New Species of Mastigodryas Amaral, 1934 from Brazilian Amazonia and Guyana (Serpentes: Colubridae)

August 13, 2017 | Autor: Giovanna Montingelli | Categoria: Zoology, Herpetology, Ecological Applications
Share Embed

Descrição do Produto

New Species of Mastigodryas Amaral, 1934 from Brazilian Amazonia and Guyana (Serpentes: Colubridae) Author(s): Giovanna G. Montingelli and Hussam Zaher Source: Journal of Herpetology, 45(1):111-119. 2011. Published By: The Society for the Study of Amphibians and Reptiles DOI: 10.1670/09-170.1 URL:

BioOne ( is an electronic aggregator of bioscience research content, and the online home to over 160 journals and books published by not-for-profit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder.

BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research.

Journal of Herpetology, Vol. 45, No. 1, pp. 111–119, 2011 Copyright 2011 Society for the Study of Amphibians and Reptiles

New Species of Mastigodryas Amaral, 1934 from Brazilian Amazonia and Guyana (Serpentes: Colubridae) GIOVANNA G. MONTINGELLI1,2





Museu de Zoologia da Universidade de Sa˜o Paulo, Avenida Nazare´ 481, 04263-000, Ipiranga, Sa˜o Paulo, Sa˜o Paulo, Brazil Programa de Po´s-Graduac¸a˜o em Zoologia, Instituto de Biocieˆncias, Universidade de Sa˜o Paulo, Rua do Mata˜o, Trav. 14, 321, 05598-00, Sa˜o Paulo, Sa˜o Paulo, Brazil

ABSTRACT.—A new species of Mastigodryas is described based on 10 specimens collected on the Upper regions of the Negro and Branco River basins, in the States of Amazonas and Roraima, Brazil, and in the northernmost portion of Guyana. Apparently this species is associated with the open areas of the Amazonian region known as ‘‘lavrados,’’ ‘‘campos,’’ and ‘‘savannas.’’ The new species is characterized by the presence of eight supralabials; five dorsal stripes situated on the anterior portion of the body that fade or disappear on the posterior half of the body and exhibit a reddish unicolor pattern; and a hemipenis with an enlarged spine situated on the left side of the sulcus spermaticus. The new species is similar to Mastigodryas pleei and, herein, included in the pleei group, along with M. pleei, Mastigodryas amarali, and Mastigodryas bruesi.

The genus Mastigodryas includes medium to large, diurnal, and predominantly terrestrial snakes (Martins and Oliveira, 1998; Giraudo, 2001). Species are usually found in humid habitats such as swamps and ponds, associated with forested or open areas (Hardy and McDiarmid, 1969; Peters and OrejasMiranda 1986; Martins and Oliveira, 1998). The genus exhibits a wide geographical distribution, occurring from southern Mexico to northern Argentina, with a few records on continental (e.g., Trinidad and Tobago) and oceanic islands (e.g., Grenada, San Vincent and the Grenadines, and in Barbados, where it was introduced; Stuart, 1941; Peters and Orejas-Miranda, 1986; Henderson and Powell, 2009). Presently, the genus contains 11 species: Mastigodryas bifossatus, Mastigodryas boddaerti, Mastigodryas melanolomus, Mastigodryas amarali, Mastigodryas bruesi, Mastigodryas cliftoni, Mastigodryas danieli, Mastigodryas dorsalis, Mastigodryas heathii, Mastigodryas pleei, and Mastigodryas pulchriceps. The first three are polytypic, whereas the remaining eight are monotypic (Peters and Orejas-Miranda, 1986; Tipton, 2005; see Table 1). All juvenile Mastigodryas retain a banded dorsal pattern, whereas adults can be banded (e.g., M. bifossatus, M. cliftoni, and M. pulchriceps) or striped (the remaining eight species of the genus; Stuart, 1941; Peters and Orejas-Miranda, 1986; Montingelli, 2009). However, bands might be present in a more or less faint condition on the anterior portion of the body in some adults of typically striped species, and stripes tend to fade or disappear in the posterior part of the body of some adult striped forms. The genus Mastigodryas was divided by Stuart (1941) into four species groups on the basis of the external morphology of adults, with the exception of M. pulchriceps that was considered of uncertain affinities. Stuart’s arrangement of the genus is maintained mainly unaltered until today, with the groups bifossatus, melanolomus, boddaerti, and pleei. The bifossatus group has a banded pattern in the adult, eight supralabials, a shorter snout, and 15 dorsal scale rows in the middle of the body, whereas the other three groups have a striped or unicolor pattern in adults, nine supralabials, a longer snout, and 17 dorsal scale rows. The pleei group is distinguished from both melanolomus and boddaerti groups by the length and position of the upper light lateral stripes (sensu Stuart, 1941) on the dorsum. In the former group, the larger upper light lateral stripes are formed by three dorsal scale rows that include the upper portion of row 3, row 4, and the inner portion of row 5, whereas in the latter two groups the upper light lateral stripes are only two dorsal scale rows wide, being formed by rows 3

Corresponding Author. E-mail: [email protected]

number 4 and 5 (except for M. heathii; Fig. 1). According to Stuart (1941), the condition present in M. pulchriceps is intermediate between both patterns because it exhibits a banded pattern, eight supralabials and 17 dorsal rows. In this study, we describe a new species of Mastigodryas that retains an adult striped pattern with the upper light lateral stripe involving three dorsal scale rows and, thus, is considered a member of the pleei group (sensu Stuart, 1941). The new species occurs in open formations of the Brazilian states of Amazonas and Roraima and of the Upper Demerera-Berbice region in Guyana (Fig. 2). MATERIALS AND METHODS We examined specimens of the genus Mastigodryas from the following institutions: American Museum of Natural History (AMNH), Academy of Natural Sciences (ANSP); California Academy of Sciences (CAS); Colec¸a˜o Herpetolo´gica da Universidade de Brası´lia (CHUNB); Field Museum of Natural History (FMNH); Instituto Butantan (IBSP); Instituto Nacional de Pesquisas da Amazoˆnia (INPA); Museum of Natural History, University of Kansas (KU); Museum of Comparative Zoology, Harvard University (MCZ); Museu Paraense Emı´lio Goeldi (MPEG); Museum of Vertebrate Zoology (MVZ); Museu de Zoologia da Universidade de Sa˜o Paulo (MZUSP); University of Michigan, Museum of Zoology (UMMZ) and United States National Museum, Smithsonian Institution (USNM). Species of the genus Mastigodryas employed in comparisons belong to the pleei group and are characterized by a striped pattern on the dorsum exhibiting an upper light lateral stripe formed by three dorsal scale rows. Also, we included M. heathii in our comparison, a striped species traditionally included in the boddaerti group. In Appendices 1 and 2, we provide a list of specimens examined and an index of localities with their geographical coordinates. Head length (HL) was measured to the nearest 0.01 mm with a digital caliper. Total length (TTL) and tail length (TAL) were measured to the nearest 1 mm by stretching the specimens along a ruler. Ventral scales were counted beginning from the first one distinctly wider than long (for details, see Myers [2003] and Zaher et al. [2008]). Sex was determined by presence or absence of hemipenes, confirmed through a small incision made at the base of the tail. Maturity of the specimens was checked according to Slip and Shine (1988). Methods for hemipenial preparation and terminology follow Zaher (1999) and Zaher and Prudente (2003). Sex and measurements of the new species are given in Table 2, and the summary of scale counts of the species employed herein is given in Table 3. Means are given 6 1 SD.



