Revalidation of Herpetodryas reticulata (Peters, 1863) (Serpentes: Colubridae) from Ecuador

Revalidation of Herpetodryas reticulata (PeteRs, 1863) (seRPentes: ColubRidae) fRom eCuadoR

Giovanna G. MontinGelli1,5, JorGe H. valencia2,3, Marco a. Benavides4 and HussaM ZaHer1

1. Museu de Zoologia da universidade de são paulo. avenida Nazaré, 481, 04263‑000, ipiranga, são paulo, sp, Brasil. e‑mails: [email protected]; [email protected]

2. Fundación Herpetológica Gustavo orcés. avenida amazonas, 3008 and rumipamba, Quito, ecuador. e‑mail: [email protected]. Museo de Zoología, pontificia universidad católica del ecuador. avenida 12 de octubre y roca, aptdo. 17‑01‑2184, Quito, ecuador.

e‑mail: [email protected]. Museo ecuatoriano de ciencias Naturales, avenida rumipamba, 341 and avenida de los shyris, Quito, ecuador.

e‑mail: [email protected]. corresponding author.

aBstract. the name Herpetodryas reticulata Peters, 1863, is revalidated and assigned to the snakes of the genus Mastigodryas from the dry forests of central and southwestern ecuador. Mastigodryas reticulatus (Peters, 1863) is characterized by the presence of a striped dorsal pattern, with the upper light lateral stripe formed by two scale rows (4 and 5); dorsal scales with dark apical edges; a light, immaculate throat and venter; a higher number of ventral scales; an elongate and nude region on the base of the hemipenis; and thin, enlarged spines located lateral to the sulcus spermaticus on the distal region of the body of the hemipenis. this species is a member of the M. boddaerti Group, along with M. boddaerti and M. heathii.

Keywords. Mastigodryas, Colubridae, taxonomy, ecuador.

introduction

Herpetodryas reticulata was described by Peters (1863), based on a specimen collected by von C. Re-isf in the Gulf of Guayaquil, ecuador; the holotype presently is housed at the Zoologische museum in berlin. according to Peters (1863), the species differs from Herpetodryas boddaerti and H. rappi by the presence of smaller and rhomboidal cephalic scales and by the absence of dark blotches on the head and throat. However, boulenger (1894) considered H. re‑ticulata to be a junior synonym of drymobius bod‑daerti. stuart (1941) accepted boulenger’s decision, but explicitly mentioned that the ecuadorian speci-mens are distinct from the other populations of dry‑mobius boddaerti.

stuart (1941) and subsequent authors (Peters and orejas-miranda, 1986; tipton, 2005) recognized three species of Mastigodryas in ecuador – the geo-graphically widespread and polytypic species Mas‑tigodryas boddaerti; M. heathii from ecuador and Peru; and M. pulchriceps from western ecuador. more recently, torres-Carvajal (2004) reported the presence of a fourth species, M. melanolomus, in the drier and lower areas of isla de la Plata. this is a polytypic, widespread Central american species in which the nominal subspecies is characterized by a predominantly uniform and unicolor pattern (stuart, 1941; Peters and orejas-miranda, 1986). the identi-fication of M. melanolomus in ecuador and the range extension that it represents from Central america to

ecuador may have been influenced by Perez-santos and moreno’s (1988) report of Colombian specimens with a unicolor dorsum from Cundinamarca and tolima.

in contrast to the aforementioned authors, we at-tribute full species status to the Mastigodryas popula-tion distributed in the dry forests of central and south-western ecuador. the available and applicable name for this population is Mastigodryas reticulatus (Pe-ters, 1863), for which we provide a detailed descrip-tion of the holotype and morphological variation.

Materials and MetHods

We compared Mastigodryas reticulatus with the andean species of the genus that belong to the M. boddaerti Group – i.e., M. heathii (Cope, 1875) and M. boddaerti (sentzen, 1796) (montingelli and Zaher, 2011: table 1). Herein, M. boddaerti refers only to the typically striped specimens from bolivian, brazilian, Colombian, and Peruvian amazonia (cis-andean samples). the unicolor trans-andean popu-lations from ecuador and Colombia, traditionally as-signed to M. boddaerti by stuart (1941) and Peters and orejas-miranda (1986), are considered here as a distinct, undescribed species of the genus (montin-gelli, 2009); herein, it is recognized as a separate tax-on that is designated Mastigodryas sp.

