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94 IGUANA  •  VOLUME 13, NUMBER 2  •  JUNE 2006

Santa Lucia is a cloud forest reserve of some 820 hectares in
a lushly forested section of Pichincha Province, Ecuador.

Draped over the steep slopes of the Andean Cordillera
Occidental, the reserve lies between elevations of about
1400–2600 m. Because the region receives about 2,000 mm of
rain per year, the precipitous slopes host an astonishing plant
and animal diversity despite the rugged terrain and the often
chilly climate. The large Maquipucuna Reserve borders Santa
Lucia to the south, protecting an additional 6000 hectares.
Together, these reserves protect an extensive complex of pristine
and regenerating subtropical cloud forest that makes up much
of the Rio Umachaca drainage.

Santa Lucia is a local conservation and eco-tourism coop-
erative established by a number of land-owning campesino fam-

ilies to find a sustainable alternative to the agriculture and
ranching for which a small fraction of the less precipitous slopes
were formerly used. About 84% of Santa Lucia’s land is primary
cloud forest. The remaining area is about evenly divided
between secondary forest and pastured hillsides.

Santa Lucia hosts an astounding variety of plant and ani-
mal species. Particularly speciose groups include birds, arthro-
pods, and orchids. The current bird list exceeds 370 species and
over 340 species of orchids (among nearly 2000 vascular plant
species) have been documented in the neighboring
Maquipucuna Reserve (Webster and Rhode 2001, 2005).

Although the reptilian fauna of Santa Lucia is not nearly
so diverse as the groups mentioned above, many of the species
are endemic and have relatively small ranges. Additionally, our

Reptiles of the 
Santa Lucia Cloud Forest, Ecuador

Aaron Z. Savit

Department of Biology, Earlham College, Richmond, Indiana

Photographs by the author.

�

SAVIT

Thick clouds of fog settle on the mountains at night and often linger well into the day.

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IGUANA  •  VOLUME 13, NUMBER 2  •  JUNE 2006 95

knowledge of most of these species is so scant that their natu-
ral history and habitat preferences usually have not been
described in the literature. Perez-Santos and Moreno (1991)
published a volume that renders the chaos of identification a
bit less imposing, at least as far as snakes are concerned, but no
comparable treatment covers Andean lizards, although some
fairly comprehensive treatments of entire genera (i.e., Harris
1994, Torres-Carvajal 2001) make diagnosis a bit more feasi-
ble. However, the following list is the first that documents the
reptilian species from an Andean cloud forest site. I have made
every effort to present information in a manner that does not
require prior knowledge of obscure Andean taxa and to gather
and present all available ecological and distributional data, both
from the literature and my own observations.

This list represents the squamate species I found at Santa
Lucia over a five-month period from November 2004 to April
2005. Most were caught along a ridge at elevations from
1800–2000 m. Several additional species are probably present
in the reserve, especially at its upper and lower reaches. I par-
ticularly suspect that one or more species of elusive, litter-
dwelling Spectacled Lizards (“microteiids” in the family
Gymnophthalmidae) escaped detection. One species in the
gymnophthalmid genus Proctoporus from the lower reaches of
the reserve remains unidentified, as does a species of Ground

Snake (Atractus, probably A. major). Additionally, the reserve’s
guides and staff reported several other species of snakes, but
none could be captured and identified.

Readers should bear in mind that considerable geographic
variation in coloration exists for many of the species described.
However, the color variations mentioned herein refer solely to
documented differences within the populations at Santa Lucia.
Any information not specifically cited is based on my own
observations and measurements.

L I Z A R D S

Anolis aequatorialis Werner 1894
Description: This relatively large anole has snout-to-vent (SVL)
lengths that can exceed 80 mm. Dorsal color patterns include
a series of dark transverse markings with mottled greens,
browns, and blacks, with juveniles exhibiting significantly more
brown and black coloration. The striking dewlap has greenish-
yellow lines where it comes into contact with the body and
throat, but these quickly degenerate into a series of brick-red
blotches on a black background. Hints of turquoise often
appear on the anterior part of the dewlap near the throat. Both
males and females of the species possess dewlaps, although
those of males are relatively larger in individuals of similar sizes.
Juveniles also have an ocellum on each side of the head poste-

Situated at the end of a ridge, the lodge at Santa Lucia overlooks lush valleys on three sides.

REPTILES OF THE SANTA LUCIA CLOUD FOREST

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rior to the tympanum. This appears to dissipate in older males,
but is retained in adult females.

