Revalidation of Herpetodryas Reticulata (Peters, 1863) (Serpentes: Colubridae) from Ecuador Author(S): Giovanna G

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Revalidation of Herpetodryas reticulata (Peters, 1863) (Serpentes: Colubridae) from Ecuador Author(s): Giovanna G. Montingelli, Jorge H. Valencia, Marco A. Benavides and Hussam Zaher Source: South American Journal of Herpetology, 6(3):189-197. Published By: Brazilian Society of Herpetology https://doi.org/10.2994/057.006.0304 URL: http://www.bioone.org/doi/full/10.2994/057.006.0304

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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] 3. Museo de Zoología, Pontificia Universidad Católica del Ecuador. Avenida 12 de Octubre y Roca, Aptdo. 17-01-2184, Quito, Ecuador. E-mail: [email protected] 4. Museo Ecuatoriano de Ciencias Naturales, Avenida Rumipamba, 341 and Avenida de Los Shyris, Quito, Ecuador. E-mail: [email protected] 5. 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 Ecuador may have been influenced by Perez-Santos and Moreno’s (1988) report of Colombian specimens Herpetodryas reticulata was described by Peters with a unicolor dorsum from Cundinamarca and (1863), based on a specimen collected by Von C. Re- Tolima. isf in the Gulf of Guayaquil, Ecuador; the holotype In contrast to the aforementioned authors, we at- presently is housed at the Zoologische Museum in tribute full species status to the Mastigodryas popula- Berlin. According to Peters (1863), the species differs tion distributed in the dry forests of central and south- from Herpetodryas boddaerti and H. rappi by the western Ecuador. The available and applicable name presence of smaller and rhomboidal cephalic scales for this population is Mastigodryas reticulatus (Pe- and by the absence of dark blotches on the head and ters, 1863), for which we provide a detailed descrip- throat. However, Boulenger (1894) considered H. re- tion of the holotype and morphological variation. ticulata to be a junior synonym of Drymobius boddaerti. Stuart (1941) accepted Boulenger’s decision, but explicitly mentioned that the Ecuadorian speci- MATERIALS AND METHODS mens are distinct from the other populations of Dry- mobius boddaerti. We compared Mastigodryas reticulatus with Stuart (1941) and subsequent authors (Peters and the Andean species of the genus that belong to the Orejas-Miranda, 1986; Tipton, 2005) recognized M. boddaerti Group – i.e., M. heathii (Cope, 1875) three species of Mastigodryas in Ecuador – the geo- and M. boddaerti (Sentzen, 1796) (Montingelli and graphically widespread and polytypic species Mas- Zaher, 2011: Table 1). Herein, M. boddaerti refers tigodryas boddaerti; M. heathii from Ecuador and only to the typically striped specimens from Bolivian, Peru; and M. pulchriceps from western Ecuador. Brazilian, Colombian, and Peruvian Amazonia (cis- More recently, Torres-Carvajal (2004) reported the Andean samples). The unicolor trans-Andean popu- presence of a fourth species, M. melanolomus, in the lations from Ecuador and Colombia, traditionally as- drier and lower areas of Isla de La Plata. This is a signed to M. boddaerti by Stuart (1941) and Peters polytypic, widespread Central American species in and Orejas-Miranda (1986), are considered here as a which the nominal subspecies is characterized by a distinct, undescribed species of the genus (Montin- predominantly uniform and unicolor pattern (Stuart, gelli, 2009); herein, it is recognized as a separate tax- 1941; Peters and Orejas-Miranda, 1986). The identi- on that is designated Mastigodryas sp. fication of M. melanolomus in Ecuador and the range Head length (HL) was measured to the nearest extension that it represents from Central America to 0.01 mm with digital calipers. Total length (TTL) 190 Revalidation of Herpetodryas reticulata

