REVISION OF THE VENOMOUS SNAKES OF BOLIVIA.

II:
THE PITVIPERS (SERPENTES: VIPERIDAE)

Authors: Harvey, Michael B., Aparicio E, James, and Gonzales A,

Lucindo
Source: Annals of Carnegie Museum, 74(1) : 1-37
Published By: Carnegie Museum of Natural History
URL: https://doi.org/10.2992/0097-4463(2005)74[1:ROTVSO]2.0.CO;2

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 ANNALS OF CARNEGIE MUSEUM
VOL. 74, NUMBER 1, PP. 1—37 30 MARCH 2005

REVISION OF THE VENOMOUS SNAKES OF BOLIVIA. II: THE PITVIPERS (SERPENTES: VIPERIDAE)

MICHAEL B. HARVEY
Natural Sciences Department, Broward Community College, 3501 S.W. Davie Road, Davie, FL 33314 (mharvey@broward.edu)

JAMES APARICIO E.
Colección Boliviana de Fauna, La Paz, Bolivia

LUCINDO GONZALES A.
Museo de Historia Natural “Noel Kempff Mercado,” Santa Cruz de la Sierra, Bolivia

ABSTRACT

Twelve species (Bothriopsis bilineata, B. oligolepis, B. taeniata, Bothrocophias microphthalmus, Bothrops andianus, B. atrox, B. jonathani, B.
moojeni, B. neuwiedi, B. sanctaecrucis, Crotalus durissus, Lachesis muta) and five genera of pitvipers are known from Bolivia. Known ranges
of several species are expanded to accommodate recently collected material and Bothrops andianus is reported from Bolivia. The holotypes of
Bothrops andianus and Lachesis peruvianus are redescribed. Bothriopsis oligolepis (Werner) is shown to be synonymous with Lachesis peru-
vianus Boulenger rather than L. chloromelas. We designate a lectotype of Bothriopsis chloromelas (Boulenger) and apply this name to the ornate-
ly patterned forest pitviper endemic to the northern and central Andes of Peru. Morphological variation and reproductive data are reported for
B. sanctaecrucis. References to Bolivian specimens of B. jararacussu are based on misidentifications of B. sanctaecrucis. Although the known
ranges of B. brazili, B. jararacussu, and Bothrocophias hyoprora approach Bolivia’s borders, these species have not yet been collected within
the country. A rectangular loreal is rare in pitvipers and may be a synapomorphy of two small Andean pitvipers: B. andianus and B. lojanus.

KEY WORDS:—Bolivia, Bothriopsis, Bothrocophias, Bothrops, Crotalus, Lachesis, systematics

INTRODUCTION

Bolivia’s venomous snake fauna was last reviewed by
Campbell and Lamar (1989) in The Venomous Reptiles of
Latin America. Campbell and Lamar’s seminal and thor-
ough review of the literature stimulated a prolific era of
research into the biology of Latin American pitvipers.
Their maps underscore our limited knowledge about
pitviper distribution in Bolivia: question marks appear on
maps for Bothriopsis oligolepis, B. peruviana, Bothrops
andianus, B. atrox, B. jararacussu, B. microphthalmus,
and B. neuwiedi. The distribution of Bothrops sanctaecru-
cis is represented by only two localities, and some species
(Bothriopsis bilineata and Lachesis muta) are restricted to
Pando and northeastern La Paz when they actually have
much wider distributions within the country. In anticipa-
tion of a revised edition of Campbell and Lamar’s book,
we began a review of the venomous snakes of Bolivia
(Harvey et al. 2003).
Limited advances in filling gaps identified by the 1989
book have already been made. A new species, Bothrops
jonathani, was discovered in high elevation, xeric condi-
tions in Cochabamba and Santa Cruz (Harvey 1994;
Dirksen et al. 1995). Montero et al. (1995) reported
B. neuwiedi from Chuquisaca and Lachesis muta from
Cochabamba. Visinoni (1995) provided numerous new
localities for Lachesis muta, effectively expanding the
range of this species across the Bolivian lowlands north of
the Gran Chaco. Harvey (1998) documented the presence
of Bothrops moojeni in Bolivia based on specimens from
the Serranía de Huanchaca.
These recent publications reflect renewed interest in
Bolivian herpetology and the growing number of herpetol-
ogists who are actively building museum collections from
this country. We draw heavily on this new material in our
review of Bolivia’s pitvipers.
MATERIALS AND METHODS
For this study, we emphasized characters used in recent
phylogenetic and taxonomic reviews of Neotropical
Viperidae. We use the scale terminology of Klauber (1972)
and characters defined by Werman (1992), Gutberlet
(1998), and Gutberlet and Harvey (2002). We counted the
number of prefoveals defined as all small scales between
the lacunals, postnasal, loreal, and supralabials (Fig. 1).
The species we examined have four types of scales associ-
ated with the pit: pre-, sub-, and supralacunals, and a scale
usually visible as a triangular projection medial to the sub-
lacunal. This last scale occasionally separates the sublacu-
nal from the prelacunal. We noted whether the prelacunal
was partially or completely fused to the second supralabi-
al (=lacunolabial), whether it contacted the second supra-
labial, or whether these scales were separated by a row of
subfoveals.
We recognize several character states for the condition
of the sublacunal and its position relative to the prelacunal
and third supralabial. Werman (1992), Gutberlet (1998),
and Gutberlet and Harvey (2002) recognized that the sub-

