands and feet longer than broad (hands and feet broader than long in Bolitoglossa), and small nares (0.007–0.010 NL/SL; 0.017–0.029 NL/SL in Cryptotriton and Dendrotriton). Distinctiveness of the new species and its assignment to the Nototriton barbouri species group is supported by analysis of sequence fragments from the mitochondrial genes 16 S and cyt b (Fig. 2). Nototriton mime is unique among described congeners in the Chortís Highlands by having a yellow-brown dorsal coloration in adult males, as well as relatively longer front and hind limbs (FLL/SL 0.195–0.246, HLL/SL 0.224–0.254; versus 0.142–0.174 and 0.153 –0.200 in N. barbouri, 0.148–0.151 and 0.166–0.180 in N. brodiei, 0.137–0.160 and 0.158–0.181 in N. lignicola, 0.156–0.183 and 0.164– 0.211 in N. limnospectator, 0.179–0.191 and 0.204–0.218 in N. picucha, 0.172 and 0.178 in N. stuarti, and 0.160 and 0.197 in N. tomamorum). The new species may be further differentiated from the other species of Nototriton found in the Chortís Highlands as follows: from N. brodiei by having a broader head (HW/SL 0.121–0.138, versus 0.120), larger nares (NL/SL 0.007–0.010, versus 0.004–0.005), a shorter tail (TL/SL 0.698 –1.117, versus 1.420– 1.440), and fewer maxillary teeth (27–46, versus 60–62); from N. lignicola by having a broader head (HW/SL 0.121–0.138, versus 0.103–0.118); from N. limnospectator by having a broader head (HW/SL 0.121–0.138, versus 0.095–0.118) and larger nares (NL/SL 0.007–0.010, versus 0.003); from N. picucha by having a narrower head (HW/SL 0.121–0.138, versus 0.140–0.148); from N. saslaya by having narrower hind feet (HFW/SL 0.044 – 0.050, versus 0.075–0.091), larger nares (NL/SL 0.007–0.010, versus 0.002–0.003), and a higher number of maxillary (27–46, versus 17–22) and vomerine (12–24, versus 3–11) teeth; from N. stuarti by having a slightly narrower head HW/SL 0.121–0.138, versus 0.138), a shorter tail (TL/SL 0.698 –1.117, versus 1.264), and smaller nares (NL/SL 0.007–0.010, versus 0.012); and from N. tomamorum by having well-developed digits with subdigital pads (versus hands and feet syndactylous and lacking subdigital pads), a narrower head (HW/SL 0.121– 0.138, versus 0.145), and smaller nares (NL/SL 0.007–0.010, versus 0.018). Description of holotype. An adult male (SL = 31.8 mm, total length = 67.3 mm) with a slender body and reduced limbs. The head is rounded, slightly broader than the body; nostrils are relatively small (NL/SL= 0.010), and the snout is acutely rounded and of moderate length. Nasolabial grooves are visible but slight, and nasolabial protuberances are not apparent. The eyes are relatively large and protuberant, and the elongate parotoid glands are relatively flat and not well-defined. There are 46 maxillary teeth, 3 slightly enlarged premaxillary teeth in line with the maxillaries, and 24 vomerine teeth. The limbs are short (CLL/SL= 0.40), with the adpressed limb interval approximately 4.5 costal grooves. The hands and feet are narrow with well-developed digits that bear poorly developed subdigital pads. The relative length of the digits is I Measurements of holotype (mm). SL 31.8; AG 16.9; TW 3.3; HL 5.8; HW 3.6; TL 35.5; HLL 6.7; FLL 5.8; CLL 12.5; FFW 0.8; HFW 0.9; NL 0.3; EL 1.8; EW 0.9; IOD 0.7; anterior rim of orbit to snout 1.4; distance separating internal nares 1.1; distance separating external nares 1.6; snout projection beyond mandible 0.3; tip of snout to axilla 9.0; snout to anterior edge of vent 27.8; tail depth at basal constriction 2.7; tail width at basal constriction 2.6. Color in life of adult male holotype. Description of the coloration of the holotype based on a series of color photographs taken in life: dorsal surfaces of paratoid region of head, body, and tail Chamois (84); lateral surfaces mottled Ground Cinnamon (270), with Dark Grayish Brown (284) spots beginning on the centrolateral portion of costal groove 4 and becoming larger posteriorly, with one large blotch present centrolaterally from costal groove 12 to the hind