Ecology: This species is most often found on well-shaded leaf
litter, although juveniles are frequently seen in low undergrowth
near the ground. These lizards seem to prefer cooler substrates
and air temperatures, both of which averaged around 18.5 ºC.
Average body temperature of 21 A. aequatorialis was 20.5 ºC.
The elevational range for the species is roughly 1000–2000 m
(Torres-Carvajal 2001).

Anolis gemmosus O’Shaughnessy 1875
Description: This anole reaches just over 60 mm SVL. An
astounding diversity of color phases can occur within a single
population. Perhaps the most common phase at Santa Lucia is
the patterned phase, in which males have a series of transverse
rows of black spots. In females, this pattern manifests itself as a
caudal series of spade-shaped black blotches. Other color phases
include a nearly solid green, which occurs in both males and
females, and a female morph with a wide, light-brown vertebral
stripe bordered by two thinner gray stripes on a green back-
ground. In a very rare variation of the male patterned phase, the
spaces between the rows of black spots are filled with large
white markings.

Ecology: These anoles appear to prefer cooler environmental
temperatures much like those associated with A. aequatorialis.
Average substrate and air temperatures were both around 19 ºC.
Body temperature of 34 A. gemmosus averaged 21.1 ºC. Anolis
gemmosus differ from A. aequatorialis in that they are completely
arboreal. The elevational range is roughly 2000–3000 m (Torres-
Carvajal 2001). These lizards are most frequently encountered
along forest edges, but never in direct sun. The preference for
edge habitat is probably an artifact of increased arthropod prey
abundance in those areas.

Cercosaura vertebralis (O’Shaughnessy 1879)
Description: A gymnophthalmid of medium size, Santa Lucia
Cercosaura vertebralis had SVLs to 69 mm, although adults were
usually below 65 mm SVL. These lizards have a prominent 2–3
scale-wide vertebral stripe that starts as light orange on top of
the head and fades through yellow to become white over the
sacrum. Dorsal background color is black to dark brown.
Golden-yellow subocular stripes run from the tip of the snout
under the tympanum and terminate at the front limbs. White-
centered ocelli often are present on the flanks around the fore-
and hindlimbs. More prominent ocelli are frequently accompa-
nied by red highlights on the flanks and a red venter mottled
with black. In other individuals, the venter would be cream

96 IGUANA  •  VOLUME 13, NUMBER 2  •  JUNE 2006 SAVIT

Edge habitats, such as this partially forested slope, provide favorable conditions for some lizards such as Anolis gemmosus and Cercosaura vertebralis.

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with extensive black mottling. This coloration was displayed far
more frequently in animals collected between November and
January than in those collected from February to April. This
suggests that red on the venter and flanks, as well as the
enhanced ocelli, may be breeding coloration.

Ecology: Cercosaura vertebralis belongs to a group of three
gymnophthalmid genera that have recently been shown to be
monophyletic (Doan 2003). The species is highly heliophilic,
and individuals are mostly active in dry leaf litter. However,
they can be seen both foraging and basking on soil, litter, and
deadfall anywhere where sunlight is sufficient. They may be
completely absent from the surface for days if weather is not
suitable. Substrate and air temperatures for sites at which C. ver-
tebralis was found averaged 26.8 and 21.9 ºC, respectively.
These lizards maintain relatively high body temperatures, which
averaged 27.9 ºC (n = 20). The species has been recorded at ele-
vations as low as 700 m and as high as 2500 m and ranges from
northern Colombia and far southern Panamá through Ecuador
and the far north of Perú (Uzzell, 1973).

Proctoporus oculatus (O’Shaughnessy 1879)
Description: This impressive gymnophthalmid can grow as
large as 88 mm SVL (Kizirian 1996) and have a tail more than
twice that long. The dorsum is dark brown with two faint dor-
solateral bands extending from the head and dissipating near
the pelvis. A series of yellow-centered ocelli extend from the
neck to the base of the tail. The black-and-white mottled ven-
ter gradually fades into a very striking pattern of red and black
along the flanks.

IGUANA  •  VOLUME 13, NUMBER 2  •  JUNE 2006 97REPTILES OF THE SANTA LUCIA CLOUD FOREST

Anolis aequatorialis. Top: male (left, 80 mm SVL) and female (right,
76 mm SVL); bottom: male (79 mm SVL).