TABLE 1. Measurements of the specimens of Mastigodryas reticu- Francisco (CAS); Coleção Herpetológica da Univer- latus in mm. HL: head length, TTL: total length, TAL: tail length. sidade de Brasília, Brasília (CHUNB); Museo de An- Missing data are represented by a dash. The holotype is desig- fibios y Reptiles, Fundación Herpetológica Gustavo nated with an asterisk. Orcés, Quito (FHGO); Field Museum of Natural His- Specimen Sex HL TTL TAL tory, Chicago (FMNH); University of Kansas, Law- AMNH 126612 F 37.60 1365 368 rence (KU); Museo Ecuatoriano de Ciencias Natura- AMNH 28984 F 32.89 1092 296 les, Quito (MECN); Museu Paraense Emílio Goeldi, CAS 8802 F 27.06 977 274 Belém do Pará (MPEG); Museu de Zoologia da Uni- FHGO 6123 F 28.24 1447 292 versidade de São Paulo, São Paulo (MZUSP); Mu- FHGO 8390 F — — 225 seum of Comparative Zoology, Harvard University, FHGO 8391 F 20.75 — — FHGO 6756 F 26.20 1342 280 Cambridge (MCZ); Museo de Zoología, Pontifícia KU 84670 F 28.08 — — Universidad Católica de Ecuador (QCAZ); Museo MZUSP 19387 F 33.20 1169 321 de la Escuela Politécnica Nacional, Quito (EPN); MZUSP 19388 F 37.49 1337 340 Universidade Federal do Ceará, Fortaleza (UFC); QCAZ 7441 F 29.09 1104 211 Universidade Federal do Mato Grasso, Cuiabá QCAZ 946 F 23.61 1163 243 (UFMT); Universidade Federal da Paraíba, João Pes- QCAZ 945 F 19.27 954 194 soa (UFPB); University of Michigan, Museum of Zo- QCAZ 3968 F 24.83 1360 267 ology, Ann Arbor (UMMZ); United States National QCAZ 4054 F 27.79 1328 367 Museum, Smithsonian Institution, Washington, D.C. USNM 211049 F 29.30 915 259 USNM 211050 F 19.23 646 191 (USNM); Zoologische Museum, Berlin (ZMB). AMNH 28987 M — — — Specimens examined are listed in Appendix I. KU 301093 M 26.00 840 236 QCAZ 7440 M 23.33 1193 229 USNM 211050 M 39.11 1257 354 RESULTS AMNH 22099 — 38.06 — — QCAZ 3945 — 18.62 — — Mastigodryas reticulatus (Peters, 1863) ZMB 4504* — — — — Figures 1-3, 4A

Herpetodryas reticulata Peters, 1863: type locality: and tail length (TAL) were measured to the nearest Guayaquil. 1 mm by carefully aligning the specimens along a Dryadophis boddaerti boddaerti: Stuart, 1941: 66, ruler. Measures of TTL, TAL, and subcaudal counts part. were not recorded for specimens with broken tails. Mastigodryas boddaerti boddaerti: Pérez-Santos and Ventral scales were counted from the first scale that Moreno, 1991: 245-246, part. is distinctly wider than long (described by Myers Mastigodryas melanolomus: Torres-Carvajal, 2004: [2003] and Zaher et al. [2008]) to the last scale an- 85. teriorly adjacent to the cloacal scale. Sex was deter- mined by noting the presence or absence of hemi- Holotype – Zoologische Museum, Berlin (ZMB penes by making a small incision at the base of the 4504), an adult specimen collected by Von C. Reisf tail. Maturity of the specimens was checked follow- near Guayaquil, Ecuador (02°10’S, 79°50’W). ing the methods of Slip and Shine (1988). Methods for hemipenial preparation and terminology fol- Diagnosis – Mastigodryas reticulatus differs from all low those of Zaher (1999) and Zaher and Prudente the other members of the genus by the following com- (2003). Scale counts, sex, and measurements are bination of characters: (1) presence of a striped dor- given in Tables 1 and 2. sal pattern, with the upper, light lateral stripe formed We examined 276 specimens, as follow: 24 Mas- by Scale Rows 4 and 5; (2) dorsal scales with dark tigodryas reticulatus (Peters, 1863); 186 M. bod- apical edges; (3) light, immaculate throat and venter; daerti; 33 to M. heathii; and 33 Mastigodryas sp. (4) higher number of ventral scales; (5) elongate bas- Specimens of Mastigodryas were provided by the al and nude region of the hemipenis; (6) presence of following institutions (institutional codes in paren- some enlarged and thinner spines lateral to the sulcus theses): American Museum of Natural History, New spermaticus, on the distal region of the body of the York (AMNH); California Academy of Sciences, San hemipenis. TABLE 2. Summary of scale counts of the species compared with Mastigodryas reticulatus. Numbers in parentheses are supralabials (SL) and infralabials (IL) in contact with the orbit and genials, respectively; temporal scales (T), primary and secondary; dorsals (D) counted on the anterior, middle and posterior regions of the body; ventral scales (V) and subcaudal scales (SC).