TABLE 1. Composition of the species groups of Mastigodryas according to Peters and Orejas-Miranda (1986) and Tipton (2005). Species group

bifossatus group

boddaerti group

melanolomus group

Peters and Orejas-Miranda, 1986

M. bifossatus bifossatus (Raddi, 1820) M. M. M. M. M. M. M. M. M. M. M.

bifossatus striatus (Amaral, 1931) bifossatus triseriatus (Amaral, 1931) bifossatus villelai (Hoge, 1952) boddaerti boddaerti (Sentzen, 1796) boddaerti dunni (Stuart, 1933) heathii (Cope, 1875) boddaerti ruthveni (Stuart, 1933) dorsalis (Bocourt, 1890) melanolomus alternatus (Bocourt, 1884) melanolomus laevis (Fischer, 1881) melanolomus melanolomus (Cope, 1868)

M. melanolomus tehuanae (Smith, 1943) pleei group

Tipton, 2005

M. M. M. M.

sanguiventris (Taylor, 1954) amarali (Stuart, 1938) bruesi (Barbour, 1914) pleei (Dume´ril et al, 1854)

Incertae sedis M. danieli Amaral, 1934 M. pulchriceps (Cope, 1868)

RESULTS Mastigodryas moratoi sp. nov. Figures 3, 4A,B, and 5 Holotype.—MZUSP 5371, an adult male collected on 4 November 1972 by the Expedic¸a˜o Permanente da Amazoˆnia (E.P.A. 72.2575); approximately 50 m of altitude, on the north bank of the Negro River in Tapera, State of Amazonas, Brazil (00u259S, 64u359W). The right hemipenis was prepared and stored separately in the hemipenial collection of the Museu de Zoologia da USP (MZUSP). Paratypes.—AMNH 141798, female collected on 26 August 1995 by Charles J. Cole and Carol R. Townsend (JC 7055); 3.2 km by road WNW Dubulay Ranch House, APPROXIMATELY 60 m of altitude (05u379N, 57u539W); INPA 16350, female collected by Rafael Bernhard, in Santa Isabel do Rio Negro, Comunidade Sa˜o Joa˜o (00u249S, 65u029W); MZUSP 5355, female collected on 12 November 1972 by the Expedic¸a˜o Permanente da Amazoˆnia (E.P.A. 72.2797), near Tapurucuara, in Paricatuba, State of Amazonas, Brazil (00u319S, 65u019W); MZUSP 5361, male collected on 12 October 1974 by the Expedic¸a˜o Permanente da Amazoˆnia (E.P.A. 72.1952), near Tapurucuara, on the south bank of Negro River in Sa˜o Joa˜o, State of Amazonas, Brazil (00u339S, 64u579W), with the right hemipenis prepared; MZUSP 5370, female collected on 6

FIG. 1. Dorsal striped pattern (sensu Stuart, 1941) showing terminology used in the present work. (A) Mastigodryas pleei; (B) Mastigodryas boddaerti ruthveni; a. upper light lateral stripe; b. inner light lateral stripe; 1. mediodorsal dark stripe; 2. dorsolateral dark stripe; 3. lateral dark stripe. Modified from Stuart (1941: figs. 1, 5, plate III).

M. M. M. M. M. M. M. M. M. M. M. M. M. M. M. M. M. M. M. M. M. M. M. M.

bifossatus bifossatus (Raddi, 1820) bifossatus lacerdai Cunha and Nascimento, 1978 bifossatus striatus (Amaral, 1931) bifossatus triseriatus (Amaral, 1931) bifossatus villelai (Hoge, 1952) boddaerti boddaerti (Sentzen, 1796) boddaerti dunni (Stuart, 1933) heathii (Cope, 1875) boddaerti ruthveni (Stuart, 1933) dorsalis (Bocourt, 1890) melanolomus alternatus (Bocourt, 1884) melanolomus laevis (Fischer, 1881) melanolomus melanolomus (Cope, 1868) melanolomus slevini (Stuart, 1933) melanolomus stuarti (Smith, 1943) melanolomus tehuanae (Smith, 1943) melanolomus veraecrucis (Stuart, 1941) sanguiventris (Taylor, 1954) amarali (Stuart, 1938) bruesi (Barbour, 1914) pleei (Dume´ril et al., 1854) cliftoni (Hardy, 1964) danieli Amaral, 1934 pulchriceps (Cope, 1868)

November 1972 by the Expedic¸a˜o Permanente da Amazoˆnia (E.P.A. 72.2628), on the north bank of Negro River in Tapera, State of Amazonas, Brazil (00u259S, 64u359W); MZUSP 5372, male collected on 6 November 1972 by the Expedic¸a˜o Permanente da Amazoˆnia (E.P.A. 72.2615), on the north bank of Negro River in Tapera, State of Amazonas, Brazil (00u259S, 64u359W); MZUSP 8190, female collected on 4 April 1982 by M. Lemos, on the north bank of the Negro River in Sa˜o Gabriel da Cachoeira, State of Amazonas, Brazil (00u089S, 67u039W); MZUSP 9154, male collected in January, 1986 by Celso M. Carvalho (C.M.C. 1120) in Boa Vista, State of Roraima, Brazil, (02u499N, 60u409W), with the right hemipenis prepared; MZUSP 10695, male collected on 1–2 January 1993, by Celso M. Carvalho (C.M.C. 3633) in Taiano, State of Roraima, Brazil (03u169N, 61u069W), with the right hemipenis prepared.