Anolis gemmosus. Color phases: transverse bars (top), white blotches
(middle), and striped (bottom).

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Ecology: This species occurs in only a few drainages along the
western Ecuadorian Andes, and has been previously recorded
in sympatry with P. unicolor (Kizirian, 1996). They are rarely
seen on the surface, and prefer the cool temperature and cover
afforded by abundant leaf litter.

Proctoporus unicolor (Gray 1858)
Description: This medium-sized gymnophthalmid grows to
slightly over 60 mm SVL. Dark brown dorsally, it has two light
dorsolateral stripes that extend over the shoulders and dissipate
quickly. A small amount of white spotting may be present on the
throat or flanks. A bright red accents the ventral part of the flanks,
and extends most of the way down the tail. The underside of the
jaw is white mottled with black. Otherwise, the venter is solid
black. Kizirian (1996) indicated that considerably more color vari-
ation exists than was observed at Santa Lucia, where all P. unicolor
generally adhered to the description above.

Ecology: Like P. oculatus, this species is limited in range to the
northwestern Ecuadorian Andes. It has been previously
reported at elevations from 2390–3300 m (Kizirian 1996).
These lizards appear to prefer cool temperatures, and quickly
turn lethargic when held in the heat of a human hand. They
forage under the thick cover of damp leaf litter and hardly ever
emerge on the surface. They have extremely powerful jaws and
a strong tail that can be up to 250% of SVL if unbroken and
not regenerated.

Proctoporus sp.
Description: Only one individual of this unknown species was
captured. It measured 49 mm SVL and 146 mm total length
(with a complete tail). This lizard had a brown dorsal back-
ground color with broken dorsolateral lines of lighter brown
blotches outlined with black. Flanks had a number of yellow-
centered ocelli that were particularly prominent near the fore-
limbs and decreased in size posteriorly. Venter was white with
regular black spotting forming rows and increasing in intensity
posteriorly. White markings remained near the cloaca, but the
ventral surface of the tail was solid black with a strong irides-
cent blue sheen.

98 IGUANA  •  VOLUME 13, NUMBER 2  •  JUNE 2006 SAVIT

Proctoporus oculatus is rarely seen on the surface. They seem to prefer
the cool temperatures and cover afforded by abundant leaf litter.

Cercosaura vertebralis is heliophilic (sun-loving), and individuals are
mostly active in dry leaf litter.

Proctoporus unicolor quickly turns lethargic when held in the heat of
a human hand. They forage under the thick cover of damp leaf litter
and hardly ever emerge on the surface.

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IGUANA  •  VOLUME 13, NUMBER 2  •  JUNE 2006 99REPTILES OF THE SANTA LUCIA CLOUD FOREST

Ecology: The one individual was found in the lower reaches of
the reserve at an elevation of about 1500 m.

Ptychoglossus plicatus (Taylor 1949)
Description: A medium-sized gymnophthalmid lizard, P. plica-
tus grows to about 65 mm SVL. Head scales are light lime
green, with the exception of supraocular scales, which are black
and have a yellow-green stripe on the very edge. These stripes,
as well as the other colored head scales, join at the back of the
skull to form a dorsal stripe 3–5 scales wide that fades towards
the tail. The dorsal ground color is dark brown. Lateral cream-
colored lines extend back from behind the eyes. These lines are
more prominent in some individuals than in others, but they
usually begin to dissipate around the forelimbs. Flanks are a
deep mahogany red. The venter is a bright salmon-red, fading

to cream around the throat and jaw, with some green tinges
near the very front of the mandible. In situ, P. plicatus can
appear quite similar to Cercosaura vertebralis, but they are eas-
ily distinguished by the distinctive cephalic green coloration
and relatively thicker vertebral stripe.

Ecology: This species is known to occur from Costa Rica to the
Panamá-Colombia border, with another population possibly
occurring slightly farther south in northwestern Colombia
(Harris 1994). Obviously, a population in northern Ecuador
represents a very considerable extension to the range of this
species. Because Santa Lucia lies at the upper extremity of their
documented elevational range of 30–1890 m (Harris 1994),
that more P. plicatus were seen several hundred meters below
the main collection area, which was located just above 1900 m,
is probably indicative of an elevational range mostly below that
of the reserve. This lizard is an active forager that can sometimes
be seen weaving in and out of the leaf litter. They will tolerate
both full shade and mixed sun, and are most often found in
mature forest.