Species SL IL T D V SC

Mastigodryas reticulatus 9 / 9 (4, 5, 6) (N = 21) 10 / 10 (1–5) (N = 13) 2+2/2+2 (N = 16) 17 / 17 / 15 181-206 102-116 9 (3, 4, 5) / 10 (4, 5, 6) (N = 1) 9 / 9 (1–5) (N = 4) 2+2/2+3 (N =6) (N = 24) (N = 21) (N = 20) 9 / 10 (4, 5, 6) (N = 1) 9 / 9 (1–4) (N = 1) 2+3/2+3 (N =1) 10 / 10 (4, 5, 6) (N = 1) 10 (1–5) / 9 (1–4) (N = 3) 1+2/1+2 (N =1) 10 (1–5) / 9 (1–5) (N = 1) 11 / 11 (1-5) (N = 1)

Mastigodryas boddaerti 9 / 9 (4, 5, 6) (N = 157) 10 (1-5) (N = 155) 2+2/2+2 (N = 112) 17 / 17 / 15 177-200 90-118 10 / 10 (4, 5, 6) (N =7) 9 (1-5) (N = 13) 2+2/2+3 (N = 35) (N = 186) (N = 185) (N = 158)

10 (5, 6, 7) / 9 (4, 5, 6) (N = 13) 10 (1-5) / 11 (1-5) (N =8) 2+3/2+3 (N = 23) Montingelli, G. 11 (5, 6, 7) / 9 (4, 5, 6) (N =4) 11 / 11 (1-5) (N =5) 1+2/1+2 (N =7) 10 (4, 5, 6) / 11 (5, 6, 7) (N =4) 10 (1-5) / 9 (1-5) (N =4) 2+2/1+2 (N =3) 8 / 8 (4, 5, 6) (N =1) 11 (1-5) / 12 (1-5) (N =1) 2+3/1+2 (N =2) 2+2/2+4 (N =1) 3+2/3+2 (N =1) al. et

Mastigodryas sp. 9 / 9 (4, 5, 6) (N = 25) 9 / 9 (1-5) (N = 16) 2+2/2+2 (N = 23) 17 / 17 / 15 173-195 92-112 9 (4, 5, 6) / 8 (3, 4, 5) (N =5) 10 / 10 (1-5) (N =11) 2+2/2+3 (N =7) (N = 33) (N = 33) (N = 24) 8 (3, 4, 5) (N =2) 8 (1-4) / 8 (1-5) (N =3) 2+3/2+3 (N =2) 10 / 10 (4, 5, 6) (N =1) 9 / 9 (1-4) (N =1) 1+2/2+1 (N =1)

Mastigodryas heathii 9 / 9 (4, 5, 6) (N = 23) 10 / 10 (1-5) (N = 21) 2+2/2+2 (N = 16) 17 / 17 / 15 179-202 98-115 8 (3, 4, 5) / 9 (4, 5, 6) (N =6) 9 / 9 (1-5) (N =9) 2+2/2+3 (N =8) (N = 33) (N = 32) (N = 25) 9 (4, 5, 6) / 10 (4, 5, 6) (N =2) 9 (1-4) / 10 (1-5) (N =3) 2+3/2+3 (N =4) 9 (4, 5, 6) / 10 (5, 6, 7) (N =1) 1+1/1+1 (N =1) 8 / 8 (4, 5) (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) 191 192 Revalidation of Herpetodryas reticulata

FIGURE 1. Holotype of Herpetodryas reticulata Peters, 1863 (ZMB 4504), from Guayaquil, Ecuador.