FIG. 2. Locality map for specimens of Mastigodryas moratoi, Mastigodryas amarali, Mastigodryas bruesi, Mastigodryas heathii, and Mastigodryas pleei. See Appendix 2 for a list of localities identified by their number in the map. The star represents the type locality of M. moratoi. Inset: map of South America indicating the area (gray square) on the larger map.

NEW SPECIES OF MASTIGODRYAS AMARAL TABLE 2. Measurements of the holotype and paratype specimens of Mastigodryas moratoi. HL: head length, TTL: total length, TAL: tail length. Missing data are represented by a hyphen. Specimen with an asterisk corresponds to the holotype. Mastigodryas moratoi


HL mm

TTL mm

TAL mm

AMNH 141798 INPA 16350 MZUSP 5355 MZUSP 5370 MZUSP 8190 MZUSP 5361 MZUSP 5371* MZUSP 5372 MZUSP 9154 MZUSP 10695


27.09 25.64 24.45 25.02 27.15 29.99 29.75 26.63 30.92

1,073 1,008 880 948 1,151 1,128 949 1,168

303 284 237 276 323 318 282 328

Etymology.—The specific name honors Celso M. de Carvalho from the Instituto Nacional de Pesquisas da Amazoˆnia (INPA) who contributed significantly to the knowledge of the herpetofauna of the State of Roraima, Brazil. Diagnosis.—Mastigodryas moratoi differs from all other species in the genus by the combined presence of five distinct dark dorsal stripes on the anterior portion of the body (one mediodorsal, two dorsolaterals, and two laterals); gular region smudged with dark or mostly dark with cream spots until the level of the first ventrals; tail uniformly reddish or yellowish; usually eight supralabials, with the third, fourth, and fifth in contact with the orbit; midregion of the hemipenis below the capitulum with several enlarged spines, one of which is significantly larger and hooklike. Description of the Holotype.—The specimen is an adult male, with a total length of 1,151 mm, head length 30 mm (2.6% total length), and tail length 323 mm (28.06% total length). Pholidosis.—Rostral scale wider than high; internasals and prefrontals paired, as wide as long; supraoculars, frontal and parietals almost twice longer than wide; one loreal approximately twice longer than high; one preocular higher than wide, with the tip visible from above; two postoculars, the upper one larger with the tip visible dorsally; eight supralabials with the third, fourth, and fifth in contact with the orbit; two anterior and two posterior temporals, longer than wide and disposed obliquely, anterior ones in contact with both supraoculars; mental triangular; 10 infralabials with the first five in contact with the first pair of genials; two pairs of genials, the posterior approximately twice longer than the first pair and with two small scales between them. Dorsal scales smooth, in 17–17–15 rows, with two apical pits. Reduction occurs on the rows two and three at the level of the 112th ventral; ventrals 178; cloacal scale divided; subcaudals 99. Pattern of Stripes.—Dorsum with five dark dorsal stripes: one mediodorsal, two dorsolaterals and two laterals (Fig. 1A). Mediodorsal stripe placed on upper portion of row number eight, on row number nine (vertebral) and upper portion of row number eight of the other side. Dorsolateral stripes occupy upper portion of the row five, the row six and inner part of row seven on each side; lateral stripes placed on superior portion of row two and inferior portion of row three, on each side (2/3; 5/ 6/7; 8/9/8; 5/6/7; 2/3). Mediodorsal and dorsolateral stripes with small and dark marks disposed obliquely on the lateral margin of each scale (like an inverse ‘‘v’’) producing the effect of a light center on the stripes (on rows six and nine). Lateral dark stripes with small and dark marks disposed along the superior margin of row two and inner margin of row three, from the anterior region of the body to 36th ventral scale. The mediodorsal stripe fades on the posterior end of body, whereas dorsolateral stripes fade on anterior portion of body. Upper light lateral stripes (Fig. 1A) situated between the dark dorsolateral and lateral stripes are more conspicuous on


anterior part of body and faded on posterior end. On the anterior part of body, these stripes cover the upper half of row number three, row four, and lower portion of row five, and on the posterior end, they cover only row four and lower part of row five, before disappearing completely at the level of tail. Coloration.—Head uniformly brown dorsally; supralabials light cream, ventrally marginated by a thin dark line; ocular band dark brown, extending from eye to posterior temporal; gulars dark gray smudged with small cream spots. Anterior portion of dorsum striped with dark brown and cream stripes that disappear posteriorly, leaving dorsum with a unicolored bluish-gray pattern and slightly reddish tail. The upper light lateral stripes appear about a third of the distance posteriorly on body and extend to posterior region. The first ventrals are smudged with dark gray like the gulars and become immaculate cream posteriorly. Hemipenis.—Hemipenis fully everted and almost maximally expanded; slightly folded on one side as a result of previous eversion and fixation (Fig. 4A, B). The injection of petroleum jelly reached the point of maximal inflation of the organ but failed to correct the fold. Hemipenis unilobed, with a simple centrolinear sulcus spermaticus that ends on the apex of the lobe. Enlarged globular lobe covered with calyces, papillated distally and spinulated proximally. Spinules enlarge while calyces reduce gradually toward the base of the lobe. Lobe not capitulated but with its proximal boundary defined by a slight constriction. Distal half of the hemipenial body covered by rows of medium-sized spines whereas the proximal half lacks any ornamentation. Sulcate surface with one hypertrophied spine present on the lateral side of the distal half of the body left of the sulcus spermaticus. Large basal pocket present throughout the length of the proximal half of the hemipenial body. Intraspecific Variation.—Five adult males, three adult, and two subadult females were available for study. We considered measurements of only two adult females (AMNH 141798, INPA 16350) because the third one had a broken tail and no skull (MZUSP 5355). The largest female has 27.09 mm HL, 1,073 mm TTL, and 303 mm TAL. The largest male has 30.92 mm HL, 1,168 mm TTL, and 328 mm TAL. Tail 28.2% of TTL for females (N 5 2) and 28.5% of TTL for males, being slightly longer in males. All measurements taken can be found in Table 2. Ventral counts range from 175–187 (mean 5 185.0 6 4.42; N 5 10) and are significantly different between females and males (females with 182–187, mean 5 185.0 6 1.58; N 5 5; males with 175–178, mean 5 177.0 6 1.22; N 5 5; t7.5 5 8.94, P 5 0.00002). The cloacal scale is divided and the subcaudals are paired in all specimens examined. Subcaudal counts range from 92–102 (mean 5 98.5 6 3.16; N 5 8) and do not differ significantly between females and males (females 92–102, mean 98.25 6 4.5; N 5 4; males 97–101, mean 5 98.75 6 1.71; N 5 4; t3.8 5 20.21, P . 0.8). All the specimens examined have smooth dorsal scales with two apical pits and an undifferentiated vertebral row, and the same dorsal formula 17/17/15 with the reduction from 17 to 15 scales occurring posteriorly (111–124 ventrals), mostly on scale rows two and three but also on scale rows three and four. Loreal always present; temporal scales vary in 2 + 2 or 2 + 3 (2 + 2 on both sides, N 5 5; 2 + 2 / 2 + 3, N 5 2; 2 + 3 on both sides, N 5 3); seven of the 10 specimens exhibit eight supralabials on both sides with the third, fourth, and fifth in contact with the orbit (except MZUSP 10695, which shows only the fourth and fifth in contact on the left side); one specimen (MZUSP 9154) has nine supralabials on both sides and two individuals exhibit individual variation, one with eight supralabials on the left side and 10 on the right (fourth, fifth, and sixth) (MZUSP 5372) and the other with nine on the left side (fourth, fifth, and sixth) and eight on the right. Nine of the 10 specimens have 10 infralabials, and only one has nine