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 2 ANNALS OF CARNEGIE MUSEUM
lacunal may be divided into anterior and posterior compo-
nents in some taxa and be polymorphic in others such
as Bothrocophias microphthalmus and B. hyoprora.
Gutberlet and Harvey (2002) found that the scale may also
have interior and exterior components. Data were taken for
many specimens before variation in this character was
understood, and we did not score interior/exterior division
for all specimens. Subfoveals may form a complete (none
of the lacunals contact the supralabials) or incomplete (the
sublacunal does not contact the supralabials; however, the
prelacunal does) row. Presence of one or two subfoveals at
the juncture of the sublacunal, prelacunal, and second and
third supralabials appears to have diagnostic value among
Bothrops and Bothriopsis (Table 1).
The loreal scale contacts the postnasal, canthal, upper
preocular, and supralacunal. We recorded its overall shape,
whether this scale was divided, and if it contacted one or
more canthal scales. In describing Bothrops andianus,
Amaral (1923) noted that the species possesses a rectangu-
lar loreal, and this useful diagnostic trait is relatively rare
in South American pitvipers (Table 1). Most species of
Bothrops and Bothriopsis have loreals that are 80–150% as
long as wide and subtriangular rather than rectangular.
In Bothriopsis, Bothrocophias, Bothrops, and
Lachesis, we recognize three preocular scales, although
the lowest preocular is always smallest and frequently
excluded from the orbit by an elongate subocular. For each
specimen, we looked for fusion of the middle preocular to
the supralacunal and scored whether the upper preocular is
divided and whether it contributes to the canthus. Counts
of interoculabials are the fewest number of scales between
the eye and mouth and include a subocular and a supralabi-
al (i.e., Klauber’s, 1972, use of the term).
On the dorsal surface of the head, we scored presence
or absence of canthorostrals. These tiny scales are posi-
tioned between the rostral and internasal and have
only recently been identified as a synapomorphy of
Bothrocophias hyoprora and B. microphthalmus (Gutber-
let and Campbell 2001; Gutberlet and Harvey 2002). In
pitvipers, the internasals are in contact medially, or one or
more small scales separate them (Fig. 2A). This character
cannot be used to distinguish Bothrops jararaca and
B. jararacussu as previously thought (Campbell and
Lamar 1989). Although across the range of both species
the scale is absent, this trait is nonetheless useful in iden-
tifying other species (Table 1). We found that width of the
canthals and number of scales between the canthals was
particularly helpful in identifying some species such as
B. sanctaecrucis (Fig. 2, Table 1). Because they have been
frequently reported in the past, intersupraoculars were
counted; however, ranges for most species overlap consid-
erably, and we found this character to be of little use in
identifying Bolivian species. We also scored the presence
of intersupraocular fusions. In most Bothrops, the inter-
supraoculars are keeled. These scales may be fused to
form “doubled” scales that are also keeled. Less frequent-
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VOL. 74
ly, several intersupraoculars may fuse to produce large, flat
plate-like scales. We counted interrictals (the number of
scales in a straight line between the corners of the mouth
and including the last supralabials) and compared the
counts to counts at midbody.
We counted the number of infralabials contacting the
chinshields and the number of gulars in a straight line from
the chinshields to the first preventral. Klauber (1972) used
the term “intergenials” to refer to the condition we found
in some Crotalus durissus where the first pair of infralabi-
als are divided to form a pair of scales between the chin-
shields and mental. Our definitions of preventrals and ven-
trals are consistent with the terminology of Dowling
(1951). A pair of postanal scales was included in subcau-
dal counts if the two sides were in contact or fused medi-
ally. For all specimens, we noted the number and location
of entire and divided subcaudals.
We measured snout-vent length and tail length with a
string and meter stick to the nearest 1 mm. Dial calipers or
an ocular micrometer were used to measure head length
(from the posterior margin of the last supralabial to the
center of the rostral), head width (at the level of the rictus),
greatest eye diameter, eye-nostril distance (from the ante-
rior corner of the eye to the center of the nostril), greatest
loreal length and height, and rostral height and width
(width measured at its ventral margin).
Maslin (1942) thought that the nasal pore was absent in
Bothrops sensu lato (“lateralis, schlegelii, atrox, num-
mifera, godmani, aurifera, and nigricauda”) except for
castelnaudi but present in the Asian genus Trimeresurus.