Head length (Hl) was measured to the nearest 0.01 mm with digital calipers. total length (ttl)

south american Journal of Herpetology, 6(3), 2011, 189-197© 2011 brazilian society of Herpetology

and tail length (tal) were measured to the nearest 1 mm by carefully aligning the specimens along a ruler. measures of ttl, tal, and subcaudal counts were not recorded for specimens with broken tails. ventral scales were counted from the first scale that is distinctly wider than long (described by myers [2003] and Zaher et al. [2008]) to the last scale an-teriorly adjacent to the cloacal scale. sex was deter-mined by noting the presence or absence of hemi-penes by making a small incision at the base of the tail. maturity of the specimens was checked follow-ing the methods of slip and shine (1988). methods for hemipenial preparation and terminology fol-low those of Zaher (1999) and Zaher and Prudente (2003). scale counts, sex, and measurements are given in tables 1 and 2.

We examined 276 specimens, as follow: 24 Mas‑tigodryas reticulatus (Peters, 1863); 186 M. bod‑daerti; 33 to M. heathii; and 33 Mastigodryas sp. specimens of Mastigodryas were provided by the following institutions (institutional codes in paren-theses): american museum of natural History, new York (amnH); California academy of sciences, san

francisco (Cas); Coleção Herpetológica da univer-sidade de brasília, brasília (CHunb); museo de an-fibios y Reptiles, fundación Herpetológica Gustavo orcés, Quito (fHGo); field museum of natural His-tory, Chicago (fmnH); university of Kansas, law-rence (Ku); museo ecuatoriano de Ciencias natura-les, Quito (meCn); museu Paraense emílio Goeldi, belém do Pará (mPeG); museu de Zoologia da uni-versidade de são Paulo, são Paulo (mZusP); mu-seum of Comparative Zoology, Harvard university, Cambridge (mCZ); museo de Zoología, Pontifícia universidad Católica de ecuador (QCaZ); museo de la escuela Politécnica nacional, Quito (ePn); universidade federal do Ceará, fortaleza (ufC); universidade federal do mato Grasso, Cuiabá (ufmt); universidade federal da Paraíba, João Pes-soa (ufPb); university of michigan, museum of Zo-ology, ann arbor (ummZ); united states national museum, smithsonian institution, Washington, d.C. (usnm); Zoologische museum, berlin (Zmb). specimens examined are listed in appendix i.

results

Mastigodryas reticulatus (Peters, 1863) figures 1-3, 4a

Herpetodryas reticulata Peters, 1863: type locality: Guayaquil.

dryadophis boddaerti boddaerti: stuart, 1941: 66, part.

Mastigodryas boddaerti boddaerti: Pérez-santos and moreno, 1991: 245-246, part.

Mastigodryas melanolomus: torres-Carvajal, 2004: 85.

Holotype – Zoologische museum, berlin (Zmb 4504), an adult specimen collected by von C. Reisf near Guayaquil, ecuador (02°10’s, 79°50’W).

diagnosis – Mastigodryas reticulatus differs from all the other members of the genus by the following com-bination of characters: (1) presence of a striped dor-sal pattern, with the upper, light lateral stripe formed by scale Rows 4 and 5; (2) dorsal scales with dark apical edges; (3) light, immaculate throat and venter; (4) higher number of ventral scales; (5) elongate bas-al and nude region of the hemipenis; (6) presence of some enlarged and thinner spines lateral to the sulcus spermaticus, on the distal region of the body of the hemipenis.

taBle 1. measurements of the specimens of Mastigodryas reticu‑latus in mm. Hl: head length, ttl: total length, tal: tail length. missing data are represented by a dash. the holotype is desig-nated with an asterisk.

specimen sex Hl ttl talamnH 126612 f 37.60 1365 368amnH 28984 f 32.89 1092 296Cas 8802 f 27.06 977 274fHGo 6123 f 28.24 1447 292fHGo 8390 f — — 225fHGo 8391 f 20.75 — —fHGo 6756 f 26.20 1342 280Ku 84670 f 28.08 — —mZusP 19387 f 33.20 1169 321mZusP 19388 f 37.49 1337 340QCaZ 7441 f 29.09 1104 211QCaZ 946 f 23.61 1163 243QCaZ 945 f 19.27 954 194QCaZ 3968 f 24.83 1360 267QCaZ 4054 f 27.79 1328 367usnm 211049 f 29.30 915 259usnm 211050 f 19.23 646 191amnH 28987 m — — —Ku 301093 m 26.00 840 236QCaZ 7440 m 23.33 1193 229usnm 211050 m 39.11 1257 354amnH 22099 — 38.06 — —QCaZ 3945 — 18.62 — —Zmb 4504* — — — —