Stenocercus varius (Boulenger 1885)
Description: This stocky lizard in the family Tropiduridae can
grow to about 8.5 cm SVL (Torres-Carvajal 2000). Basic col-
oration consists of regular black markings along the dorsum,
with a dark greenish-brown background and occasional yellow
spots. The top of the head has the same dark background col-
oration, with the region around the mouth and eyes generally
a much lighter shade of green. The flanks also are a lighter
shade of green, with yellow-green spots arranged in regular
transverse rows. The venter is green to greenish-white with a
light orange color on the ventral surface of the tail. These lizards
are capable of rapid color change. They can appear nearly
melanistic at times, with the light green of the mouth and eyes
and the yellow spotting on the flanks completely obscured.

Ecology: Stenocercus varius occurs at elevations from
1460–2200 m. They are restricted to montane forests with high
rainfall, and are frequently found on tree trunks and fallen logs,
particularly in cleared areas. They also occur along forest edges
(Torres-Carvajal 2000). They are quite rare at Santa Lucia,
although they appear to be quite territorial, with one individ-
ual recaptured 3 m from the release site after a period of four
months.

Proctoporus sp.: Only one individual of this unidentified species was found
in the lower reaches of the reserve at an elevation of about 1500 m.

Ptychoglossus plicatus is an active forager that can sometimes be seen
weaving in and out of the leaf litter. They will tolerate both full shade
and mixed sun, and are most often found in mature forest.

Stenocercus varius appears to be restricted to montane forests with high
rainfall, and is frequently found on tree trunks and fallen logs, partic-
ularly in cleared areas.

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S N A K E S

Atractus occidentalis Savage, 1955
Description: This rather small snake in the family Colubridae
reaches lengths a bit under 35 cm total length in mature
females. They are thick snakes with small heads and no obvi-
ous neck differentiation. Symmetrical pairs of black spots mark
the dorsum. Dorsal ground color exhibits some variation, with
both brown and burgundy phases documented. Additional
variation may exist. The venter is black posteriorly and mottled
with a rose-tinged cream anteriorly.

Ecology: This species has been documented as occurring at ele-
vations from 1200–3000 m (Perez-Santos and Moreno 1991).
They are terrestrial snakes that generally forage in the litter of
the forest interior. In mid-February, a heavily gravid female with
SVL = 31 cm (34 cm total length) was encountered basking on
a trail and responded to capture by playing dead. She was kept

for observation, and, about 12 hours after capture, laid two
eggs, each measuring about 3 x 1 cm. Both eggs were fertile and
appeared to be developing when candled. However, they had
not hatched by the time I left the reserve about 3 months later.
They were kept moist and still appeared healthy at that time.

Atractus (major?) Boulenger, 1893
Description: This tentative identification is based on a single
specimen that was captured, but could not be photographed.
Consequently, the species is only tentatively identified as A.
major. This individual displayed a very distinct pattern of bro-
ken, irregular, longitudinal bands running the length of its
body. Markings were brown with black bordering on a brick-
red ground color. Venter was white with some dark blotches.
The single specimen measured about 45 cm total length (38 cm
SVL), which is over the maximum length of most species of
Atractus, but well under the maximum known for the presumed
species (72 cm; Perez-Santos and Moreno, 1991).

Ecology: Atractus major is very widespread throughout the
Amazonian slopes and eastern lowlands of Venezuela,
Colombia, Ecuador, Perú, Bolivia, and Brazil. This species has
been documented at elevations from near sea level to almost
3000 m (Perez-Santos and Moreno, 2001). Given this large ele-
vational range, A. major or a closely related species could very
easily be present on the Pacific slopes of the Andes. This is par-
ticularly likely in light of the Huancabamba Depression, an area
in northern Perú where both Andean cordilleras drop well
below 3000 m.

100 IGUANA  •  VOLUME 13, NUMBER 2  •  JUNE 2006 SAVIT

Atractus occidentalis is a small inconspicuous snake that forages in leaf
litter on the forest floor.

Bothrocophias campbelli are small, stout pit-vipers that are perfectly
camouflaged and almost invisible in leaf litter.

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IGUANA  •  VOLUME 13, NUMBER 2  •  JUNE 2006 101REPTILES OF THE SANTA LUCIA CLOUD FOREST

Bothrocophias campbelli (Freire 1991)
Description: These stout pit-vipers can grow to over half a
meter in length. A pattern of brown angular blotches on a gray
background extends laterally from either side of the middor-
sum. These coalesce into a more cohesive banded pattern
toward the tail. The venter is gray with black speckling, and a
series of regular black splotches occur ventrolaterally.