We scored presence or absence of the nasal pore and con-
firmed the observations of Gutberlet and Harvey (2002)
that this trait is present in all Bothriopsis, Bothrops,
Lachesis, and Bothrocophias. It is absent in Crotalus
durissus.
In the accounts, a dash is used for ranges, whereas a
slash (/) is used to designate counts from opposite sides of
the same specimen. Wherever ranges appear, they are fol-
lowed by means ± standard deviation. Specimens were fre-
quently damaged, and we follow each range or character
frequency with the sample size. Hemipenial descriptions
employ the terminology of Myers and Campbell (1981).
Except for the Colección Boliviana de Fauna (CBF, La
Paz, Bolivia), the Centro de Biodiversidad y Genética
(CBG, Cochabamba, Bolivia), the Museo de Historia
Natural “Noel Kempff Mercado” (NK, Santa Cruz,
Bolivia), and the Estación Biológica Doñana (EBD,
Sevilla, Spain), institutional abbreviations are those in
Leviton et al. (1985). We do not supply extensive syn-
onymies of most wide-ranging pitvipers occurring in
Bolivia: Bothriopsis bilineata, B. taeniata, Bothrops atrox,
B. neuwiedi, Crotalus durissus, and Lachesis muta (inter-
ested readers are referred to McDiarmid et al. 1999 for
recent synonymies of these species). Synonymies for the
remaining species include all known synonyms and refer-
ences consulted during this study. Where identifications
 2005 HARVEY ET AL.—BOLIVIAN PITVIPERS
may have been in doubt, we examined specimens on
which the citations are based. In the interest of space, the
synonymies are arranged according to chronology of
names rather than chronology of bibliographic references.
KEY TO BOLIVIAN PITVIPERS
1. Dorsal pattern of diamond-shaped marks; tail short, 7–12% as
long as snout-vent length . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1'. Dorsal pattern uniform, with bands, or with X-shaped marks, but
never with large middorsal diamond-shaped marks; tail more
than 12% as long as snout-vent length . . . . . . . . . . . . . . . . . 3
2(1). Tail ending in a rattle or button, two preoculars, nasal pore
absent, subcaudals entire . . . . . . . . . . . . . . . Crotalus durissus
2'. Tail not ending in rattle, distal caudals extensively divided, three
preoculars, nasal pore present, subcaudals mostly divided . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachesis muta
3(1'). Dorsal keels tuberculate on posterior part of body, infralabials
and gulars brown with cream round or C-shaped marks, ventrals
fewer than 160, canthorostrals present or absent . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . .Bothrocophias microphthalmus
3' Dorsal keels well-developed but not tuberculate, infralabials and
gulars variously patterned or immaculate but not brown with
well-defined round or C-shaped marks, ventrals more than 160,
canthorostrals absent except as a very rare anomoly . . . . . . . 4
4(3'). Dorsum bright green; tail prehensile; loreal subtriangular . . 5
4'. Dorsum olive green, brown, or gray but if green then loreal
elongate and subrectangular; tail not prehensile . . . . . . . . . . 7
5(4). Dorsum banded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5'. Dorsum uniform green with black specks on each scale . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bothriopsis bilineata
6(5). Dorsal pattern of short black and yellow bands on an immacu-
late green background; venter immaculate; subcaudals divided
with a few entire subcaudals located proximally in some speci-
mens . . . . . . . . . . . . . . . . . . . . . . . . . . . Bothriopsis oligolepis
6'. Dorsum lichenose, interspaces heavily stippled in yellow and
black; venter heavily pigmented and maroon in color posterior-
ly; subcaudals mostly entire (a few divided subcaudals usually
present distally, rarely present proximally) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Bothriopsis taeniata
7(4). Second supralabial usually not fused to prelacunal . . . . . . . . 8
Fig. 1.—Left: Facial scalation of Bothrops neuwiedi (UTA 39112) showing three rows of interoculabials (gray). Right: Specimen of Crotalus durissus
(UTA 38042) with a divided loreal (gray) and with the upper loreal partially fused to the canthal.
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3
7'. Second supralabial usually fused to prelacunal . . . . . . . . . . . 9
8(7). Postocular stripe high (covering about 4.5 temporals at its high-
est point); dorsals 27–33; supralabials 9–12; infralabials 13–16;
interoculabials 4 . . . . . . . . . . . . . . . . . . . . Bothrops jonathani
8'. Postocular stripe moderate (1.5–3 temporals at its highest
point); dorsals 23–27; supralabials 8–10; infralabials 10–13;
interoculabials usually 3 . . . . . . . . . . . . . . Bothrops neuwiedi
9(7'). Loreal subrectangular and elongate; dorsal blotches becoming
distinctive row of 2–4 black spots on posterior body and tail;
ventrals 166–179; internasals usually (91%) separated by 1–3