1 1 1 2 2 2 2

1/1 2/1 2/2 1/2 2/2 2/2 3/2

+ + + + + + +

+ + + + + + + 1 2 2 3 2 3 4

(N (N (N (N (N (N (N

5 5 5 5 5 5 5

5 5 5 5 5 5 5 5 5

1) 2) 3) 1) 29) 19) 4)

4) 1) 18) 5) 1) 11) 10) 13) 10)

17/17/15 (N 5 83)

17/17/15 (N 5 23)

17/17/15 (N 5 52)

9/9(1–4) (N 5 7) 9/9(1–5) (N 5 21) 10/10(1–5) (N 5 5 2) 11(1–5) (N 5 1)

(N (N (N (N (N (N (N (N (N

7(3,4)/8(4,5) (N 5 1) 8/8(3,4,5) (N 5 11) 8(3,4,5)/9(4,5,6) (N 5 10) 9/9(4,5,6) (N 5 63)

1 2 2 2 2 2 3 2 3

M. pleei

+ + + + + + + + +

9/9(1–5) (N 5 9) 10/10(1–5) (N 5 14)

1/1 1/1 2/1 2/2 1/2 2/2 2/2 2/2 3/2

+ + + + + + + + +

8/8(4,5) (N 5 1) 8(3,4,5)/9(3,4,5) (N 5 1) 8(4,5,6)/9(4,5,6) (N 5 2) 9/9(4,5,6) (N 5 15) 9(4,5,6)/10(4,5,6) (N 5 4)

1 1 1 1 2 2 2 2 2

M. heathii

M. bruesi

9/9(1–5) (N 5 6) 9/10(1–5) (N 5 1) 10/10(1–5) (N 5 43)

17/17/15 (N 5 15)

2 + 2/2 + 2 (N 5 8) 2 + 2/2 + 3 (N 5 5) 2 + 3/2 + 3 (N 5 2)

9(1–5) (N 5 6) 10(1–5) (N 5 9)

M. amarali


17/17/15 (N 5 10)


2 + 2/2 + 2 (N 5 5) 2 + 3/2 + 3 (N 5 3) 2 + 2/2 + 3 (N 5 2)


9/9(1–5) (N 5 1) 10/10 (1–4) (N 5 1) 10/10 (1–5) (N 5 8)


8/8(3,4,5) (N 5 6) 8(3,4,5)/8(3,4) (N 5 1) 8(3,4,5)/9(4,5,6) (N 5 1) 9/9(4,5,6) (N 5 1) 8(3,4,5)/10(4,5,6) (N 5 1) 8(4,5)/9(4,5,6) (N 5 1) 9/9(4,5,6) (N 5 12) 9(4,5,6/10(4,5,6) (N 5 1) 9(4,5,6)/10(5,6,7) (N 5 1) 9/9(4,5,6) (N 5 51) 9/10(4,5,6) (N 5 2)


M. moratoi

115–117,2/3r (N 5 3) 107–121,3/4r (N 5 5) 103,4/5r (N 5 1) 112,2/3l (N 5 1) 110–121,3/4l (N 5 6) 106,4/5 l(N 5 1) 120,1/2 r(N 5 1) 109–111,2/3r (N 5 3) 80–125,3/4r (N 5 29) 111–117,4/5r (N 5 3) 109–120,2/3l (N 5 3) 80–125,3/4l (N 5 28) 103–120,4/5l (N 5 5)

115–130,2/3r (N 5 6) 110–140,3/4r (N 5 36) 118–126,4/5r (N 5 4) 115–130,2/3l (N 5 7) 110–140,3/4l (N 5 28) 122–125,4/5l (N 5 4)

97–119, 3/4r,l (N 5 12)

111–123,2/3r (N 5 9) 122,3/4r (N 5 1) 112–124,2/3l (N 5 6) 118–121,3/4l (N 5 4)

D Red

168–201 (N 5 83)

178–200 (N 5 22)

188–209 (N 5 41)

183–196 (N 5 12)

175–187 (N 5 10)


85–110 (N 5 64)

98–115 (N 5 16)

113–133 (N 5 38)

98–119 (N 5 10)

92–102 (N 5 8)


TABLE 3. Summary of scale counts of the species employed on the comparisons with Mastigodryas moratoi. Numbers in parentheses represent supralabials (SL) and infralabials (IL) in contact with the orbit and genials, respectively. Temporal, primary, and secondary (T). Dorsals counted on the anterior, middle and posterior regions of the body (D). Dorsal reduction formula (D Red) shows the number of the ventral scale that is at the level of the reduction and the number of the dorsal row which suffers the reduction on the right (r) and on the left (l) sides of the body. Ventral scales (V). Subcaudal scales (SC).




FIG. 3. Holotype of Mastigodryas moratoi (MZUSP 5371). Scale bars: 2 cm (head details) and 5 cm (body).