Revalidation of Herpetodryas reticulata190

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Mas

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retic

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us9

/ 9 (4

, 5, 6

) (N

= 2

1)

9 (3

, 4, 5

) / 1

0 (4

, 5, 6

) (N

= 1

)

9 / 1

0 (4

, 5, 6

) (N

= 1

)

10 /

10 (4

, 5, 6

) (N

= 1

)

10 /

10 (1

–5) (

N =

13)

9 / 9

(1–5

) (N

= 4

)

9 / 9

(1–4

) (N

= 1

)

10 (1

–5) /

9 (1

–4) (

N =

3)

10 (1

–5) /

9 (1

–5) (

N =

1)

11 /

11 (1

-5) (

N =

1)

2 +

2 / 2

+ 2

(N =

16)

2 +

2 / 2

+ 3

(N =

6)

2 +

3 / 2

+ 3

(N =

1)

1 +

2 / 1

+ 2

(N =

1)

17 /

17 /

15

(N =

24)

181-

206

(N =

21)

102-

116

(N =

20)

Mas

tigod

ryas

bod

daer

ti9

/ 9 (4

, 5, 6

) (N

= 1

57)

10 /

10 (4

, 5, 6

) (N

= 7

)

10 (5

, 6, 7

) / 9

(4, 5

, 6) (

N =

13)

11 (5

, 6, 7

) / 9

(4, 5

, 6) (

N =

4)

10 (4

, 5, 6

) / 1

1 (5

, 6, 7

) (N

= 4

)

8 / 8

(4, 5

, 6) (

N =

1)

10 (1

-5)

(N =

155

)

9 (1

-5) (

N =

13)

10 (1

-5) /

11

(1-5

) (N

= 8

)

11 /

11 (1

-5) (

N =

5)

10 (1

-5) /

9 (1

-5) (

N =

4)

11 (1

-5) /

12

(1-5

) (N

= 1

)

2 +

2 / 2

+ 2

(N =

112

)

2 +

2 / 2

+ 3

(N =

35)

2 +

3 / 2

+ 3

(N =

23)

1 +

2 / 1

+ 2

(N =

7)

2 +

2 / 1

+ 2

(N =

3)

2 +

3 / 1

+ 2

(N =

2)

2 +

2 / 2

+ 4

(N =

1)

3 +

2 / 3

+ 2

(N =

1)

17 /

17 /

15

(N =

186

)

177-

200

(N =

185

)

90-1

18

(N =

158

)

Mas

tigod

ryas

sp.

9 / 9

(4, 5

, 6) (

N =

25)

9 (4

, 5, 6

) / 8

(3, 4

, 5) (

N =

5)

8 (3

, 4, 5

) (N

= 2

)

10 /

10 (4

, 5, 6

) (N

= 1

)

9 / 9

(1-5

) (N

= 1

6)

10 /

10 (1

-5) (

N =

11)

8 (1

-4) /

8 (1

-5) (

N =

3)

9 / 9

(1-4

) (N

= 1

)

2 +

2 / 2

+ 2

(N =

23)

2 +

2 / 2

+ 3

(N =

7)

2 +

3 / 2

+ 3

(N =

2)

1 +

2 / 2

+ 1

(N =

1)

17 /

17 /

15

(N =

33)

173-

195

(N =

33)

92-1

12

(N =

24)

Mas

tigod

ryas

hea

thii

9 / 9

(4, 5

, 6) (

N =

23)

8 (3

, 4, 5

) / 9

(4, 5

, 6) (

N =

6)

9 (4

, 5, 6

) / 1

0 (4

, 5, 6

) (N

= 2

)

9 (4

, 5, 6

) / 1

0 (5

, 6, 7

) (N

= 1

)

8 / 8

(4, 5

) (N

= 1

)

10 /

10 (1

-5) (

N =

21)