small scales . . . . . . . . . . . . . . . . . . . . . . . . Bothrops andianus
9'. Loreal subtriangular; dorsal pattern not modified in front of
vent; ventrals usually 175 or more; internasals in contact or sep-
arated by 1–3 small scales at low frequencies . . . . . . . . . . . 10
10(9'). Canthals wide, separated by 0–2 (rarely 3) keeled scales; iris
pale green; distal one-half of tail black (in adults), sharply con-
trasting with pale dorsal pattern . . . . . Bothrops sanctaecrucis
10'. Canthals narrow, separated by three or more keeled scales; iris
bronze; distal one-half of tail heavily pigmented but same color
as dorsal blotches and not sharply contrasting with dorsal
pattern . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11(10'). Postocular stripe two or more temporals high and well defined;
venter heavily pigmented, uniformly stippled or with alternating
charcoal and cream squares . . . . . . . . . . . . . . . Bothrops atrox
11'. Postocular stripe narrow (less than one temporal high); venter
immaculate or nearly so . . . . . . . . . . . . . . . Bothrops moojeni
SYSTEMATIC ZOOLOGY
Class Reptilia Laurenti, 1768
Order Squamata Merrem, 1820
Suborder Serpentes Linnaeus, 1766
Family Viperidae Oppel, 1811
Bothriopsis Peters, 1861
Remarks.—In recent years, the systematics of Neotropical
pitvipers has been in a state of flux. The arboreal species
referred to the genus Bothriopsis (Campbell and Lamar
1989; McDiarmid et al. 1999) consistently cluster with
 4 ANNALS OF CARNEGIE MUSEUM
Fig. 2.—Intercanthal scales (gray) of (left) Bothrops sanctaecrucis (MCZ 17699, also with small scale separating internasals) and (right) B. atrox
(LSUMZ 45975).
species of Bothrops in phylogenetic analyses using mito-
chondrial DNA (Kraus et al. 1996; Parkinson et al. 2002;
Salomão et al. 1999; Wüster et al. 2002), allozymes
(Werman 1992), and combined molecular and morpholog-
ical characters (Werman 1992, 1999). Usually, Bothriopsis
species are nested within a clade containing species of
Bothrops with a lacunolabial. Recognition of Bothriopsis
renders Bothrops paraphyletic, and Bothriopsis as current-
ly defined (i.e., including Bothriopsis punctata,
McDiarmid et al. 1999) may not be monophyletic.
However, we follow other recent revisors (Campbell and
Lamar 1989; McDiarmid et al. 1999) and recognize the
genus with the anticipation that Bothrops will be further
partitioned. Nonetheless, we note that not only is
Bothriopsis nested within Bothrops, but it is nested within
the B. atrox group. Resolving the problem of paraphyly is
not a simple matter of recognizing three genera, one each
for the terrestrial species with lacunolabials, the species
lacking a lacunolabial, and the arboreal species.
Our study is based almost exclusively on external char-
acters, and we can contribute little to this debate. Green col-
oration was thought to be a synapomorphy of Bothriopsis;
however, we found that some specimens of B. andianus
from Santa Cruz and Cochabamba and all specimens from
La Paz are green. When examining comparative material of
the terrestrial species Bothrops osbornei, we noted that this
species is yellow as a juvenile and green as an adult. Other
than a prehensile tail, we know of no other unambiguous
morphological (molecular data are available for only three
of the seven species) synapomorphies of Bothriopsis. For
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VOL. 74
this reason, and because the genus requires further revision,
we will not provide a definition of the genus here.
Content.—Seven species restricted to South America:
Bothriopsis bilineata (Wied-Neuwied); B. chloromelas
(Boulenger); B. medusa (Sternfeld); B. oligolepis (Werner);
B. pulchra (Peters); B. punctata (García); B. taeniata Wagler.
Bothriopsis bilineata smaragdina Hoge, 1966
Figure 3
Bothrops bilineatus smaragdinus Hoge, 1966:114. HOLOTYPE female
from “upper Purus river, State Amazonas, Brasil” (IBH 14731)
Diagnosis.—Bothriopsis bilineata smaragdina is distin-
guished from all congeners and from species of
Bothrocophias and Bothrops by the following combina-
tion of characters: (1) lacunolabial present; (2) dorsal
keels nontuberculate; (3) infralabials and gulars pig-
mented to varying degrees but lacking regular series of
cream round or C-shaped marks; (4) canthorostrals
absent; (5) most subcaudals divided; (6) ventrals
192–220 and subcaudals 55–76 (Campbell and Lamar
1989); (7) loreal subtriangular, higher than long; (8) can-
thals broad and separated by 2–5 scales; (9) internasals
in contact; (10) postocular stripe poorly defined or
absent; (11) dorsum pastel green, finely peppered with
black; dorsal bands absent; broken yellow paraventral
stripe extending length of body.
 2005 HARVEY ET AL.—BOLIVIAN PITVIPERS
Fig. 3.—Top, Juvenile Bothrops atrox (UTA 38035) from near Yolosa in
the Andes of La Paz. Bottom, Bothriopsis bilineata from Parque