(AMNH 141798); genials 2 + 2, 1–5 in contact with the first pair (N 5 9) or 1–4 in contact (N 5 1); one or two pairs of scales between the second pair of genials. Gular color varies in respect to the number of spots, which can be more numerous and brighter in some individuals. Mediodorsal stripe is usually present only on the anterior half of the body, rarely extending through the posterior half as in the holotype. In the majority of the specimens, the mediodorsal stripe disappears together with the dorsolateral stripes (N 5 8). In the holotype, this stripe extends until the posterior region of the body, and in one individual (MZUSP 10695), the mediodorsal and dorsolateral stripes reach the posterior region separately but faded. The dark lateral stripes are very thin and discrete in the majority but in three individuals are more evident (AMNH 141798; MZUSP 5355; 5370). Belly light and immaculate (N 5 7) or slightly smudged (N 5 3). Coloration in Life.—Celso M. de Carvalho provided a photograph of a recently dead specimen, which is used to describe the color pattern of the species in life (Fig. 5). In general, the color is similar to that described for the 10 fixed specimens but with more conspicuous colors. The dorsum is brown with dark brown stripes; mediodorsal stripe changes to a lighter tone posteriorly and disappears on the posterior half of the body. The posterior half of the body is uniformly reddish. Comparisons.—Comparisons are here restricted to the species belonging to the pleei group and with M. heathii. Although the latter species is traditionally included in the boddaerti group, it shares a striped pattern with the members of the pleei group.

Mastigodryas moratoi, M. amarali, M. heathii, and M. pleei share the presence of an upper light lateral stripe formed by three scale rows whereas the remaining species of the genus exhibit the same lateral stripe formed only by two rows (Fig. 1). Mastigodryas bruesi shows a variable condition of the length of the upper light lateral stripe, which can be composed either of three, two, or only one dorsal rows (Montingelli, 2009). Mastigodryas moratoi, as well as M. amarali and M. bruesi, can be distinguished from M. heathii by the lower position of the light lateral stripe on the dorsum involving dorsal rows number three, four, and five instead of four, five, and six in the latter species. Mastigodryas moratoi differs from M. amarali, M. bruesi, and M. pleei by the presence of eight supralabial scales instead of nine, a dorsal scale row reduction that occurs mainly on the second and third rows instead of third and fourth or fourth and fifth rows and a lower number of ventral and subcaudal scales. Additionally, M. amarali and M. bruesi retain only one broad dark dorsal stripe on the anterior region of body instead of five dark dorsal stripes as in M. moratoi (Montingelli, 2009; Table 3). Although M. pleei also retains five dark dorsal stripes (Stuart, 1941; Roze, 1966; Montingelli, 2009, Fig. 1), the three more central ones (the mediodorsal and the dorsolaterals) fuse on the neck to form a broad dark stripe that extends until the posterior end of the body; a pattern similar to the one exhibited by M. amarali and M. bruesi. Hemipenial Comparisons.—Mastigodryas moratoi shares with M. amarali, M. bruesi, and M. heathii the presence of enlarged spines



FIG. 4. Everted and inflated hemipenes of (A) Mastigodryas moratoi MZUSP 5371; (B) M. moratoi MZUSP 9154; (C) Mastigodryas heathii MZUSP 5708; (D) Mastigodryas pleei MZUSP 6187. Sulcate (right) and asulcate (left) surfaces. Scale bars: 0.5 cm.

on the distal region of the body, laterally to the sulcus spermaticus. Mastigodryas pleei lacks such a condition, retaining only smaller spines uniformly distributed around the distal half of the hemipenial body (Figs. 4, 6). Mastigodryas moratoi and M. heathii differ from M. amarali and M. bruesi by a lower number of medium-sized spines and by the presence of three significantly larger hooklike spines. Mastigodryas moratoi differs from M. heathii by the presence of smaller spines on the asulcate surface and smaller papillae ornamenting the calyces of the lobe (Fig. 4). Geographical Distribution and Habitat.—Mastigodryas amarali, M. heathii, M. moratoi, and M. pleei exhibit restricted geographic distributions, inhabiting semiarid and arid regions of South America (Tipton, 2005; Montingelli, 2009). Mastigodryas amarali occurs in northeastern Venezuela and the islands of Trinidad,

Tobago, and Margarita (Hernandez and Radda, 1992; Montingelli, 2009); M. heathii inhabits arid regions along the Cordillera Occidental of Peru (Tipton, 2005; Montingelli, 2009); M. moratoi is associated with areas of open vegetation in the Amazonian basin of Brazil and Guyana (Fig. 2). In Brazil, specimens were collected on the north and south bank of the Negro River, in the state of Amazonas, and on the south bank of the Branco River, in the state of Roraima. According to field notes obtained from Celso M. de Carvalho and from documents belonging to the E.P.A. Expedition to Amazonia (see Vanzolini and Morato, 1991), specimens from Roraima were collected in open areas called ‘‘lavrados,’’ whereas specimens from Amazonas were collected in an area of savanna locally called ‘‘capoeira de terra firme.’’ Also, the


FIG. 5. Recently killed specimen of Mastigodryas moratoi Taiano, Roraima, Brazil.

specimen from Dubulay Ranch House is associated with a region of savanna and was collected approximately 60 m of altitude. Mastigodryas pleei is found in arid regions of northern South America (Colombia and Venezuela, including Islas Margarita and Testigos) and, also, in southern Central America (Panama; Montingelli, 2009). Mastigodryas bruesi is an insular species that occurs in Grenada, Saint Vincent and the Grenadines, and Barbados in the Lesser Antilles (Tipton, 2005; Henderson and Powell, 2009; Montingelli, 2009).


Acknowledgments.—We are grateful to the following curators, collection managers, and institutions for permission to examine the specimens under their care and for allowing the preparation of hemipenial material: D. Frost and D. Kizirian (AMNH); Ned Guilmore (ANSP); A. Leviton, R. Drewes, and J. Vindum (CAS); G. R. Colli (CHUNB); M. Kearney and A. Resetar (FMNH); F. L. Franco and V. Germano (IBSP); R. Vogt and V. T. de Carvalho (INPA); W. E. Duellman and L. Trueb (KU); J. B. Losos and J. Rosado (MCZ); A. L. Prudente (MPEG); J. McGuire and C. Spencer (MVZ); R. Nussbaum and G. Schneider (UMMZ); and G. Zug, R. Heyer, R. McDiarmid, R. Wilson, and T. Harstell (USNM). Also, we are grateful to C. M. de Carvalho who provided field data and photographs on specimens belonging to the new species; D. Mesquita who provided field data for his new record of M. pleei collected in Monte Alegre, Para´, Brazil; A. R. Percequillo and V. Verdade who revised an earlier version of this manuscript; and C. de Castro-Mello who provided technical support and helped with the geographical coordinates. We are grateful to R. Scartozzoni for his help determining the maturity of the specimens. This research was supported by grants from the Coordenac¸a˜o de Aperfeic¸oamento de Pessoal de Nı´vel Superior, American Museum of Natural History, California Academy of Sciences and United States National Museum to GGM, and the Fundac¸a˜o de Amparo a` Pesquisa do Estado de Sa˜o Paulo (BIOTA/FAPESP grant 02/13602-4) and Conselho Nacional de Desenvolvimento Cientı´fico e Tecnolo´gico to HZ. Two anonymous referees offered constructive criticism on the manuscript. LITERATURE CITED