Ecology: Four species of snakes, including B. campbelli, were
recently allocated to a new genus, with this species being
restricted to the Ecuadorian Cordillera Occidental (Gutberlet
and Campbell 2001). This species has a very restricted range,
especially in comparison with two of its congeners, both of
which have extensive ranges on the eastern side of the Andes
and in parts of the Amazonian lowlands. These snakes are per-
fectly camouflaged and are almost invisible in leaf-litter. They
are inconspicuous, secretive, and nocturnal.

Chironius monticola Roze 1952
Description: This impressive colubrid snake can grow to over
1.5 m in total length. The dorsum is a solid forest-green with
dark outlines of each scale. The venter is a light gray-green,
turning yellow anteriorly. Throat, jaw, and upper labials are a
bright yellow.

Ecology: This terrestrial snake is most often seen basking in the
open on warm, sunny days. It is very alert and can disappear
rapidly into the leaf litter and undergrowth. They range from

Venezuela through Bolivia, and have been documented at ele-
vation from 1200–2000 m (Perez-Santos and Moreno 1991).

Dipsas oreas elegans (Boulenger 1896)
Description: This small, elegant colubrid has a laterally com-
pressed body and a large, blunt head. The body is covered by
alternating bands of brownish-gray and black. The venter is a
silvery gray with black blotches. The long, prehensile tail
accounts for about a third of the snake’s total length.

Ecology: Dipsas oreas is an arboreal snake with an elevational
range of 1200–3000 m (Perez-Santos and Moreno 1991). The
diet consists mostly of invertebrates, especially soft-bodied
species. They are active nocturnally.

Liophis epinephelus albiventris Jan 1863
Description: This medium-sized colubrid snake grows to just
under 80 cm (Perez-Santos and Moreno 1991). Ground color
of the dorsum is deep green. A series of black spots along the

Chironius monticola is very alert and can disappear rapidly into the leaf
litter and undergrowth.

Dipsas oreas elegans is arboreal and has a laterally compressed body and
a large, blunt head. The former character facilitates climbing and the
latter the consumption of relatively large prey.

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dorsum merge posteriorly to form two lateral stripes and a
thicker dorsal stripe. The venter is bright yellow, as are the supe-
rior and inferior labials. Bright orange skin can sometimes be
seen between dorsal scutes.

Ecology: This terrestrial species responds to threats by flatten-
ing its head and body in a convincing viper impersonation.
This particular subspecies is restricted to Ecuador, whereas two
other subspecies occur throughout northern South America as
far north as Panamá and as far south as Perú (Perez-Santos and
Moreno 1991). Virtually nothing is known about the ecology
of this species.

Mastigodryas boddaerti (Sentzen 1796)
Description: A relatively large colubrid snake, Mastigodryas bod-
daerti can reach 160 cm in total length (Perez-Santos and
Moreno 1991). It has a brown dorsum, sometimes with dark
longitudinal stripes. A white subocular stripe extends from the
rostrum to the nuchal region, where it fades into the brown of
the dorsum. The venter is a creamy-yellow with a small amount
of black spotting. Some variation in coloring has been docu-
mented for this species.

Ecology: A fairly widespread species, M. boddaerti ranges from
Colombia to Bolivia and Brazil. It has been documented at ele-
vations ranging from close to sea level to about 2000 m (Perez-
Santos and Moreno 1991). Aside from the fact that it is mostly
terrestrial, not much is known about behavior, habitat use, or
life history.

102 IGUANA  •  VOLUME 13, NUMBER 2  •  JUNE 2006 SAVIT

Mastigodryas boddaerti. Other than being largely terrestrial, little is
known about behavior, habitat use, or life history.

Liophis epinephelus albiventris (female, 41 cm SVL): This terrestrial
species responds to threats by flattening its head and body in a con-
vincing viper impersonation.

Mastigodryas pulchireps is an alert, terrestrial snake found mainly in
forest interior.