Nacional Amboró (NK 328, Photo by I. De la Riva).
Description.—The largest Bolivian specimen (NK 328)
has a snout-vent length of 812 mm and tail length of 136
mm. The tail is 16–19% (17.0 ± 0.9, 4) of snout-vent
length in females and 16% of snout-vent length in a single
juvenile specimen. In the adults, the head accounts for
3.6–4.2% (4.1 ± 0.5, 3) of snout-vent length, and eye-nos-
tril distance is 24.3–25.8% (24.6 ± 1.1, 3) of head length.
Bothriopsis bilineata lacks canthorostrals and scales
between the internasals. The canthals are broad and sepa-
rated from one another by 2–3 keeled scales in Bolivian
specimens (2–5 scales elsewhere in South America). Part
of the upper preocular and the internasal and canthal form
the canthus. The lateral margins of the last two scales are
thickened and upturned. Five to eight (6.8 ± 1.3, 5) keeled
scales separate the supraoculars. A “doubled” inter-
supraocular is present in one specimen. The surfaces of the
internasals, canthals, and supraoculars are rugose.
Interrictal counts (26–30, 28 ± 2, 5) were one to three
scales more than number of dorsals at midbody.
In our sample, the rostral is slightly higher than wide;
four or five prefoveals are present. The second supralabial
and prelacunal are discrete on one side of AMNH 6773,
and only a small part of the prelacunal and second supral-
abial are fused in NK 328. The loreal is single, subtriangu-
lar, and about as long as high. It broadly contacts the can-
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5
thal and supralacunal. The middle preocular is partially
fused to the supralacunal on both sides of the paratypes.
Neither the upper preocular nor the sublacunal is divided.
Subfoveals are absent. An elongate subocular and two pos-
toculars are present in all but one specimen; postoculars
are absent in NK 452, apparently having fused to the sub-
ocular. A lower preocular is absent in FMNH 161512 and
appears to have fused to the sublacunal on both sides. In
two specimens (40%, 5), the subocular contacts the third
supralabial, whereas the subocular and supralabials are
separated by a row of keeled scales (=3 interoculabials) in
the remaining specimens.
Bolivian specimens have seven or eight supralabials
and 10–12 infralabials. Number of infralabials contacting
the chinshields is more variable in this species than in
other South American pitvipers. In the Bolivian sample,
the first three (50%, 5), first two (30%, 5), first four (10%,
5) or first five (10%, 5) infralabials contact the chinshields.
Four or five gulars and one to three preventrals are pres-
ent. Female specimens have 192–198 (195.8 ± 2.6, 4) ven-
trals and 58–68 (63.8 ± 4.6, 4) subcaudals; the juvenile has
198 ventrals and 65 subcaudals. Twenty-five to 29 (27.0 ±
1.6, 5) dorsals at midbody reduce to 17–21 (19.4 ± 1.7, 5)
dorsals one head-length in front of the vent.
In adults of our sample, each dorsal body and head scale
is green with many small black spots. Spotting extends to
the supralabials and infralabials as well. The gulars are
mostly cream, but some have a few black spots. In the juve-
nile paratype (FMNH 161512), black spotting is reduced on
the flanks and concentrated anteriorly on the body. The pos-
terior one-third of the dorsum and the tail are uniform green
without spots. A few supralabials and infralabials of the
juvenile have scattered black flecks; its gulars and chin-
shields are immaculate. In adults, the yellow paraventral
stripe appears broken, because each paraventral is yellow
distally and green proximally. In the juvenile, each par-
aventral is entirely yellow and the paraventral stripe is
unbroken. Medially the ventrals are cream; however, green
stripes on the edges of the ventrals are slightly narrower
than and border the yellow paraventral stripes. The ventral
surface of the tail is cream; dorsally, its distal one-fourth is
pale pink or red. In the juvenile, the distal one-fourth of the
tail is marked by a series of eight short brown bands.
Distribution and Comparative Material.—BOLIVIA.
COCHABAMBA. Chapare: “Alto Chipiriri” (FMNH
161512, Paratype of Bothrops bilineatus smaragdinus).
Unknown: (AMNH 6773, Paratype of Bothrops bilineatus
smaragdinus, specific locality unknown). PANDO.
Nicolás Suárez: canton Mukden (NK live collection).
SANTA CRUZ. Ichilo: Parque Nacional Amboró (NK
328 and NK 1429). Unknown: NK 452 (specific locality
unknown).
ECUADOR. MORONA-SANTIAGO. USNM
165278. NAPO. USNM 165279-80. PASTAZA. USNM
165262-77.
 6 ANNALS OF CARNEGIE MUSEUM
Specimens not examined: BOLIVIA. COCHABAM-
BA. Chapare: “Villa Tunari” (EBD, no museum number
given, Fugler and De la Riva, 1990). Based on Miranda et
al. (1991), Middendorf and Reynolds (2000) report this
species from the Beni Biological Station Biosphere
Reserve (Gral. José Ballivián and Yacuma provinces,
14°40'S, 66°30'W, 220 m) (Fig. 8).
Bothriopsis oligolepis (Werner, 1901)
Figure 4
Lachesis bilineatus var. oligolepis Werner, 1901:13. HOLOTYPE male