limb; narrow middorsal stripe Pale Pinkish Buff (3), with faint herringbone pattern evident dorsally; dorsal and lateral surfaces of head mottled Vandyke Brown (282) with some Smoky White (261) and Pearl Gray (262) stippling above the mouth; dorsal surface of limbs Sepia (286) with Cinnamon-Rufous (31), Yellow-Ocher (14), and Chamois (84) stippling, becoming more profuse towards body; dorsal surface of tail with Pale Mauve (204) spots beginning about halfway down the length of the tail, with tail becoming almost completely Pale Mauve (204) towards the tip; ventral surface of body with a Dark Mauve (208) to Cyanine Blue (189) cast, with scattered Smoky White (261) and Light Lavender (201) stippling; ventral ground color of tail Fuscous (283), darker than body, with a series of three pairs of Salmon Color (58) blotches behind the vent with spacing equivalent to that of the costal grooves, distal portion of ventral surface of tail with Smoky White (261) and Pale Mauve (204) stippling; color of iris Gem Ruby (65). Variation. In all individuals except the holotype, the tail is shorter than the body (TL/SVL= 0.780 in USNM 579871, a male, 0.817 in MVZ 269306, a female, and 0.698 in USNM 579872, a juvenile) In each case, the tail does not appear to be incomplete, and thus the significance of the degree of variation seen in tail length is not clear. The male paratype (USNM 579871) has 27 maxillary, 5 premaxillary, and 12 vomerine teeth; the female paratype (MVZ 269306) has 37 maxillary, 7 premaxillary, and 17 vomerine teeth. The female (MVZ 269306) and juvenile (USNM 579872) paratypes both exhibit a different color pattern than that seen in the male holotype (USNM 579870) and paratype (USNM 579871). The adult female paratype MVZ 269306 has the following coloration in life: dorsal surface of paratoid region of head and body Hazel (26) with darker Brick Red (36) and paler Robin Rufous (29) blotches forming a herringbone pattern, with a narrow Robin Rufous (29) middorsal stripe; irregular Dark Buff Yellow (54) dorsolateral stripes originate approximately halfway down the body and become better defined and continue onto the tail, bordered laterally by a series of small Vandyke Brown (281) blotches positioned on costal grooves 4, 5, 8, 9, and 10, and becoming a continuous blotch over costal grooves 11, 12, and 13 that extends onto the proximal half of the tail; lateral surfaces of body mottled Dark Buff Yellow (54) and Robin Rufous (29) with Pale Bluish Gray (287) stippling; dorsal surface of head with Raw Umber (280) mottling on the parietal region and between the eyes, lateral surfaces of head Russet (44) with Pale Bluish Gray (287) stippling, ventral surface Glaucous (291) with scattered small Pale Bluish Gray (287) spots; ventral surface of body Vandyke Brown (282) with scattered Smoky White (261) and Pale Bluish Gray (287) stippling most prominent near the head and along the centerline of the venter; dorsal surface of tail mottled Hazel (26) and Robin Rufous (29), with Raw Umber (280) to Vandyke Brown (282) lateral and ventral surfaces with Smoky White (261) and Pale Bluish Gray (287) stippling; iris Mahogany Red (34). Etymology. The specific epithet “ mime ” is a noun in apposition to the generic name, given to honor our late friend Arquimides Gabriel Rosales Martinez, or “ Mime ” (pronounced me-may), a young Honduran biologist passionate about amphibians. Mime and his sister, Novy Hortensia Rosales Martinez, were killed by a drunken driver on 17 December 2010 in Tegucigalpa. Geographic distribution. This species is known only from the immediate vicinity of the type locality, 1,705– 1,720 m on Cerro Ulloa, the highest ridge in the Sierra de Botaderos, forming the border between the departments of Colón and Olancho (Figs. 1, 3). Natural history. The habitat at the type locality is tropical montane cloud forest, in the Lower Montane Wet Forest formation. The male holotype (USNM 579870; Fig. 3 A) was active at night among the leaves of an orchid growing on a small