Redescription of the Holotype Intraspecific variation

Pholidosis – Rostral scale wider than high; interna- Measurements (mm) – (N = 20; 12 females, 4 males, sals and prefrontals paired, as wide as long; supra- 4 juveniles). Largest female with following measure- oculars, frontal, and parietals about twice as long ments: HL 28.24, TTL 1.45, TAL 292. Largest male: as wide; one loreal approximately twice as long as HL 39.11, TTL 1.26, TAL 354. Tail 19.8% TTL for high; one preocular higher than wide, with the tip vis- females and 28.2% for males (Table 1). ible from above; two postoculars, upper scale larger with the tip visible dorsally; nine supralabials, with Pholidosis – Total ventral count range 181-206 (mean the fourth, fifth and sixth in contact with the orbit; 194.8 ± 6.26; N = 21), not significantly different be- two anterior and two posterior temporals, longer than tween females and males (females with 181-206, wide and obliquely oriented, anterior scales in contact mean 195.7 ± 5.98; N = 16; males with 185-198, mean with supraoculars and postoculars; mental triangular; 192.6 ± 6.80; N = 3). Cloacal scale divided and sub- 10 infralabials with first five in contact with first pair caudals paired in all specimens examined. Total sub- of genials; two pairs of genials, posterior genial about caudal range 102-116 (mean 106.0 ± 3.46; N = 20), twice as long as the first pair. Dorsal scales smooth, not significantly different between males and females in 17-17-15 rows, with two apical pits; ventrals 198; (females 102-116, mean 105.8 ± 3.61; N = 15; males cloacal scale divided; subcaudals 106. 103-112, mean 106.7 ± 3.77; N = 4). All specimens with smooth dorsal scales with two apical pits and Dorsal Pattern and Coloration – Head uniform undifferentiated vertebral row, and same formula brown dorsally; first supralabials light cream, and 17/17/15 with reduction from 17 to 15 scales occur- the posterior four smudged with the same color as ring posteriorly on Scale Rows 3 and 4. Loreals al- the dorsum, bluish gray; ocular band dark brown, ways present, approximately twice longer than high; extending from the eye to the posterior temporals temporal scales vary in 1 + 2, 2 + 2 and 2 + 3 (2 + 2, (slightly faded); infralabials and gulars light cream, N =16; 2+2 / 2+3,N =6; 2+3,N =1; 1+2,N = 1); immaculate as the entire venter; lateral portion of almost all specimens (including holotype) with nine the ventral scales bluish gray as the dorsum; dorsum supralabials with the Supralabials 4-6 in contact with without vestiges of light lateral stripes, scales with the orbit (except for one female with 10 supralabi- dark apical edges. als on both sides, and two females with individual Montingelli, G. G. et al. 193 variation, with nine and ten scales). Most specimens Hemipenis (Figure 4A) – We examined fully everted with 10 infralabials (females, N = 9; males, N =1; and almost maximally expanded hemipenes of three three unsexed specimens), one female with 11 scales, individuals (KU 301093, USNM 211050, QCAZ three females and two males with nine scales, and 7441). The unilobed hemipenis has a simple centro- four specimens exhibiting individual variation with linear sulcus spermaticus, which terminates at the nine and ten scales; genials 2 + 2, 1-5 in contact with apex of the lobe. The lobe is covered with calyces, the first pair (N = 20) or 1-4 in contact (N = 3); one and is spinulated proximally and papillated distally. or two pairs of scales between second pair of genials. Spinules become larger and thinner toward the base of the lobe. The distal end of the hemipenial body Dorsal Pattern and Coloration – The dorsum is is covered with medium-sized spines that are larger brown or bluish gray in all specimens. Two speci- on the left side of the sulcus. The remaining portion mens (FHGO 8390, QCAZ 946) exhibit vestiges of a of the body lacks ornamentation. Basal pockets pres- juvenile banded pattern on the anterior portion of the ent on the proximal end of the body. The papillated body. Some specimens (AMNH 28984, KU 84670, and naked portions of the organ are longer and have 301093; MZUSP 3091) have a faint upper light lateral approximately the same size, whereas the spinulated stripe situated on dorsal Scale Rows 4 and 5 on the an- part of the body represents one third of the organ. 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. Comparisons Only three individuals have a slightly smudged venter (AMNH 28984, FHGO 6756, QCAZ 946), which Samples here assigned to Mastigodryas reticula- usually is darker than in the remaining specimens. tus differ from cis-Andean M. boddaerti by the dark interstitial skin and dark apical edges of dorsal scales Coloration in Life – Photographs of a freshly pre- (vs. light cream interstitial skin and without dark served specimen from Guayas (Figures 2 and 3) pro- apical edges; by a higher number of ventral scales vide information on the coloration in life. The dorsum (181-206, X = 194.8, N =21vs. 177-200, X = 189, 4, is light brown with the anterior and posterior region N = 185); by the elongate basal and nude region of of the body slightly olive or yellow (MZUSP 19387, the hemipenis; and by the presence of some enlarged, FHGO 3223) and with dark apical edges on the dor- thinner spines lateral to the sulcus spermaticus on the sal scales. The Dorsal Row 1 is light cream, as is the distal region of the body (vs. a short, basal nude re- venter, which is light and immaculate. The supralabi- gion lacking enlarged spines distally; Figure 4). The als are yellowish and slightly smudged dorsally. The species differs from cis-Andean Mastigodryas sp. by yellowish color pattern predominates on the ventral a light, immaculate throat and venter (vs. dark, spot- surface of the head and first ventrals, fading posteri- ted throat, and dark-edged ventrals); by more ventrals orly toward an immaculate cream color that extends (181-206, X = 194.8, N =21vs. 173-195, X = 182.3, to the tip of the tail ventrally. N = 33); and by the presence of enlarged, thin spines

