South American Journal of Herpetology, 6(3), 2011, 189-197
© 2011 Brazilian Society of Herpetology
Revalidation of Herpetodryas reticulata (Peters, 1863)
(Serpentes: Colubridae) from Ecuador
Giovanna G. Montingelli1,5, Jorge H. Valencia2,3, Marco A. Benavides4 and Hussam Zaher1
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.
1.
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. Reisf 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 specimens 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 geographically 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 identification 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 attribute full species status to the Mastigodryas population distributed in the dry forests of central and southwestern Ecuador. The available and applicable name
for this population is Mastigodryas reticulatus (Peters, 1863), for which we provide a detailed description 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 (cisAndean samples). The unicolor trans-Andean populations from Ecuador and Colombia, traditionally assigned to M. boddaerti by Stuart (1941) and Peters
and Orejas-Miranda (1986), are considered here as a
distinct, undescribed species of the genus (Montingelli, 2009); herein, it is recognized as a separate taxon that is designated Mastigodryas sp.
Head length (HL) was measured to the nearest
0.01 mm with digital calipers. Total length (TTL)
Revalidation of Herpetodryas reticulata
190
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 designated with an asterisk.
Specimen
AMNH 126612
AMNH 28984
CAS 8802
FHGO 6123
FHGO 8390
FHGO 8391
FHGO 6756
KU 84670
MZUSP 19387
MZUSP 19388
QCAZ 7441
QCAZ 946
QCAZ 945
QCAZ 3968
QCAZ 4054
USNM 211049
USNM 211050
AMNH 28987
KU 301093
QCAZ 7440
USNM 211050
AMNH 22099
QCAZ 3945
ZMB 4504*
Sex
F
F
F
F
F
F
F
F
F
F
F
F
F
F
F
F
F
M
M
M
M
—
—
—
HL
37.60
32.89
27.06
28.24
—
20.75
26.20
28.08
33.20
37.49
29.09
23.61
19.27
24.83
27.79
29.30
19.23
—
26.00
23.33
39.11
38.06
18.62
—
TTL
1365
1092
977
1447
—
—
1342
—
1169
1337
1104
1163
954
1360
1328
915
646
—
840
1193
1257
—
—
—
TAL
368
296
274
292
225
—
280
—
321
340
211
243
194
267
367
259
191
—
236
229
354
—
—
—
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 anteriorly adjacent to the cloacal scale. Sex was determined by noting the presence or absence of hemipenes by making a small incision at the base of the
tail. Maturity of the specimens was checked following the methods of Slip and Shine (1988). Methods
for hemipenial preparation and terminology follow 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 parentheses): American Museum of Natural History, New
York (AMNH); California Academy of Sciences, San
Francisco (CAS); Coleção Herpetológica da Universidade de Brasília, Brasília (CHUNB); Museo de Anfibios y Reptiles, Fundación Herpetológica Gustavo
Orcés, Quito (FHGO); Field Museum of Natural History, Chicago (FMNH); University of Kansas, Lawrence (KU); Museo Ecuatoriano de Ciencias Naturales, Quito (MECN); Museu Paraense Emílio Goeldi,
Belém do Pará (MPEG); Museu de Zoologia da Universidade de São Paulo, São Paulo (MZUSP); Museum 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 Pessoa (UFPB); University of Michigan, Museum of Zoology, 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 combination of characters: (1) presence of a striped dorsal 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 basal 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.
Mastigodryas heathii
Mastigodryas sp.