stump on the ground along a wet ridge at 1,705 m elevation. The male paratype (USNM 579871; Fig. 3 C) was collected from underneath a small log embedded in the ground at 1,720 m elevation. The female paratype (MVZ 269306; Fig. 3 B) was found in a bromeliad on the ground at 1,705 m elevation, near the holotype. The juvenile paratype (USNM 579872; Fig. 3 D) was collected during the late afternoon from within a small bromeliad approximately 2 m above the ground on a large fallen branch at 1,710 m elevation. Conservation status. Based on the extremely limited extent of intact cloud forest around the only known locality (approximately 300 hectares), and the continuing deforestation surrounding this locality, we recommend Nototriton mime be immediately considered Critically Endangered (CR B 1 ab[i, ii, iii]+ 2 ab[i, ii, iii]), following the IUCN Red List Criteria (IUCN 2012). The broadleaf forests of the Sierra de Botaderos are approximately 33,000 hectares in total extent, and ranges from 100 to 1,724 m elevation. However, less than 1,000 total hectares of forest above 1,500 m elevation exists, and is fragmented into approximately a dozen discrete patches along the spine of the cordillera. The type locality, Cerro Ulloa, is one of these patches, and is the highest elevation forest in the mountain range. Cerro Ulloa is already deforested up to and above 1,500 m elevation on the southern slopes (Fig. 4 A), and preexisting trails made by farmers cross the summit of the peak (Fig. 4 B) and access inholdings deep with the forest on the Colón side of the ridge. The approximately 300 hectares of intact cloud forest habitat surrounding the type locality are already under direct threat from migratory agriculture and cattle ranching, and, most recently, are indirectly threatened by the dramatic increase in international narcotrafficking in adjacent portions of Colón and Olancho. The recent declaration of the remaining premontane and lower montane rainforests as a national park is a positive development; however, the remote location of the park and high-risk nature of conflicts related to enforcing protection of its remaining forests make conservation here, and throughout eastern Honduras, a resourceintensive challenge. Remarks. With the description of Nototriton mime, there are 17 named species of Nototriton distributed from Guatemala to Costa Rica, six of which are endemic to Honduras (Townsend et al. 2011; Boza-Oviedo et al. 2012). Each of these six endemic species have distributions restricted to cloud forest habitat in the upper reaches of isolated mountain ranges, with only one species (N. limnospectator) known to occur at premontane elevations as low as 800 m. In addition to these six species, there are two as yet undescribed species currently referred to the taxon N. barbouri, one from Parque Nacional Pico Bonito (N. sp A in Townsend et al. 2011) and one from Refugio de Vida Silvestre Texíguat (N. sp. B, op. cit.). The coloration seen in Nototriton mime appears to be the first documented case of sexual dichromatism in the genus Nototriton, and one of few documented cases in tropical salamanders. Another endemic Honduras salamander, Bolitoglossa diaphora, from the Sierra de Omoa in northwestern Honduras, also exhibits marked dichromatism, with males being bluish-gray and females being a mottled red-orange coloration (McCranie & Wilson 1995).

Nototriton mime sp. nov. Figures 3 A���D Holotype. USNM 579870 (Fig. 3 A), an adult male from Cerro Ulloa (15.3833 ��N, 86.0399 ��W), Parque Nacional Monta��a de Botaderos, 1,705 m, Departamento de Olancho, Honduras, collected 16 April 2011 by M. Medina- Flores, O. A. Reyes-Calder��n, and J. H. Townsend; original field number JHT 3400; GenBank accession numbers KC 905090 (16 S) and KC 905094 (cyt b). Paratopotypes. Three (same collectors as holotype); a female (Fig. 3 B; MVZ 269306) and a juvenile (Fig. 3 D; USNM 579872), collected 16 April 2011, 1,710 m, 15.3843 ��N, 86.0396 ��W; and a male (Fig. 3 C; USNM 579871), 