9 / 9

(1-5

) (N

= 9

)

9 (1

-4) /

10

(1-5

) (N

= 3

)

2 +

2 / 2

+ 2

(N =

16)

2 +

2 / 2

+ 3

(N =

8)

2 +

3 / 2

+ 3

(N =

4)

1 +

1 / 1

+ 1

(N =

1)

1 +

3 / 2

+ 1

(N =

1)

1 +

3 / 1

+ 3

(N =

1)

2 +

2 / 1

+ 1

(N =

1)

2 +

2 / 1

+ 2

(N =

1)

17 /

17 /

15

(N =

33)

179-

202

(N =

32)

98-1

15

(N =

25)

montingelli, G. G. et al. 191

Redescription of the Holotype

pholidosis – Rostral scale wider than high; interna-sals and prefrontals paired, as wide as long; supra-oculars, frontal, and parietals about twice as long as wide; one loreal approximately twice as long as high; one preocular higher than wide, with the tip vis-ible from above; two postoculars, upper scale larger with the tip visible dorsally; nine supralabials, with the fourth, fifth and sixth in contact with the orbit; two anterior and two posterior temporals, longer than wide and obliquely oriented, anterior scales in contact with supraoculars and postoculars; mental triangular; 10 infralabials with first five in contact with first pair of genials; two pairs of genials, posterior genial about twice as long as the first pair. dorsal scales smooth, in 17-17-15 rows, with two apical pits; ventrals 198; cloacal scale divided; subcaudals 106.

dorsal pattern and coloration – Head uniform brown dorsally; first supralabials light cream, and the posterior four smudged with the same color as the dorsum, bluish gray; ocular band dark brown, extending from the eye to the posterior temporals (slightly faded); infralabials and gulars light cream, immaculate as the entire venter; lateral portion of the ventral scales bluish gray as the dorsum; dorsum without vestiges of light lateral stripes, scales with dark apical edges.

intraspecific variation

Measurements (mm) – (N = 20; 12 females, 4 males, 4 juveniles). largest female with following measure-ments: Hl 28.24, ttl 1.45, tal 292. largest male: Hl 39.11, ttl 1.26, tal 354. tail 19.8% ttl for females and 28.2% for males (table 1).

pholidosis – total ventral count range 181-206 (mean 194.8 ± 6.26; N = 21), not significantly different be-tween females and males (females with 181-206, mean 195.7 ± 5.98; N = 16; males with 185-198, mean 192.6 ± 6.80; N = 3). Cloacal scale divided and sub-caudals paired in all specimens examined. total sub-caudal range 102-116 (mean 106.0 ± 3.46; N = 20), not significantly different between males and females (females 102-116, mean 105.8 ± 3.61; N = 15; males 103-112, mean 106.7 ± 3.77; N = 4). all specimens with smooth dorsal scales with two apical pits and undifferentiated vertebral row, and same formula 17/17/15 with reduction from 17 to 15 scales occur-ring posteriorly on scale Rows 3 and 4. loreals al-ways present, approximately twice longer than high; temporal scales vary in 1 + 2, 2 + 2 and 2 + 3 (2 + 2, N = 16; 2 + 2 / 2 + 3, N = 6; 2 + 3, N = 1; 1 + 2, N = 1); almost all specimens (including holotype) with nine supralabials with the supralabials 4-6 in contact with the orbit (except for one female with 10 supralabi-als on both sides, and two females with individual

FiGure 1. Holotype of Herpetodryas reticulata Peters, 1863 (Zmb 4504), from Guayaquil, ecuador.


variation, with nine and ten scales). most specimens with 10 infralabials (females, N = 9; males, N = 1; three unsexed specimens), one female with 11 scales, three females and two males with nine scales, and four specimens exhibiting individual variation with nine and ten scales; genials 2 + 2, 1-5 in contact with the first pair (N = 20) or 1-4 in contact (N = 3); one or two pairs of scales between second pair of genials.

dorsal pattern and coloration – the dorsum is brown or bluish gray in all specimens. two speci-mens (fHGo 8390, QCaZ 946) exhibit vestiges of a juvenile banded pattern on the anterior portion of the body. some specimens (amnH 28984, Ku 84670, 301093; mZusP 3091) have a faint upper light lateral stripe situated on dorsal scale Rows 4 and 5 on the an-terior half of the body; the stripe gradually disappears posteriorly. in most specimens, the ventral surfaces of the head and body are light cream and immaculate. only three individuals have a slightly smudged ven-ter (amnH 28984, fHGo 6756, QCaZ 946), which usually is darker than in the remaining specimens.