from “Bolivien” (=Bolivia) (MTKD D 1714) [destroyed or lost,
Obst, 1977]. Amaral 1930a:58.
Lachesis peruvianus Boulenger, 1903:354. HOLOTYPE female from
“La Oroya, Carabaya, S.E. Peru” (BMNH 1946.1.19.27, formerly
BMNH 1903.6.30.6) [examined]. Griffin, 1916:226. [Carrillo de
Espinoza (1983, her note p.18) thought that La Oroya, a locality in
Junin, might be in error and suspected that the holotype came from
Carabaya, Puno].
Bothrops peruviana (Boulenger): Amaral, 1930b:240; Pifano and Römer,
1949:294.
Bothrops peruvianus (Boulenger): Hoge, 1966:131; Peters and Orejas-
Miranda, 1970:53; Hoge and Romano-Hoge, 1981:217; Carrillo de
Espinoza, 1983:18 (part, also includes B. chloromelas); David and
Ineich, 1999:233 (part, also includes B. chloromelas).
Bothrops oligolepis (Werner): Peters and Orejas-Miranda, 1970:53 (part);
Hoge and Romano-Hoge, 1981:217 (part); Carrillo de Espinoza
1983:17; Fugler, 1986:48; David and Ineich, 1999:233 (part).
Bothriopsis peruviana (Boulenger): Campbell and Lamar, 1989:175;
Carrillo de Espinoza and Icochea, 1995:20; McDiarmid et al.,
1999:249
Bothriopsis oligolepis (Werner): Campbell and Lamar, 1989:174 (part);
Fugler and De la Riva, 1990:37; Carillo de Espinoza and Icochea,
1995:20 (part); Fugler et al., 1995:58; McDiarmid et al., 1999:249.
Bothriechis oligolepis (Werner): Schätti and Kramer, 1993:244 (part).
Bothriechis oligolepis oligolepis (Werner): Golay et al., 1993:37 (part).
Diagnosis.—Bothriopsis oligolepis is distinguished from
all congeners and from species of Bothrocophias and
Fig. 4.—Female holotype of Lachesis peruvianus Boulenger
(=Bothriopsis oligolepsis, BMNH 1945.1.19.27, SVL 465 mm) from the
Cordillera de Carabaya, Peru.
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VOL. 74
Bothrops by the following combination of characters: (1)
lacunolabial present; (2) dorsal keels nontuberculate; (3)
infralabials and gulars immaculate in adults; (4) can-
thorostrals absent; (5) subcaudals divided or with few
entire subcaudals anteriorly; (6) ventrals 189–193 and sub-
caudals 54–65 (Campbell and Lamar, 1989, report
188–196 ventrals and 53–66 subcaudals for this species);
(7) loreal subtriangular; (8) canthals narrow; (9) inter-
nasals in contact; (10) postocular stripe black and 2–3 tem-
porals high; (11) dorsum bright green with black and yel-
low bands, each about one dorsal long; dorsal surface of
head green with few black and yellow blotches.
Description.—The holotype of Lachesis peruvianus is a
male with a total length of 547 mm; two additional speci-
mens are females 940 and 714 mm long. The tail length is
15–18% (17 ± 1, 3) of snout-vent length. This species has
a triangular head, strongly distinct from its neck. The head
is longer than it is wide, its width being 42–79% (59 ± 19,
3) of its length. Dorsal head scales are keeled; a few scales
in the parietal region clearly result from fusion of two
scales and have doubled keels; flat, plate-like scales are
absent from the intersupraocular and parietal regions. The
dorsal surfaces of the internasals, canthals, upper preocu-
lars, and supraoculars are rugose. The supraoculars are
70% as wide as long, roughly subtriangular, and separated
medially by 6–8 (7 ± 1, 3) scales. Twenty-eight to thirty-
one (29 ± 2, 3) interrictals are present.
Thickened, central portions of the internasal, canthal,
and upper preocular form a sharp and raised canthus ros-
tralis. The rostral is about as high as wide and about as
wide as the mental. The loreal is entire and as long as or
slightly longer than high (loreal height is 86–100% of its
length, 92 ± 7, 3); it broadly contacts the canthal and pre-
ocular. The foveal pit is bound by entire upper and lower
lacunals and the fused second supralabial and prelacunal.
The specimens have 2–3 prefoveals and lack canthoros-
trals. The eye is large in the relatively small holotype
where its diameter is 85% of the distance between the
anterior border of the orbit and the center of the naris. The
orbit is bound by 1/1 supraoculars, 2/2 preoculars, 1–2
elongate suboculars, and 2/2 postoculars. An elongate,
jagged subocular excludes a small, rectangular lower pre-
ocular from the orbit in all specimens. Each of the preocu-
lars is entire; the upper is subtriangular and nearly four
times as large as the nearly square middle preocular. This
species has 7–8 supralabials most of which are subtriangu-
lar and high. Supralabials 3 and 4 contact the subocular on
both sides of the holotype (=2/2 interoculabials) and on
one side only of each of the other two specimens. When
not contacting the supralabials, the subocular is separated
by a single row of keeled scales. The foveal pit lies below
an imaginary line drawn between the eye and naris. A
nasal pore is present.
This species has 9–10 infralabials. The chinshields are
about twice as long as wide, contact the first three infral-
 2005 HARVEY ET AL.—BOLIVIAN PITVIPERS 7