17 April 2011, 1,720 m elevation, 15.3874 ��N, 86.0467 ��W. Diagnosis. A member of the genus Nototriton diagnosed by possessing 13 costal grooves (> 16 costal grooves in Oedipina), hands and feet longer than broad (hands and feet broader than long in Bolitoglossa), and small nares (0.007���0.010 NL/SL; 0.017���0.029 NL/SL in Cryptotriton and Dendrotriton). Distinctiveness of the new species and its assignment to the Nototriton barbouri species group is supported by analysis of sequence fragments from the mitochondrial genes 16 S and cyt b (Fig. 2). Nototriton mime is unique among described congeners in the Chort��s Highlands by having a yellow-brown dorsal coloration in adult males, as well as relatively longer front and hind limbs (FLL/SL 0.195���0.246, HLL/SL 0.224���0.254; versus 0.142���0.174 and 0.153 ���0.200 in N. barbouri, 0.148���0.151 and 0.166���0.180 in N. brodiei, 0.137���0.160 and 0.158���0.181 in N. lignicola, 0.156���0.183 and 0.164��� 0.211 in N. limnospectator, 0.179���0.191 and 0.204���0.218 in N. picucha, 0.172 and 0.178 in N. stuarti, and 0.160 and 0.197 in N. tomamorum). The new species may be further differentiated from the other species of Nototriton found in the Chort��s Highlands as follows: from N. brodiei by having a broader head (HW/SL 0.121���0.138, versus 0.120), larger nares (NL/SL 0.007���0.010, versus 0.004���0.005), a shorter tail (TL/SL 0.698 ���1.117, versus 1.420��� 1.440), and fewer maxillary teeth (27���46, versus 60���62); from N. lignicola by having a broader head (HW/SL 0.121���0.138, versus 0.103���0.118); from N. limnospectator by having a broader head (HW/SL 0.121���0.138, versus 0.095���0.118) and larger nares (NL/SL 0.007���0.010, versus 0.003); from N. picucha by having a narrower head (HW/SL 0.121���0.138, versus 0.140���0.148); from N. saslaya by having narrower hind feet (HFW/SL 0.044 ��� 0.050, versus 0.075���0.091), larger nares (NL/SL 0.007���0.010, versus 0.002���0.003), and a higher number of maxillary (27���46, versus 17���22) and vomerine (12���24, versus 3���11) teeth; from N. stuarti by having a slightly narrower head HW/SL 0.121���0.138, versus 0.138), a shorter tail (TL/SL 0.698 ���1.117, versus 1.264), and smaller nares (NL/SL 0.007���0.010, versus 0.012); and from N. tomamorum by having well-developed digits with subdigital pads (versus hands and feet syndactylous and lacking subdigital pads), a narrower head (HW/SL 0.121��� 0.138, versus 0.145), and smaller nares (NL/SL 0.007���0.010, versus 0.018). Description of holotype. An adult male (SL = 31.8 mm, total length = 67.3 mm) with a slender body and reduced limbs. The head is rounded, slightly broader than the body; nostrils are relatively small (NL/SL= 0.010), and the snout is acutely rounded and of moderate length. Nasolabial grooves are visible but slight, and nasolabial protuberances are not apparent. The eyes are relatively large and protuberant, and the elongate parotoid glands are relatively flat and not well-defined. There are 46 maxillary teeth, 3 slightly enlarged premaxillary teeth in line with the maxillaries, and 24 vomerine teeth. The limbs are short (CLL/SL= 0.40), with the adpressed limb interval approximately 4.5 costal grooves. The hands and feet are narrow with well-developed digits that bear poorly developed subdigital pads. The relative length of the digits is IMeasurements of holotype (mm). SL 31.8; AG 16.9; TW 3.3; HL 5.8; HW 3.6; TL 35.5; HLL 6.7; FLL 5.8; CLL 12.5; FFW 0.8; HFW 0.9; NL 0.3; EL 1.8; EW 0.9; IOD 0.7; anterior rim of orbit to snout 1.4; distance separating internal nares 1.1; distance separating external nares 1.6; snout projection beyond mandible 0.3; tip of snout to axilla 9.0; snout to anterior edge of vent 27.8; tail depth at basal constriction 2.7; tail width at basal constriction 2.6. Color in life of adult male holotype. Description of the coloration of the holotype based on a series of color photographs taken in life: dorsal surfaces of paratoid region of head, body, and tail Chamois (84); lateral surfaces mottled Ground Cinnamon (270), with Dark Grayish Brown (284) spots beginning on the centrolateral portion of costal groove 4 and becoming larger posteriorly, with one large blotch present centrolaterally from costal groove 12 to the hind limb; narrow middorsal stripe Pale Pinkish Buff (3), with faint herringbone pattern evident dorsally; dorsal and lateral surfaces of head mottled Vandyke Brown (282) with some Smoky White (261) and Pearl Gray (262) stippling above the mouth; dorsal surface of limbs Sepia (286) with Cinnamon-Rufous (31), Yellow-Ocher (14), and Chamois (84) stippling, becoming more profuse towards body; dorsal surface of tail with Pale Mauve (204) spots beginning about halfway down the length of the tail, with tail becoming almost completely Pale Mauve (204) towards the tip; ventral surface of body with a Dark Mauve (208) to Cyanine Blue (189) cast, with scattered Smoky White (261) and Light Lavender (201) stippling; ventral ground color of tail Fuscous (283), darker than body, with a series of three pairs of Salmon Color (58) blotches behind the vent with spacing equivalent to that of the costal grooves, distal portion of ventral surface of tail with Smoky White (261) and Pale Mauve (204) stippling; color of iris Gem Ruby (65). Variation. In all individuals except the holotype, the tail is shorter than the body (TL/SVL= 0.780 in USNM 579871, a male, 0.817 in MVZ 269306, a female, and 0.698 in USNM 579872, a juvenile) In each case, the tail does not appear to be incomplete, and thus the significance of the degree of variation seen in tail length is not clear. The male paratype (USNM 579871) has 27 maxillary, 5 premaxillary, and 12 vomerine teeth; the female paratype (MVZ 269306) has 37 maxillary, 7 premaxillary, and 17 vomerine teeth. The female (MVZ 269306) and juvenile (USNM 579872) paratypes both exhibit a different color pattern than that seen in the male holotype (USNM 579870) and paratype (USNM 579871). The adult female paratype MVZ 269306 has the following coloration in life: dorsal surface of paratoid region of head and body Hazel (26) with darker Brick Red (36) and paler Robin Rufous (29) blotches forming a herringbone pattern, with a narrow Robin Rufous (29) middorsal stripe; irregular Dark Buff Yellow (54) dorsolateral stripes originate approximately halfway down the body and become better defined and continue onto the tail, bordered laterally by a series of small Vandyke Brown (281) blotches positioned on costal grooves 4, 5, 8, 9, and 10, and becoming a continuous blotch over costal grooves 11, 12, and 13 that extends onto the proximal half of the tail; lateral surfaces of body mottled Dark Buff Yellow (54) and Robin Rufous (29) with Pale Bluish Gray (287) stippling; dorsal surface of head with Raw Umber (280) mottling on the parietal region and between the eyes, lateral surfaces of head Russet (44) with Pale Bluish Gray (287) stippling, ventral surface Glaucous (291) with scattered small Pale Bluish Gray (287) spots; ventral surface of body Vandyke Brown (282) with scattered Smoky White (261) and Pale Bluish Gray (287) stippling most prominent near the head and along the centerline of the venter; dorsal surface of tail mottled Hazel (26) and Robin Rufous (29), with Raw Umber (280) to Vandyke Brown (282) lateral and ventral surfaces with Smoky White (261) and Pale Bluish Gray (287) stippling; iris Mahogany Red (34). Etymology. The specific epithet ��� mime ��� is a noun in apposition to the generic name, given to honor our late friend Arquimides Gabriel Rosales Martinez, or ��� Mime ��� (pronounced me-may), a young Honduran biologist passionate about amphibians. Mime and his sister, Novy Hortensia Rosales Martinez, were killed by a drunken driver on 17 December 2010 in Tegucigalpa. Geographic distribution. This species is known only from the immediate vicinity of the type locality, 1,705��� 1,720 m on Cerro Ulloa, the highest ridge in the Sierra de Botaderos, forming the border between the