coloration in life – Photographs of a freshly pre-served specimen from Guayas (figures 2 and 3) pro-vide information on the coloration in life. the dorsum is light brown with the anterior and posterior region of the body slightly olive or yellow (mZusP 19387, fHGo 3223) and with dark apical edges on the dor-sal scales. the dorsal Row 1 is light cream, as is the venter, which is light and immaculate. the supralabi-als are yellowish and slightly smudged dorsally. the yellowish color pattern predominates on the ventral surface of the head and first ventrals, fading posteri-orly toward an immaculate cream color that extends to the tip of the tail ventrally.

Hemipenis (figure 4a) – We examined fully everted and almost maximally expanded hemipenes of three individuals (Ku 301093, usnm 211050, QCaZ 7441). the unilobed hemipenis has a simple centro-linear sulcus spermaticus, which terminates at the apex of the lobe. the lobe is covered with calyces, and is spinulated proximally and papillated distally. spinules become larger and thinner toward the base of the lobe. the distal end of the hemipenial body is covered with medium-sized spines that are larger on the left side of the sulcus. the remaining portion of the body lacks ornamentation. basal pockets pres-ent on the proximal end of the body. the papillated and naked portions of the organ are longer and have approximately the same size, whereas the spinulated part of the body represents one third of the organ.

Comparisons

samples here assigned to Mastigodryas reticula‑tus differ from cis-andean M. boddaerti by the dark interstitial skin and dark apical edges of dorsal scales (vs. light cream interstitial skin and without dark apical edges; by a higher number of ventral scales (181-206, X = 194.8, N = 21 vs. 177-200, X = 189, 4, N = 185); by the elongate basal and nude region of the hemipenis; and by the presence of some enlarged, thinner spines lateral to the sulcus spermaticus on the distal region of the body (vs. a short, basal nude re-gion lacking enlarged spines distally; figure 4). the species differs from cis-andean Mastigodryas sp. by a light, immaculate throat and venter (vs. dark, spot-ted throat, and dark-edged ventrals); by more ventrals (181-206, X = 194.8, N = 21 vs. 173-195, X = 182.3, N = 33); and by the presence of enlarged, thin spines

FiGure 2. freshly preserved specimen of Mastigodryas reticula‑tus from Guayas, ecuador (mZusP 19387).

FiGure 3. freshly preserved specimen of Mastigodryas reticula‑tus from Guayas, ecuador (mZusP 19387).


lateral to the sulcus spermaticus on the distal re-gion of the body (vs. 1 or 2 well-developed, hook-like spines; figure 4). Mastigodryas reticulatus dif-fers from M. heathii by the width and the position of the upper light lateral stripe that occupies two dorsal scale Rows 4 and 5 (vs. 3 dorsal scale Rows, 3-5) and by the absence of hooklike spines lateral to the sul-cus spermaticus (vs. the presence of hook-like spines) (figure 4).

Geographical distribution (figure 5)

Mastigodryas reticulatus, M. heathii, and Mas‑tigodryas sp. have restricted ranges. Mastigodryas

reticulatus occurs in lowland (up to 600 m a.s.l.), de-ciduous and semideciduous forests (sierra 1999) from central and southwestern ecuador, south to Caraquez bay (manabí Province) and the provinces of santa elena, Guayas, and loja, and in the continental is-lands of Puna and la Plata. its distribution coincides with one of the most important and threatened areas of western ecuador – the tumbesian Region, which harbors a high number of endemic species that in-cludes plants, birds, mammals, and reptiles (aguirre-mendoza and Kvist 2005, Cisneros-Heredia 2006, valencia et al., 2010). Mastigodryas heathii inhabits arid regions of southwestern ecuador, occurring in the provinces of loja and el oro, and arid regions of northwestern Peru, along the Cordillera occidental,

FiGure 4. fully everted and expanded hemipenes of (a) Mastigodryas reticulatus, isla de la Plata, ecuador (Ku 301093); (b) Mastigodry‑as sp., esmeraldas, ecuador (usnm 211054); (C) M. boddaerti, oriximiná, Pa, brasil (mZusP 5077), and (d) M. heathii, la libertad, Peru (mZusP 5708). sulcate (left) and asulcate (right) surfaces. scale bars = 5 mm.