abials, and are separated from the mental by medial con-

tact of the first pair of infralabials. The mental groove is
shallow. Four to six gulars separate the chinshields from
the preventrals and ventrals. The paraventrals and parasub-
caudals are keeled. This species has 23 reducing to 19 dor-
sals, 189–193 (191 ± 2, 3) ventrals and preventrals, and
54–65 (60 ± 6, 3) subcaudals. Subcaudals 2–8 are entire in
the holotype; all are divided in the other specimens.
In spite of its age, the coloration of the holotype does
not appear to have faded substantially. Its dorsum is blue-
green with a black and yellow pattern. Ventrally the spec-
imen is yellow with black and blue-green edging on most
ventrals, especially posteriorly. A small black blotch over-
laps the medial contact between the internasals and the
first couple of keeled dorsal scales just behind these inter-
nasals. A prominent, black V-shaped mark begins as a bar
extending between the anterior edge of the supraoculars;
the mark continues as wide black stripes edging the
supraoculars and paralleling the postocular stripe onto the
neck. The dorsal surface of the scales making up the can-
thus and the supraoculars is largely immaculate except for
yellow blotches associated with surface rugosities. A pair
of black, curved bands is edged in yellow medially and
overlaps the neck and some parietal scales between the
“arms” of the cephalic V-shaped mark.
Scales of the face are blue-green with yellow edges.
Some scales have scant and diffuse black pigment. The
anterior supralabials are yellow with blue-green and Fig. 5.—Female specimen of Bothriopsis pulchra (ZFMK 41480) from
black edges. The postocular stripe covers part of the sub- Andoas, Peru.
ocular, 2–3 rows of temporals, the upper one-fourth of
supralabial 5/6, and more than half of supralabials
6–7/7–8. Scales of the chin are immaculate yellow except ing can be discerned on the posterior third of their bodies.
for the last infralabials, which are overlapped by the pos-
tocular stripes. Specimens Examined and Distribution.—PERU. JUNIN.
Well-defined black and yellow bands, one scale long “Chanchamayo: La Merced region” (MCZ 45906. FMNH
cover the dorsal body. The bands are separated from the 68597).
paraventrals by a single row of green dorsals. Yellow PUNO. “La Oroya, Carabaya, S.E. Peru” (=Cordillera
bands are adjacent to each other: the sequence of bands de Carabaya) (BMNH 1946.1.19.27, formerly
is yellow–black, black–yellow, yellow–black, etc. 1903.6.30.6, holotype of Lachesis peruvianus).
Paraventral spots are yellow and occupy one and one-half
to three paraventrals. These spots are separated by 1–1.5 Remarks.—Werner (1901) described a green snake with
black and green scales. The dorsal pattern is largely con- black and yellow bands from Bolivia. In addition to the
sistent except on the anterior part of the body where the presence of green pigmentation, it possessed other charac-
bands are broken and irregular. Notably, well defined teristics of the forest pitvipers: few interoculars (6) and
bands extend to the vent. The dorsal and paraventral pat- labials (7/8 supralabials, 10/11 infralabials). He thought
terns on the tail are quite different from that of the dorsal his specimen was a color variant of Bothriopsis bilineata
body. Most notably, parasubcaudal yellow and black and proposed the new name oligolepis for it.
stripes extend for three quarters of the length of the tail. Later, Werner (1922) believed that oligolepis was syn-
The distal one-fourth of the tail is black both ventrally and onymous with Boulenger’s Lachesis chloromelas and sug-
dorsally. Most of the tail is blue-green dorsally, although gested that oligolepis be treated as a junior synonym of
four yellow bands are present on its proximal one-fourth. that species. However, Parker (1934) noted that Werner’s
The buccal epithelium is unpigmented; the tongue was name predated that of Boulenger and most revisors have
retracted, and its color could not be ascertained without followed Parker’s view that oligolepis is the oldest name
dissection. The other two specimens are in poorer condi- applicable to the intricately patterned species currently
tion and their coloration has faded (MCZ 45906) or black- represented by several specimens from northern and cen-
ened (FMNH 68597) with age. Nonetheless, yellow band- tral Peru. When redescribing Bothriopsis medusa, Pifano