departments of Col��n and Olancho (Figs. 1, 3). Natural history. The habitat at the type locality is tropical montane cloud forest, in the Lower Montane Wet Forest formation. The male holotype (USNM 579870; Fig. 3 A) was active at night among the leaves of an orchid growing on a small stump on the ground along a wet ridge at 1,705 m elevation. The male paratype (USNM 579871; Fig. 3 C) was collected from underneath a small log embedded in the ground at 1,720 m elevation. The female paratype (MVZ 269306; Fig. 3 B) was found in a bromeliad on the ground at 1,705 m elevation, near the holotype. The juvenile paratype (USNM 579872; Fig. 3 D) was collected during the late afternoon from within a small bromeliad approximately 2 m above the ground on a large fallen branch at 1,710 m elevation. Conservation status. Based on the extremely limited extent of intact cloud forest around the only known locality (approximately 300 hectares), and the continuing deforestation surrounding this locality, we recommend Nototriton mime be immediately considered Critically Endangered (CR B 1 ab[i, ii, iii]+ 2 ab[i, ii, iii]), following the IUCN Red List Criteria (IUCN 2012). The broadleaf forests of the Sierra de Botaderos are approximately 33,000 hectares in total extent, and ranges from 100 to 1,724 m elevation. However, less than 1,000 total hectares of forest above 1,500 m elevation exists, and is fragmented into approximately a dozen discrete patches along the spine of the cordillera. The type locality, Cerro Ulloa, is one of these patches, and is the highest elevation forest in the mountain range. Cerro Ulloa is already deforested up to and above 1,500 m elevation on the southern slopes (Fig. 4 A), and preexisting trails made by farmers cross the summit of the peak (Fig. 4 B) and access inholdings deep with the forest on the Col��n side of the ridge. The approximately 300 hectares of intact cloud forest habitat surrounding the type locality are already under direct threat from migratory agriculture and cattle ranching, and, most recently, are indirectly threatened by the dramatic increase in international narcotrafficking in adjacent portions of Col��n and Olancho. The recent declaration of the remaining premontane and lower montane rainforests as a national park is a positive development; however, the remote location of the park and high-risk nature of conflicts related to enforcing protection of its remaining forests make conservation here, and throughout eastern Honduras, a resourceintensive challenge. Remarks. With the description of Nototriton mime, there are 17 named species of Nototriton distributed from Guatemala to Costa Rica, six of which are endemic to Honduras (Townsend et al. 2011; Boza-Oviedo et al. 2012). Each of these six endemic species have distributions restricted to cloud forest habitat in the upper reaches of isolated mountain ranges, with only one species (N. limnospectator) known to occur at premontane elevations as low as 800 m. In addition to these six species, there are two as yet undescribed species currently referred to the taxon N. barbouri, one from Parque Nacional Pico Bonito (N. sp A in Townsend et al. 2011) and one from Refugio de Vida Silvestre Tex��guat (N. sp. B, op. cit.). The coloration seen in Nototriton mime appears to be the first documented case of sexual dichromatism in the genus Nototriton, and one of few documented cases in tropical salamanders. Another endemic Honduras salamander, Bolitoglossa diaphora, from the Sierra de Omoa in northwestern Honduras, also exhibits marked dichromatism, with males being bluish-gray and females being a mottled red-orange coloration (McCranie & Wilson 1995)., Published as part of Townsend, Josiah H., Medina-Flores, Melissa, Reyes-Calder��n, On��n & Austin, James D., 2013, A new Nototriton (Caudata: Plethodontidae) from Parque Nacional Monta��a de Botaderos in northeastern Honduras, pp. 358-368 in Zootaxa 3666 (3) on pages 361-366, DOI: 10.11646/zootaxa.3666.3.6, http://zenodo.org/record/215885