from lima to Piura provinces, from sea level to near-ly 2000 m. Mastigodryas sp. occurs along the humid forests of northwestern ecuador and southwestern Colombia, on the western slopes of the Cordillera occidental, from sea level to nearly 2000 m. Mas‑tigodryas boddaerti has one of the most widespread distributions in the genus, occurring predominantly on low, humid areas of western south america, from bolivia, Peru, and Colombian and brazilian amazo-nia, to the eastern portion of venezuela.

discussion

We consider the unicolored specimens identified as Mastigodryas melanolomus by torres-Carvajal (2004; QCaZ 945, 946) to represent M. reticulatus, rather than M. melanolomus. the latter probably does not occur in ecuador. the Colombian specimens of “M. melanolomus” examined by Perez-santos and moreno (1988) also are thought to belong to a dis-tinct, undescribed species assignable to the M. bod‑daerti Group. therefore, we restrict the distribution of the M. melanolomus Group to Central america, with its southernmost limit in Panamá.

in summary, we hypothesize that three species of genus Mastigodryas inhabit ecuador. We recognize the presence of M. heathii and M. pulchriceps, but resurrect M. reticulatus as a valid species restricted geographically to southwestern ecuador. moreover, a probable fourth, predominantly unicolored species occurs in western ecuador. Populations of this new species have been incorrectly identified as belong-ing to M. boddaerti. Given the recognition of M. re‑ticulatus and Mastigodryas sp. as distinct species, the distributional range of the formerly polytypic M. boddaerti (montingelli and Zaher, 2011) should exclude the trans-andean portion of ecuador and Colombia.

resuMo

na presente contribuição, o nome Herpetodryas reticulata é revalidado e atribuído a população do gê-nero Mastigodryas distribuída nas florestas secas da região central e sudoeste do equador. Mastigodryas reticulatus (Peters, 1863) se caracteriza por apresen-tar padrão dorsal estriado, com a listra lateral superior formada por duas fileiras de escamas, números quarto

FiGure 5. Geographical distribution of Mastigodryas boddaerti, M. heathii, M. reticulatus, and Mastigodryas sp.


e cinco; escamas dorsais possuem ápices escuros; ventre da cabeça e do corpo claros e imaculados; nú-mero elevado de escamas ventrais; região basal e nua do hemipênis longa, e presença de espinhos aumenta-dos e finos, presentes na lateral do sulco espermático, na região distal do órgão. esta espécie é incluída no Grupo M. boddaerti, juntamente com M. boddaerti e M. heathii.

acKnowledGMents

We thank the following curators and colleagues for provid-ing access to the specimens under their care and for allowing the preparation of hemipenial material: d. frost and d. Kizirian (amnH); a. leviton, R. drewes, and J. vindum (Cas); G. R. Colli (CHunb); ana almendariz (ePn); Katty Garzón (fHGo); m. Kearney and a. Resetar (fmnH); W. e. duellman and l. trueb (Ku); J. losos, J. Rosado, and J. Woodward (mCZ); mario Yánez-muñoz y manuel morals-mite (meCn); a. l. Prudente (mPeG); omar torres-Carvajal (QCaZ); d. m. borges-nojosa (ufC); m. a. de Carvalho (ufmt); G. Calazans (ufPb); R. nussbaum and G. schneider (ummZ); G. Zug, R. Heyer, R. mc-diarmid, R. Wilson, and t. Harstell (usnm); W. Günther and C. Kucharzewsky (Zmb). We are also grateful to a. Percequillo and l. trueb for revising an earlier version of this manuscript, C. de Castro-mello (mZusP) for providing technical support, and J. Prado for map construction. this research was supported by grants from the Coordenação de aperfeiçoamento de Pessoal de nível superior (CaPes), Conselho nacional de Pesquisa e de-senvolvimento Científico (CnPq), american museum of natural History (amnH), California academy of sciences (Cas) and united states national museum (usnm) to GGm, fundação de amparo à Pesquisa do estado de são Paulo (biota/faPesP) and Conselho nacional de desenvolvimento Científico e tec-nológico (CnPq) to HZ, and fundación Herpetológica Gustavo orcés and secretaria nacional de educación superior, Ciencia y tecnología del ecuador (senesCYt) Project no. PiC-08-00470 granted to Pontifícia universidad Católica del ecuador.