DISCUSSION Based on its external morphology, we suggest that M. moratoi belongs to the pleei group that includes M. amarali, M. bruesi, and M. pleei, all exhibiting a light dorsolateral stripe that involves dorsal scales rows three, four, and five. If our assignment of M. moratoi to the pleei group proves to be correct, the distribution of group from the open and semiarid and arid areas of Panama, Colombia and Venezuela (including Margarita and Testigos islands; Stuart, 1941; Peters and OrejasMiranda, 1986; Montingelli, 2009) is greatly extended to the open areas of Brazil, more precisely to the junction of the Negro and Branco Rivers and the savannas of the Upper DemeraraBerbice region in Guyana. Also, this geographical extension for the group includes one new record of M. pleei (CHUNB 31188), recently published for the northern region of Brazil (savannas of Monte Alegre in the state of Para´; Franc¸a et al., 2006).

FRANC¸A, F. G. R., D. O. MESQUITA, AND G. R. COLLI. 2006. A checklist of snakes from Amazonian savannas in Brazil, housed in the Colec¸a˜o Herpetolo´gica da Universidade de Brası´lia, with new distribution records. Occasional Papers of the Oklahoma Museum of Natural History 17:1–13. GIRAUDO, A. R. 2001. Serpientes de la Selva Paranaense y del Chaco Hu´medo. L.O.L.A., Buenos Aires, Argentina. HARDY, L. M., AND R. W. MCDIARMID. 1969. The amphibians and reptiles of Sinaloa, Mexico. University of Kansas Publications of the Museum of Natural History 18:39–252. HENDERSON, R. W., AND R. POWELL. 2009. Natural History of West Indian Reptiles and Amphibians. University Press of Florida, Gainesville. HERNANDEZ, V. M., AND M. DE RADA. 1992. Contribucion al conocimiento del genero Mastigodryas (Serpentes: Colubridae) en Venezuela. Acta Biologica Venezuelica 13:67–81. MARTINS, M., AND M. E. OLIVEIRA. 1998. Natural history of snakes in forests of the Manaus region, Central Amazonia, Brazil. Herpetological Natural History 6:78–150.

FIG. 6. Everted and inflated hemipenes of (A) Mastigodryas bruesi KU 288661; (B) Mastigodryas amarali USNM 217216. Sulcate (right) and asulcate (left) surfaces. Scales: 0.5 cm.



MONTINGELLI, G. G. 2009. Revisa˜o Taxonoˆmica do Geˆnero Mastigodryas Amaral, 1934 (Serpentes, Colubridae). Unpubl. Ph. D. thesis, Universidade de Sa˜o Paulo, Sa˜o Paulo, Brasil. MYERS, C. W. 2003. Rare snakes—five new species from eastern Panama: reviews of northern Atractus and southern Geophis (Colubridae: Dipsadinae). American Museum Novitates 3391:1–47. PETERS, J. A., AND B. OREJAS-MIRANDA. 1986. Catalogue of the Neotropical Squamata, Part I. Snakes. Rev. Smithsonian Institution Press, Washington, DC. ROZE, J. A. 1966. La Taxonomia y Zoogeografia de los Ophidios en Venezuela. Caracas, Ed. Biblioteca Central Universidad Central Venezuela, Caracas. SLIP, D. J., AND R. SHINE. 1988. The reproductive biology and mating system of Diamond Pythons, Morelia spilota (Serpentes, Boidae). Herpetologica 44:396–404. STUART, L. C. 1941. Studies on Neotropical Colubrinae. VIII. A revision of the genus Dryadophis Stuart 1939. Miscellaneous Publications Museum of Zoology, University of Michigan 49:1–106. TIPTON, B. L. 2005. Snakes of the Americas: Checklist end Lexicon. Krieger Publishing Company, Melbourne, Victoria, Australia. VANZOLINI, P. E., AND C. M. C. MORATO. 1991. Two sibling and sympatric species of Gymnophthalmus in Roraima, Brazil (Sauria, Teiidae). Pape´is Avulsos de Zoologia 37:173–226. ZAHER, H. 1999. Hemipenial morphology of the South American xenodontine snakes, with a proposal for a monophyletic Xenodontinae and a reappraisal of colubroid hemimpenes. Bulletin of the American Museum of Natural History 240:1–168. ZAHER, H., AND A. L. C. PRUDENTE. 2003. Hemipenes of Siphlophis (Serpentes: Xenodontinae) and techniques of hemipenial preparation in snakes: a response to Dowling. Herpetological Review 34:295–302. ZAHER, H., M. E. OLIVEIRA, AND F. L. FRANCO. 2008. A new brightly colored species of Pseudoboa Schneider, 1801 from the Amazon Basin (Serpentes: Xenodontinae). Zootaxa 1674:27–37. Accepted: 29 June 2010.