FIGURE 2. Freshly preserved specimen of Mastigodryas reticula- FIGURE 3. Freshly preserved specimen of Mastigodryas reticulatus from Guayas, Ecuador (MZUSP 19387). tus from Guayas, Ecuador (MZUSP 19387). 194 Revalidation of Herpetodryas reticulata

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. lateral to the sulcus spermaticus on the distal re- reticulatus occurs in lowland (up to 600 m a.s.l.), de- gion of the body (vs. 1 or 2 well-developed, hook- ciduous and semideciduous forests (Sierra 1999) from like spines; Figure 4). Mastigodryas reticulatus dif- central and southwestern Ecuador, south to Caraquez fers from M. heathii by the width and the position of Bay (Manabí Province) and the provinces of Santa the upper light lateral stripe that occupies two dorsal Elena, Guayas, and Loja, and in the continental is- Scale Rows 4 and 5 (vs. 3 dorsal Scale Rows, 3-5) and lands of Puna and La Plata. Its distribution coincides by the absence of hooklike spines lateral to the sul- with one of the most important and threatened areas cus spermaticus (vs. the presence of hook-like spines) of western Ecuador – the Tumbesian Region, which (Figure 4). harbors a high number of endemic species that in- cludes plants, birds, mammals, and reptiles (Aguirre- Mendoza and Kvist 2005, Cisneros-Heredia 2006, Geographical Distribution (Figure 5) Valencia et al., 2010). Mastigodryas heathii inhabits arid regions of southwestern Ecuador, occurring in Mastigodryas reticulatus, M. heathii, and Mas- the provinces of Loja and El Oro, and arid regions of tigodryas sp. have restricted ranges. Mastigodryas northwestern Peru, along the Cordillera Occidental, Montingelli, G. G. et al. 195