Mastigodryas boddaerti
Mastigodryas reticulatus
Species
(N = 21)
(N = 24)
2 + 2 / 2 + 3 (N = 6)
2 + 3 / 2 + 3 (N = 1)
1 + 2 / 1 + 2 (N = 1)
9 / 9 (1–5) (N = 4)
9 / 9 (1–4) (N = 1)
10 (1–5) / 9 (1–4) (N = 3)
9 (3, 4, 5) / 10 (4, 5, 6) (N = 1)
9 / 10 (4, 5, 6) (N = 1)
10 / 10 (4, 5, 6) (N = 1)
(N = 185)
(N = 186)
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)
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)
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) / 11 (5, 6, 7) (N = 4)
8 / 8 (4, 5, 6) (N = 1)
1 + 3 / 2 + 1 (N = 1)
8 / 8 (4, 5) (N = 1)
2 + 2 / 1 + 2 (N = 1)
2 + 2 / 1 + 1 (N = 1)
1 + 3 / 1 + 3 (N = 1)
1 + 1 / 1 + 1 (N = 1)
2 + 3 / 2 + 3 (N = 4)
9 (1‑4) / 10 (1‑5) (N = 3)
9 (4, 5, 6) / 10 (4, 5, 6) (N = 2)
9 (4, 5, 6) / 10 (5, 6, 7) (N = 1)
2 + 2 / 2 + 3 (N = 8)
9 / 9 (1‑5) (N = 9)
8 (3, 4, 5) / 9 (4, 5, 6) (N = 6)
179‑202
(N = 32)
17 / 17 / 15
(N = 33)
1 + 2 / 2 + 1 (N = 1)
9 / 9 (1‑4) (N = 1)
10 / 10 (4, 5, 6) (N = 1)
2 + 2 / 2 + 2 (N = 16)
2 + 3 / 2 + 3 (N = 2)
8 (1‑4) / 8 (1‑5) (N = 3)
8 (3, 4, 5) (N = 2)
10 / 10 (1‑5) (N = 21)
(N = 33)
(N = 33)
2 + 2 / 2 + 3 (N = 7)
10 / 10 (1‑5) (N = 11)
9 (4, 5, 6) / 8 (3, 4, 5) (N = 5)
9 / 9 (4, 5, 6) (N = 23)
173‑195
17 / 17 / 15
2 + 2 / 2 + 2 (N = 23)
9 / 9 (1‑5) (N = 16)
9 / 9 (4, 5, 6) (N = 25)
3 + 2 / 3 + 2 (N = 1)
2 + 2 / 2 + 4 (N = 1)
177‑200
17 / 17 / 15
2 + 2 / 2 + 2 (N = 112)
10 (1‑5) (N = 155)
9 / 9 (4, 5, 6) (N = 157)
11 / 11 (1-5) (N = 1)
10 (1–5) / 9 (1–5) (N = 1)
181‑206
V
17 / 17 / 15
D
2 + 2 / 2 + 2 (N = 16)
T
10 / 10 (1–5) (N = 13)
IL
9 / 9 (4, 5, 6) (N = 21)
SL
(N = 25)
98‑115
(N = 24)
92‑112
(N = 158)
90‑118
(N = 20)
102‑116
SC
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).
Montingelli, G. G. et al.
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; internasals and prefrontals paired, as wide as long; supraoculars, 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 visible 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.
Measurements (mm) – (N = 20; 12 females, 4 males,
4 juveniles). Largest female with following measurements: 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).
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.
Pholidosis – Total ventral count range 181‑206 (mean
194.8 ± 6.26; N = 21), not significantly different between 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 subcaudals paired in all specimens examined. Total subcaudal 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 occurring posteriorly on Scale Rows 3 and 4. Loreals always 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 supralabials on both sides, and two females with individual
Montingelli, G. G. et al.
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 specimens (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 anterior 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 venter (AMNH 28984, FHGO 6756, QCAZ 946), which
usually is darker than in the remaining specimens.
193
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 centrolinear 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 present 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
Coloration in Life – Photographs of a freshly preserved specimen from Guayas (Figures 2 and 3) provide 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 dorsal scales. The Dorsal Row 1 is light cream, as is the
venter, which is light and immaculate. The supralabials are yellowish and slightly smudged dorsally. The
yellowish color pattern predominates on the ventral
surface of the head and first ventrals, fading posteriorly toward an immaculate cream color that extends
to the tip of the tail ventrally.
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 region lacking enlarged spines distally; Figure 4). The
species differs from cis-Andean Mastigodryas sp. by
a light, immaculate throat and venter (vs. dark, spotted 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).
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 region of the body (vs. 1 or 2 well-developed, hooklike spines; Figure 4). Mastigodryas reticulatus differs 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 sulcus 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.), deciduous 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 islands 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 includes plants, birds, mammals, and reptiles (AguirreMendoza 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,
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 nearly 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 Amazonia, 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 distinct, 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 belonging 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 gênero Mastigodryas distribuída nas florestas secas da
região central e sudoeste do Equador. Mastigodryas
reticulatus (Peters, 1863) se caracteriza por apresentar padrão dorsal estriado, com a listra lateral superior
formada por duas fileiras de escamas, números quarto
Revalidation of Herpetodryas reticulata
196
e cinco; escamas dorsais possuem ápices escuros;
ventre da cabeça e do corpo claros e imaculados; número elevado de escamas ventrais; região basal e nua
do hemipênis longa, e presença de espinhos aumentados 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 providing 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. McDiarmid, 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 Desenvolvimento 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 Tecnológico (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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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: Humaitá: 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: Fazenda 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; Rondônia: 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.