Rio Zana, near the basecamp on trail btw Monte Seco and Chorro Blanco, about 2,5 km northeast of Monte Seco: FMNH 231772; Piura: FMNH 41593. Mastigodryas moratoi: BRAZIL: Amazonas: Paricatuba, near Tapurucuara: MZUSP 5355. Sa˜o Joa˜o, near Tapurucuara: MZUSP 5361. Tapera, Rio Negro: MZUSP 5370, MZUSP 5371, MZUSP 5372. Santa Isabel do Rio Negro, Comunidade Sa˜o Joa˜o: INPA 16350. Sa˜o Gabriel da Cachoeira, BR 210, Km 07: MZUSP 8190. Roraima: Boa Vista: MZUSP 9154. Taiano: MZUSP 10695. GUYANA: Upper DemeraraBerbice Region: approximately 2 miles (approximately 3 km) by Road from Dubulay Ranch House: AMNH 141798. Mastigodryas pleei: BRAZIL: Para´: Monte Alegre: CHUNB 31188; COLOMBIA: Atlantico: FMNH 165643, Barranquilla: MPEG 18332, MZUSP 2115; Bolı´var: El Manteco: Hato Terrecay, 16 km north of El Manteco: AMNH 114734, 114736, Parupa: La Gran Sabana, 40 km east of Cavanayen: AMNH 115676, 1 km (by road) of Rio Aro on route 19: MVZ 176259, Santa Rosa region: MZUSP 6187; Cundinamarca: Girardot: MPEG 18346; El Cesar: Valencia: UMMZ 54956, 16 km southwest of Valledupar: KU 169961; Huila: 5 km north of Villavieja: MVZ 71529, 3 km north of Cerbatana, near Road to San Alfonzo from Villavieja: MVZ 71527–71528; La Guajira: FMNH 165641, Nazaret: USNM 194740; Magdalena: Pozo Colorado, 11 km west of Santa Marta: CAS 116168–116170, 116181– 116182, 10 south of Santa Marta: CAS 113873; Santander: San Gil: UMMZ 74807; Tolima: Coyaima: MVZ 42048, Road between Coyaima and Chaparra: MVZ 42044, Espinal: IB 8596; Valle de Cauca: KU 117025, Obando: AMNH 91810, Cali: USNM 151727–151730, Rio Raposo, south of Buenaventura: USNM 154047–154048; Antioquia: Lomera: MPEG 18330, Sampedro, village some 12 mi north of Medellin: AMNH 35747, 35750, Sabanalarga: AMNH 35791, San Martin: Remedios: MPEG 18331; ´ : AMNH 89775, El Espino: KU 110640, Herrera: 5 km eastPANAMA northeast of Pese: KU 107688, Parita: USNM 127301, Pearl Islands: AMNH 11259; Nueva Gorgona: AMNH 89976–89978, 88980–88990, KU 110635; VENEZUELA: Zulia: El Laberinto: IB 19436, Mission El Tukuko, Sierra de Perija´: MPEG 18333, La Guajira: CAS 94628; Nueva Esparta: USNM 22533, Isla de Margarita: Cueva El Convento 1 km north, 1 km west of San Francisco de Macanao: KU 117045, La Asuncion: 3 km northnortheast of Salamanca: KU 117041, UNSM 217218, Los Robles: USNM 79225, Porlamar: KU 117044; TESTIGOS: MCZ 6146.

APPENDIX 1 Specimens Examined Mastigodryas amarali: VENEZUELA: Bolivar: forest along Rio Aro, between mouth of Can˜a Azul and rapids near Cerro Coroba, estado. 1,200 ft elevation (approximately 360 m): AMNH 75545; Gua´rico: Cariquito: ANSP 18286–18287; Martinique: Martinique: ANSP 5596; Monagas: Caripito: AMNH 65562; Cocollar: probably near Mount Turumiquire: FMNH 17838; San Antonio de Maturin: MCZ 9988–9990; Nueva Esparta: USNM 22534, Salamanca: USNM 217216; Sucre: Muelle de Cariaco: 6 km (by road) east of Muelle de Cariaco: KU 117038; Penı´nsula de Paria: Yacua: MCZ 43852; Trinidad: Princes Town: AMNH 2943; Tobago: USNM 10137. Mastigodryas bruesi: GRENADA: Carriocoa: MCZ 79101–79102, USNM 104229, Isle de Caille: USNM 79161, Southwest Point: KU268659; Saint Andrew: Balthazar: USNM 79193, Birch Grove: KU 268654, Pearl’s Airport: MVZ 84059; Saint David: Corinth: MCZ 79714; Saint George: CAS 39460–39461, MCZ 185667, near of: MCZ 7792, MCZ 133176–133179, 0,5 mi south (approximately 800 m) of Annandale falls: KU 268656, Annandale Estate: USNM 79191, Beausejour: MCZ 79712– 79713, Beavrejour: MCZ 96717, Botanic Garden: KU 268657–268658, MCZ 88052, Constantine: KU 268655, Perseverance: MCZ 88051, Ross Point: UMMZ 127073, Tempe, northeast of Saint George: MCZ 79715; Saint Patrick: Green Island: KU 268653; GRENADINES: Bequia: Park: KU 268660–61, USNM 104230–104231, Mustique Island: Grand Bay: KU 268662, Quatre Island: USNM 79099, SAINT VINCENT: Saint George: Admiralty Bay: MVZ 84061, Kingston: MCZ 6142, MCZ 157560–157563, 177561, Lowman’s Hill: MCZ 79708–79709, Princess Margaret Bay: MCZ 79711, Cane Garden: MCZ 79710; Saint Patrick: Belle Isle: KU 268663, Tobago Cays: MCZ 79103, Union Island: USNM 79166, Chatham Bay: KU 268664, Yong’s Island: MCZ 38195. Mastigodryas heathii: PERU: Lima: Nan˜a: MCZ 132768, Verrugas Canyon: USNM 51513, Paramonga, Rocky beach at mouth of Rio Fortaleza: AMNH 74740; Ancash: FMNH 81534; La Libertad: FMNH 34296–34298, 34316–34317, 34319, Trujillo, vic. of, AMNH 116323– 116324, on the coast, near Trujillo, 325 km from Punta Aguja, Moche: MZUSP 5708, Pacasmayo: FMNH 5706; Cajamarca: 3 km south (by road) San Pablo, MCZ 183598, Asuncio´n, KU 221724, Rio Zana, FMNH 222664, Rio Zana, 1 m north (airline) Monte Seco, 1,380 m: ANSP 31804,

APPENDIX 2 Gazetteer The numbers preceding collecting localities are the same as presented in Figure 2. BRAZIL Amazonas 1. Paricatuba, near Tapurucuara, 00u319S, 65u019W. 2. Santa Isabel do Rio Negro, comunidade Sa˜o Joa˜o, 00u249S, 65u029W. 3. Sa˜o Gabriel da Cachoeira (BR 210, Km 07), 00u089S, 67u039W. 4. Sa˜o Joa˜o, near Tapurucuara, 00u339S, 64u579W. 5. Tapera, north bank of Negro River, 00u259S, 64u359W. Para´ 6. Monte Alegre, 02u019S, 54u049W. Roraima 7. Boa Vista, 02u499N, 60u409W. 8. Taiano, 03u169N, 61u069W. COLOMBIA Antioquia 9. Remedios, San Martin, 07u019S, 74u329W. 10. Sabanalarga, 06u519S, 48u249W. 11. San Pedro, village some 12 miles north of Medellin, altitude slightly higher than Santa Elena, San Pedro, 06u289S, 75u339W. Atlantico 12. Barranquilla, 11u009N, 74u499W. Bolivar 13. Region of Santa Rosa, 10u279N, 72u139W. 14. Gulf of Morrosquillo, west of Sincelejo, 09u249N, 75u409W. 15. Arenal, Village in front of Soplaviento, Canal del Dique, 10u239N, 75u089W. Cundinamarca 16. Girardot, 04u189S, 74u489W. El Cesar