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submitted: 11 november 2011 accepted: 13 december 2011


appendix i

specimens examined

Mastigodryas boddaerti: bolivia: mZusP 6447; Cochabamba: villa tunari: Ku 183485; el beni: ivon: amnH 22490, 22500; Rurrenabageu: amnH 22476; vacadiez: tumi Chucua: usnm 280403-08; Guayaramerin: usnm 123969; la Paz: mapiri: amnH 32995; santa Cruz: fmnH 35623, 35625, 35659, 195906, ummZ 69377-78; buenavista: ummZ 60793, 63921, 63923, 64002, 67925a, 67925b. bRasil: amapá: CHunb 3639; macapá: CHunb 13157; amazonas: Humai-tá: CHunb 33929; manaus: mZusP 7983; Reserva ducke: mZusP 8490; Reserva inPa-WWf: mZusP 8476, 9309; Ceará: serra de ibiapaba: 7 Km nW de ipú: ufC 3996-98; ubajara: sítio santana: ufC 1348; mato Grosso: mZusP 3798; barra do bugres: estação ecológica serra das araras: ufmt 1637; barra do tapirapés: mZusP 4750; Cáceres: mZusP 1390; Cuiabá: ufmt 1636; vale de são domingos: margem direita do Rio Guaporé: ufmt 1638; mato Grosso do sul: Corumbá: acurizal: serra do amolar: ufmt 1397, 1404, 1408, 1416; serra do urucum: entre urucum e são domingo: mZusP 4434; Pará: alter do Chão: mZusP 8207; ananindeua: biotério evandro Chagas: ufC 2035; augusto Corrêa: fa-zenda Cacoal: mPeG 6475, 6506, 9992; baixo trombetas, margem esquerda em frente a foz do nhamundá: mZusP 4784; barcarena: arredores da cidade: mPeG 16370; barreirinha: Rio tapajós: mZusP 5133, 5142; boca do Cuminá-mirim: mZusP 4819; bragança: bom Jesus: mPeG 2499, 2503, 5091; Capitão Poço: são Pedro: mPeG 12151; Carajás: mPeG 16705, 16920; Colônia nova, próximo ao Rio Gurupi, bR 316: mPeG 4335, 7752, 7778; Curuçá: marauá: mPeG 5889; dom eliseu: mPeG 13394, 13417, 14960, 14961; estação utiariti: ufC 1267; estrada de rodagem, entre Castanhal e são francisco do Pará: mPeG 18177; Gancho do arari: bR 222, entre miranda e arari: mPeG 14861; igarapé-açú: mPeG 936; inhangapi: arraial do Carmo: mPeG 1461, 2729; marabá: CHunb 30421; novo Progresso: serra do Cachimbo: CHunb 33923, 35063; oriximiná: mZusP 4801, 5075-77, 5082, 5488, 5489, 10639–41; ourém: limão Grande: mPeG 1257, 6146, 6163; Parauapebas: serra dos Carajás: CHunb 6668; ca 66 Km sW of Parque nacional da amazonia, Rio tapajós: usnm 288932; Rio Gurupi, nova vida, 25 km de distância do rio, bR-316: mPeG 13679, 14989, 14990, 14992, 14993, 16206; salenópolis: Restinga da praia do maçarico: mZusP 15593; santarém: ummZ 56312; santo antonio do tauá: sítio do Paiva: ufC 2038; são Geraldo do araguaia: CHunb 30420; sítio bela vista, bR-222: mPeG 15204; taboleiro leonardo: Rio trombetas: mZusP 7420; tomé açú: mZusP 3725; uruá: Rio tapajós, Parque nacional da amazônia: mZusP 7298, 7406; vigia: santa Rosa: mPeG 5605, 6827; viseu: bela vista: mPeG 15962, fazenda Real: mPeG 4496, 5292; Rondô-nia: Porto velho: mZusP 3203, 3204, 3205, 3206, 3207; simbal: Guajará-mirim: CHunb 22055, 22056; Roraima: apiaú: mZusP 10915-16, 10321; boa vista: mZusP 9856, 10376, 10479; Cachoeira do Cujubim: Rio Catrimani: mZusP 6392, 8031, margem esquerda do Rio branco: mZusP 9784; Colônia Confiança: mZusP 9968; fazenda salvamento: mZusP 9990; missão Catrimani: mZusP 10481, 10896; tocantins: Palmas: uHe luis eduardo magalhães: mZusP 14435-40, 15623-25, 15729-30. PeRu: fmnH 40024, 134472; Junin: fmnH 40023; loreto: ucayali: Contamana: amnH 52241, 52697, 71615; iquitos: Rio itaya: amnH 52073; monte Carmelo: Requena (uresti): amnH 53165, 55625, 55649; orellana: amnH 52911, 55669; Pampa Hermosa: Rio Cushabatay: amnH 53548, 53552, 55458-59, 55463, 55469, 55475-76, 55719, 55723-24, 55733, 55752, 55779, 55802, 55855, Rio ucayali: amnH 52029, 52058, 53289, 54425, fmnH 45595, 56112, 56134-35, 56160-62, ummZ 71625; Panya: Rio Ponasa: amnH 52344, 52627.