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IGUANA  •  VOLUME 13, NUMBER 2  •  JUNE 2006 103REPTILES OF THE SANTA LUCIA CLOUD FOREST

Mastigodryas pulchireps (Cope 1868)
Description: This striking snake has a beautiful pattern consist-
ing of a row of blackish-brown rectangles running along the
dorsum, with another lateral row of rectangles along either
flank. Each rectangle is separated by a whitish-gray stripe. The
tail is almost entirely black. The venter is black with a small
white spot on either side of each scale. Black ocular stripes help
to camouflage the eyes. Maximum size for this species is not
known. The only specimen I caught was 24 cm SVL (32 cm
TL). Specimens at the zoology museum at the Pontifica
Universidad Católica del Ecuador were “somewhat larger” than
the one I caught, but I did not measure them.

Ecology: This species is known from Colombia and Ecuador,
although it has heretofore been documented only as high as
1500 m (Perez-Santos and Moreno 1991). The specimen I
caught in the reserve was at an elevation of about 1750 m. It is
an alert, terrestrial snake found mainly in forest interior. Little
else is known about its life history.

Tantilla melanocephala (Linnaeus 1758)
Description: This small colubrid reaches over 40 cm in total
length. Neonates can be as small as 5 cm SVL. The neck is not
distinctly differentiated. The head is black with a light snout,
postocular stripes, and two white spots on the back of the head.

Dorsal background color is a light brown, and some individu-
als may have a varying number of longitudinal stripes. The ven-
ter is yellow.

Ecology: This is an extremely widespread species, ranging from
Central America to northern Argentina and Uruguay. They can
occur from sea level to 2500 m (Perez-Santos and Moreno
1991). They eat mostly small arthropods and arthropod eggs,
and are often found in association with leaf litter. They were the
most abundant snakes I encountered at Santa Lucia, with four
of 13 snakes belonging to this species.1

Acknowledgments
I thank all the staff and community members of Santa Lucia,
especially Santa Lucia’s administrator, Francisco Molina. I am
indebted to Jorge Noé Morales, Julio Ayala, Carlos Calvache,
Eduardo Tapia, and Arturo Alba Zuñiga for help in collecting
animals. I also thank Dr. Luís Coloma and the Pontifica
Universidad Católica del Ecuador for access to their collections.
Robert Henderson generously helped with identification, advice,
and material support. Robert Powell provided useful advice dur-
ing my time at Santa Lucia and since.

Literature Cited
Doan, T.M. 2003. A new phylogenetic classification for the gymnophthalmid

genera Cercosaura, Pantodactylus and Prionodactylus (Reptilia: Squamata).
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Gutberlet, R.L. and J.A. Campbell. 2001. Generic recognition for a neglected
lineage of South American pitvipers (Squamata: Viperidae: Crotalinae),
with the description of a new species from the Colombian Chocó.
American Museum Novitates (3316): 1–15.

Harris, D.M. 1994. Review of the teiid lizard genus Ptychoglossus. Herpetological
Monographs 8: 226–275.

Kizirian, D.A. 1996. A review of Ecuadorian Proctoporus (Squamata:
Gymnophthalmidae) with descriptions of nine new species. Herpetological
Monographs 10: 85–155.

Perez-Santos, C, and A.G. Moreno. 1991. Serpientes de Ecuador. Museo
Regionale di Scienze Naturali, Torino, Italy.

Torres-Carvajal, O. 2000. Ecuadorian lizards of the genus Stenocercus
(Squamata: Tropiduridae). Scientific Papers, Natural History Museum,
University of Kansas (15): 1–38.

Torres-Carvajal, O. 2001. Lizards of Ecuador: Checklist, distribution, and sys-
tematic references. Smithsonian Herpetological Information Service (131):
1–35.

Uzzell, T. 1973. A revision of lizards of the genus Prionodactylus with a new
genus for P. leucostictus and notes on the genus Euspondylus (Sauria,
Teiidae). Postilla (159): 1–7.

Webster, G.L. and R.M. Rhode. 2001. Plant Diversity of an Andean Cloud
Forest: Inventory of the Vascular Plants of Maquipucuna, Ecuador.
University of California Publications in Botany, Volume 82. Berkeley,
California.

Webster, G.L. and R.M. Rhode. 2005. Maquipucuna monocots.
http://maqui.ucdavis.edu/newest_maqui_monocots.html.

Tantilla melanocephala eats mostly small arthropods and arthropod
eggs, and is often found in association with leaf litter.

1 Editor’s note: Snakes in the mainland Neotropics often are difficult
to find, apparently due to a combination of low population densi-
ties and secretive habits. The low number of encounters at the
reserve also reflects conditions at high elevations, which are rarely
ideal for squamate reptiles.

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