FIGURE 5. Geographical distribution of Mastigodryas boddaerti, M. heathii, M. reticulatus, and Mastigodryas sp. from Lima to Piura provinces, from sea level to near- In summary, we hypothesize that three species of ly 2000 m. Mastigodryas sp. occurs along the humid genus Mastigodryas inhabit Ecuador. We recognize forests of northwestern Ecuador and southwestern the presence of M. heathii and M. pulchriceps, but Colombia, on the western slopes of the Cordillera resurrect M. reticulatus as a valid species restricted Occidental, from sea level to nearly 2000 m. Mas- geographically to southwestern Ecuador. Moreover, tigodryas boddaerti has one of the most widespread a probable fourth, predominantly unicolored species distributions in the genus, occurring predominantly occurs in western Ecuador. Populations of this new on low, humid areas of western South America, from species have been incorrectly identified as belong- Bolivia, Peru, and Colombian and Brazilian Amazo- ing to M. boddaerti. Given the recognition of M. re- nia, to the eastern portion of Venezuela. ticulatus and Mastigodryas sp. as distinct species, the distributional range of the formerly polytypic M. boddaerti (Montingelli and Zaher, 2011) should DISCUSSION exclude the trans-Andean portion of Ecuador and Colombia. We consider the unicolored specimens identified as Mastigodryas melanolomus by Torres-Carvajal (2004; QCAZ 945, 946) to represent M. reticulatus, RESUMO rather than M. melanolomus. The latter probably does not occur in Ecuador. The Colombian specimens of Na presente contribuição, o nome Herpetodryas “M. melanolomus” examined by Perez-Santos and reticulata é revalidado e atribuído a população do gê- Moreno (1988) also are thought to belong to a dis- nero Mastigodryas distribuída nas florestas secas da tinct, undescribed species assignable to the M. bod- região central e sudoeste do Equador. Mastigodryas daerti Group. Therefore, we restrict the distribution reticulatus (Peters, 1863) se caracteriza por apresen- of the M. melanolomus Group to Central America, tar padrão dorsal estriado, com a listra lateral superior with its southernmost limit in Panamá. formada por duas fileiras de escamas, números quarto 196 Revalidation of Herpetodryas reticulata