NEW SPECIES OF MASTIGODRYAS AMARAL 17. 16 km southwest of Valledupar, Valledupar, 10u249S, 73u229W. 18. Valencia, 10u189S, 73u249W. Huila 19. 3 km north of Cerbatana, near road to San Alfonzo from Villavieja, 03u199S, 75u109W. La Guajira 20. Nazaret, 12u119S, 71u179W. 21. Caserio de Chimare, 3 km from sea shore, 12u219S, 71u279W. Magdalena 22. 10 km south of Santa Marta, 11u099S, 74u139W. 23. Pozo Colorado, 11 km west of Santa Marta, 11u129S, 74u149W. Santander 24. San Gil, 06u339S, 73u089W. Tolima 25. Coyaima, 03u489S, 75u129W. 26. Espinal, 04u169N, 74u899W. 27. Road between Coyaima and Chaparra, 03u489S, 75u129W. Valle de Cauca 28. Cali, 03u279N, 76u319W. 29. Obando, 04u359N, 75u599W. 30. Rio Raposo, south of Buenaventura, 03u539N, 77u049W. GUYANA Upper Demerara-Berbice Region 31. Dubulay Ranch House, approximately 2 miles by road from, 05u379N, 57u539W. Upper Takutu-Upper Essquibo Region 32. Kuyuwini Landing, 2u049N, 59u159W.


57. 1 km (airline) northeast of Monte Seco, Rio Zana, 06u529S, 79u059W. 58. Cerro Condoroaz near Quebrada San Isidro, approximately 6 km (airline) west-southwest of Monte Seco, 06u519S, 79u099W. La Libertad 59. Chiclin, Hacienda Chiclin, 07u509S, 79u099W. 60. Moche, 08u119S, 79u029W. 61. Pacasmayo, 07u249S, 79u339W. 62. Trujillo, 08u079S, 79u029W. Lima 63. Verrugas, Canyon, 11u529S, 76u329W. 64. Nana, 11u599S, 76u509W. 65. Lima, Garden in Magdalena Vieja, 12u059S, 77u049W. SAINT VINCENT AND GRENADINE ISLANDS Bequia 66. Park, 13u169N, 61u079W. Grenadines 67. Carriocoa, 12u299N, 61u289W. 68. Quatre Island, 12u579N, 61u159W. 69. Mustique Island, Grand Bay, 12u539N, 61u119W. 70. Chathan Bay, Union Island, 12u369N, 61u269W. 71. Tobago Cays, 12u389N, 61u229W. Saint George 72. Cane Garden, south of Kingston, 13u89N, 61u139W. 73. Kingston, 13u109N, 61u149W. 74. Princess Margaret Bay, 13u09N, 61u149W. Trinidad 75. Princes Town, 10u169N, 61u239W.

GRENADA VENEZUELA Saint Andrew 33. Birch Grove, 12u069N, 61u409W. 34. Balthazar, 12u79N, 61u399W. Saint David 35. Corinth, 12u29N, 61u409W. Saint George 36. near Saint George, 12u079N, 61u409W. 37. Annandale Estate, 12u59N, 61u439W. 38. 0.5 miles south of Annandale Falls, 12u59N, 61u439W. 39. Beausejour, 12u69N, 61u449W. 40. Botanic Garden, St George, 12u39N, 61u449W. 41. Constantine, 12u49N, 61u439W. 42. Perseverance, 12u69N, 61u449W. 43. Ross Point, 12u29N, 61u459W. 44. Tempe, northeast of Saint George, 12u39N, 61u449W. Saint Patrick 45. Green Island, 12u139N, 61u359W. PANAMA Canal Zone 46. Cocle, 08u279N, 80u269W. Cocle 47. 8 km southwest of Penonome´, 08u279N, 80u249W. Herrera 48. 22 km northwest of Chitre, 08u049N, 80u359W. 49. 24 km southwest of Aguadulce, 08u059N, 80u419W. 50. 5 km east-northeast of Pese, 07u559N, 80u349W. 51. Parita, 08u009N, 80u319W. Panama 52. El Espino, 08u509N, 79u519W. 53. Nueva Gorgona, 08u339N, 79u529W. PERU Ancash 54. Yungay, 09u099S, 77u449W. Cajamarca 55. Assuncion, 07u199S, 78u319W. 56. 3 km south (by road) of San Pablo, 07u089S, 78u499W.

Bolivar 76. Forest along Rio Aro, between mouth of Can˜a Azul and rapids near Cerro Coroba estate. 1,200 ft elevation 06u419N, 63u529 W. 77. 1 km east (by road) of Rio Aro on route 19, 07u309N, 64u059W. 78. Hato Terrecay (16 km north of El Manteco), El Manteco, 07u359N, 62u309W. 79. La Gran Sabana, 40 km east of Cavanayen, Parupa, 05u409N, 61u239W. Dependencia Federal 80. Testigos Island, northwest from Trinidad, 11u229N, 63u059W. Guarico 81. Caraquito, 9u219N, 65u229W. Monagas 82. Caripito, Jungle approximately 100 m, 10u089N, 63u069W. 83. Cocollar, near Mount Turumiquire, 10u079N, 63u529W. 84. San Antonio de Maturin, 10u079N, 63u439W. Nueva Esparta 85. Cueva El Convento, 1 km north, 1 km west of San Francisco de Macanao, Isla Margarita, 11u019N, 64u189W. 86. La Asuncion, 11u029N, 63u539W. 87. La Asuncion, 3 km north-northeast of Salamanca, Isla Margarita, 07u309N, 64u059W. 88. Los Robles, Isla Margarita, 10u589N, 63u509W. 89. Porlamar, Isla Margarita, 10u579N, 63u519W. Sucre 90. 16 km east of Quetepearita, Cumana´, 10u269N, 64u029 W. 91. 2.5 km southwest of Cumana´, Cumana´, 10u269N, 64u119W. 92. 21 km east of Cumana´, 10u279N, 63u579W. 93. 25 km (by road) south of Cumana´, Cumana´, 10u139N, 64u119W. 94. 5 km (by road) west of Muelle de Cariaco, Muelle de Cariaco, 10u299N, 63u339W. 95. San Fernando, Montes, 10u189N, 63u569W. 96. Yacua, Penı´nsula de Paria, 10u409N, 62u309W. Trujillo 97. 46 km west-northwest of Valera, Valle Verde, 09u269N, 70u599W. 98. Sabana de Mendoza, 09u269N, 70u469W. Zulia 99. El Laberinto, 10u339N, 72u139W. 100. Mission El Tukuko, Sierra de Perija, 09u599N, 72u489W.

Lihat lebih banyak...


Copyright © 2017 DADOSPDF Inc.