Mastigodryas heathii: PeRu: fmnH 38108, 39643, 229361, usnm 38557; ancash: fmnH 81534; Cajamarca: asunción: Ku 221724; Río Zana: fmnH 231772, 232664; 3 km s by road san Pablo: mCZ 183598; la libertad: Chiclin (the same as Hacienda Chiclin, a sugar-hacienda between trujillo and Chicama): fmnH 34296–98, 34316–17, 34319, amnH 116323; moche: on the coast, near trujillo, 325 km from Punta aguja: mZusP 5708; Pacasmayo: fmnH 5706; trujillo: amnH 116324; lima: nana: mCZ 132768; Paramonga: rocky beach at mouth of Rio fortaleza: amnH 74740; verrugas Canyon: usnm 51513; Piura: fmnH 41593. eCuadoR: el oro: Piñas, villa elvita: fHGo 1083; loja: macará: QCaZ 7993, Km 7 via loja-Cuenca: fHGo 639, san Pedro: fHGo 673-74, vilcabamba: QCaZ 8985, fHGo 2904, fHGo 5552; Zapotillo, Cabeza de toro: fHGo 2673, Quebrada el Cañaberales: fHGo 3485.

Mastigodryas reticulatus: eCuadoR: Guayas: Cerro masvale: QCaZ 7440; via daule-samborondon: QCaZ 7441; el milagro, naldivieso: usnm 211049; Guayaquil: amnH 126612, mZusP 19387 -88, fHGo 6123, QCaZ 3945, Zmb 4504; Playas: usnm 211050; Punas id: Cas 8802; santa elena: amnH 22099, 28987; manabí: manta: fHGo 3223, 6756, 8390, 8391; isla de la Plata: amnH 28984, Ku 84670, Ku 301093, QCaZ 945 -46, 3968; 11 km of Puertoviejo: usnm 211051; Km 5 via Colón-Quimis: QCaZ 4054.

Mastigodryas sp.: Colombia: bolivar: road to Pepinal: mZusP 6132; darien: lago Calima: Ku 169953; Caldas: fmnH 54966; Cauca: fmnH 54967–70, 61665, Gorgona island: amnH 63564; nariño: fmnH 165338; valle de Cauca: san antonio: usnm 151660–62, 267274. eCuadoR: fmnH 16940; Chaquarapata: amnH 23031; bolivar: balzapamba: ummZ 88958; el oro: amnH 18321; esmeraldas: muisne: san francisco: meCn 2898, Río blanco, at equator: usnm 211054, bulun: amnH 13590, san Javier: amnH 13589; imbabura: Paramba: amnH 13433, Pimampiro: meCn 3607, ummZ 83705, san nicolas: ummZ 83707; los Ríos: Playas de montalvo: ummZ 83950; Pichincha: Centro Científico Río Palenque: amnH 119839, maquipucuna: fHGo 3488; Pacto: usnm 211052, domingo de los Colorados: Río baba, 10 km s, 4 km e santo: Ku 142812, 47 km s of Centro Científico Río Palenque: usnm 285490; san miguel de los Colorados: usnm 211053.


Revalidation of Herpetodryas reticulata (Peters, 1863) (Serpentes: Colubridae) from Ecuador

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