e cinco; escamas dorsais possuem ápices escuros; (Colubridae: Dipsadinae). American Museum Novitates, ventre da cabeça e do corpo claros e imaculados; nú- 3391:1-47. mero elevado de escamas ventrais; região basal e nua Pérez-Santos, C. and A. G. Moreno. 1988. Ofidios de Colombia. Monografie. VI. Museo Regionale de Scienze Naturali, do hemipênis longa, e presença de espinhos aumenta- Torino. dos e finos, presentes na lateral do sulco espermático, Peters, J. A. and B. Orejas-Miranda. 1986. Catalogue of the na região distal do órgão. Esta espécie é incluída no Neotropical Squamata, Part I. Snakes. Revised Smithsonian Grupo M. boddaerti, juntamente com M. boddaerti e Institution Press, Washington. M. heathii. Peters, W. 1863. Über einige neue oder weniger bekannte Schlangenarten des zoologischen Museums zu Berlin. Monatsberichte Akademie der Wissenschaften, Berlin: 272-289. ACKNOWLEDGMENTS Slip, D. J. and R. Shine. 1988. The reproductive biology and mating system of Diamond Pythons, Morelia spilota We thank the following curators and colleagues for provid- (Serpentes, Boidae). Herpetologica, 44:396-404. ing access to the specimens under their care and for allowing Sierra, R. 1999. Propuesta Preliminar de un Sistema the preparation of hemipenial material: D. Frost and D. Kizirian de Clasificación de la Vegetación para el Ecuador (AMNH); A. Leviton, R. Drewes, and J. Vindum (CAS); G. R. Continental, Proyecto INEFAN/GEF-BIRF, Ecociencia. Colli (CHUNB); Ana Almendariz (EPN); Katty Garzón (FHGO); Quito, Ecuador. M. Kearney and A. Resetar (FMNH); W. E. Duellman and L. Stuart, L. C. 1932. Studies on Neotropical Colubrinae. I. Trueb (KU); J. Losos, J. Rosado, and J. Woodward (MCZ); Mario The taxonomic status of the genus Drymobius Fitzinger. Yánez-Muñoz y Manuel Morals-Mite (MECN); A. L. Prudente Occasional Papers of the Museum of Zoology, University of (MPEG); Omar Torres-Carvajal (QCAZ); D. M. Borges-Nojosa Michigan, 236:1-16. (UFC); M. A. de Carvalho (UFMT); G. Calazans (UFPB); R. Stuart, L. C. 1933a. Studies on Neotropical Colubrinae. II. Some Nussbaum and G. Schneider (UMMZ); G. Zug, R. Heyer, R. Mc- new species and subspecies of Eudryas Fitzinger, 1843, Diarmid, R. Wilson, and T. Harstell (USNM); W. Günther and with an annotated list of the forms of Eudryas boddaertii C. Kucharzewsky (ZMB). We are also grateful to A. Percequillo (Sentzen). Occasional Papers of the Museum of Zoology, and L. Trueb for revising an earlier version of this manuscript, C. University of Michigan, 254:1-10. de Castro-Mello (MZUSP) for providing technical support, and Stuart, L. C. 1933b. Studies on Neotropical Colubrinae. III. The J. Prado for map construction. This research was supported by Taxonomic status of certain Neotropical racers. Copeia, grants from the Coordenação de Aperfeiçoamento de Pessoal de 1933:9-10. Nível Superior (CAPES), Conselho Nacional de Pesquisa e De- Stuart, L. C. 1938. Studies on Neotropical Colubrinae. VI. A new senvolvimento Científico (CNPq), American Museum of Natural species of Eudryas from South America. Copeia, 1938:7-8. History (AMNH), California Academy of Sciences (CAS) and Stuart, L. C. 1939. A new name for the genus Eudryas Fitzinger United States National Museum (USNM) to GGM, Fundação de 1843. Copeia, 1939:55. Amparo à Pesquisa do Estado de São Paulo (BIOTA/FAPESP) Stuart, L. C. 1941. Studies on Neotropical Colubrinae. VIII. A and Conselho Nacional de Desenvolvimento Científico e Tec- revision of the genus Dryadophis Stuart 1939. Miscellaneous nológico (CNPq) to HZ, and Fundación Herpetológica Gustavo Publications Museum of Zoology, University of Michigan, Orcés and Secretaria Nacional de Educación Superior, Ciencia y 49:1-106. Tecnología del Ecuador (SENESCYT) Project No. PIC-08-00470 Tipton, B. L. 2005. Snakes of the America: Checklist and Lexicon. granted to Pontifícia Universidad Católica del Ecuador. Krieger Publishing Company, Melbourne. Torres-Carvajal, O. 2004. The herpetofauna of Isla de La Plata, Ecuador. Herpetological Review, 35:85. LITERATURE CITED Valencia, J. H., G. Vaca-Guerrero, and K. Garzón. 2010. Natural history, potential distribution and conservation status of the Aguirre-Mendoza, Z. and L. P. Kivist. 2005. Florisitic composition Manabi Hognose Pitviper Porthidium arcosae (Schatti and and conservation status of the dry forest in Ecuador. Lyonia, Kramer, 1993) in Ecuador. Herpetozoa, 23:31-43. 8:41-67. Zaher, H. 1999. Hemipenial morphology of the South American Boulenger, G. A. 1894. Catalogue of snakes in the British xenodontine snakes, with a proposal for a monophyletic Museum (Natural History). London: BMNH. v. 2, i-ix, Xenodontinae and a reappraisal of colubroid hemipenes. 382 p.; pls. I-XX. Bulletin of the American Museum of Natural History, Cisneros-Heredia, D. F. 2006. Distribution and ecology of the 240:1-168. western Ecuador frog Leptodactylus labrosus (Amphibia: Zaher, H. and A. L. C. Prudente. 2003. Hemipenes of Siphlophis Anura: Leptodactylidae). Zoological Research, 27:225-234 (Serpentes: Xenodontinae) and techniques of hemipenial Montingelli, G. G. 2009. Revisão Taxonômica do Gênero preparation in snakes: a response to Dowling. Herpetological Mastigodryas Amaral, 1934 (Serpentes, Colubridae). Unpubl. Review, 34:295-302. Ph.D. thesis, Universidade de São Paulo, São Paulo, Brasil. Zaher, H., M. E. Oliveira, and F. L. Franco. 2008. A new Montingelli, G. G. and H. Zaher. 2011. New species of brightly colored species of Pseudoboa Schneider, 1801 from Mastigodryas Amaral, 1934 from Brazilian Amazonia and the Amazon Basin (Serpentes: Xenodontinae). Zootaxa, Guyana (Serpentes: Colubridae). Journal of Herpetology, 1674:27-37. 45:111-119. Myers, C. W. 2003. Rare snakes-five new species from eastern Submitted: 11 November 2011 Panama: reviews of northern Atractus and southern Geophis Accepted: 13 December 2011 Montingelli, G. G. et al. 197

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.