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 8 ANNALS OF CARNEGIE MUSEUM
and Römer (1949) provided a Spanish translation of
Boulenger’s description and used chloromelas to refer to
the Peruvian species, however these authors appear to
have been unaware of Parker’s synonymization of
Werner’s and Boulenger’s names.
Other than Werner, no revisor of this species has ever
examined the holotype of oligolepis. Werner’s specimen
differed morphologically from all known specimens of
chloromelas. Werner described the head of the holotype as
being green with a few black spots; whereas, the head of
chloromelas is distinctly black and heavily mottled with
yellow and green. He reports that all of the scales on the
top of the head of his type specimen were strongly keeled.
Usually (MCZ 156347 is an exception), specimens of
chloromelas possess one or more flat, plate-like scales,
especially in the interocular and parietal regions. Werner
describes the venter as being light green; the venter of
chloromelas is green but heavily speckled from the chin
throughout the body and tail. All of the subcaudals were
divided in Werner’s specimen; this is true for two of the
three specimens of Bothriopsis oligolepis but not for any
B. chloromelas that we examined. Finally, there is no men-
tion of the distinctive lichenose dorsal pattern of
B. chloromelas; Werner only mentions yellow and black
bands, the same pattern found in the holotype of
L. peruvianus.
One might wonder why in 1922 Werner would suggest
that chloromelas and oligolepis are synonymous rather
than peruvianus and oligolepis since both chloromelas in
1912 and peruvianus in 1903 had already been named. In
fact, Werner confused Boulenger’s species. In a later pub-
lication, he (Werner 1927) referred a specimen from
“Amable Maria, Peru” to peruvianus when the specimen
could only have been chloromelas. Werner mentions the
black head, distinctive yellow lines on the anterior head
shields, heavily punctate labial scales, and heavily speck-
led venter characteristic of all specimens of chloromelas
[with 194 ventrals and 60 subcaudals, the specimen was
probably not B. taeniatus, which has 203–254 ventrals and
66–91 subcaudals (Campbell and Lamar 1989)].
To our knowledge, no specimens of Bothriopsis
chloromelas are known from south of Pasco. Although the
possibility that B. chloromelas occurs in Bolivia should
not be completely discounted, we think it unlikely that
even this cryptic species should evade detection for so
long. Werner’s type of oligolepis and the holotype of
Lachesis peruvianus are the same species. Accordingly,
we relegate Lachesis peruvianus Boulenger to the syn-
onymy of Bothriopsis oligolepis and revalidate
Bothriopsis chloromelas (Boulenger).
Campbell and Lamar (1989, fig. 155 and 156) provid-
ed two color photos of specimens they referred to
Bothriopsis peruviana. We are not sure if these specimens
are Bothriopsis oligolepis. In both specimens, the yellow
pigment appears to be replaced by white. The iris is silvery
green.
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VOL. 74
Henle and Ehrle (1991) reported a specimen of
Bothrops peruvianus from Andoas, a town on the Río
Pastaza near the Ecuadorian border. In all substantive
characteristics the female specimen (ZFMK 41480, Fig. 5)
agrees with specimens of B. pulchra examined by us
(Appendix). Since descriptions of Peruvian B. pulchra are
unavailable, we provide a brief account of this rare and
poorly known species. The facial scales are dessicated and
scales in this area could not be counted with accuracy. On
each side, an elongate subocular contacts the supralabials
and is separated from the entire supraocular by 2/4 postoc-
ulars. The internasal, canthal, and upper preocular form a
thickened (“raised”) canthal border. Behind the inter-
nasals, intercanthals increase from three to four to six;
there are five intersupraoculars and 26 interrictals. Three
of the intersupraoculars are fused pairs of scales. The spec-
imen has 7/7 supralabials and 10/10 infralabials with
infralabials 1–3/1–4 contacting the first pair of chin-
shields. Four pairs of scales border the mental groove and
are separated from the preventrals by two gulars. The
specimen has 20 + 23 + 17 dorsals, four preventrals, 175
ventrals, and 61 subcaudals. The anal and subcaudals
1–41, 44–45, and 47–48 are entire.
The intercanthals, intersupraoculars, and dorsals are
heavily keeled, and the supraoculars are weakly rugose.
The paraventrals are nearly smooth anteriorly with weak
keels nearing the vent. The parasubcaudals are keeled.
In preservative, the dorsum is blue-green with black
bands so diffuse as to be barely discernible. All dorsal
body and head scales have white keels. Terra cotta colored
bands are restricted to the posterior half of the specimen:
16 on the body, five on the tail. The tip of the tail is yellow
for about 27 subcaudals. A black postocular stripe occu-
pies the upper one-third of supralabials 6–7 and 1.5 rows
of temporals. The supralabials are grayish cream with
large black spots overlapping their junctures. The chin is
white except for black pigment along the edges of all
infralabials. The specimen has a prominent row of para-
ventral spots, each occupying about 1–3 paraventrals and
the upper portions of adjacent ventrals. The spots are sep-
arated from one another by single paraventrals, and each is
bound ventrally by a thick, gray to blue-green band.
Midventrally, the venter is white with few small gray
blotches.
Measurements (in mm): SVL 346, tail-length 76,
head length 16.08, head width 10.25, eye-diameter
3.25.
This species was once thought to differ from other
forest pitvipers by having fewer dorsals than all species
except Bothriopsis medusa (19–21 vs. 23–35, Campbell
and Lamar 1989). However, specimens with 23 dorsals
are not uncommon and account for 30% (n = 10) of the
specimens we examined (also see Campbell and Lamar
1992). The species is most readily distinguished from
all other forest pitvipers by the white keels, terra cotta
bands on the posterior body, and distinct labial pattern.
 2005 HARVEY ET AL.—BOLIVIAN PITVIPERS
Referred to in the past by its junior synonyms
Bothriopsis albocarinatus and B. alticola, the confused
taxonomic history of this species was recently reviewed by
Kuch (1997), Gutberlet and Harvey (1998), and Wüster
(1998). The International Commission of Zoological
Nomenclature, opinion 1939, placed the specific name
pulcher on the official list as defined by the holotype
Trigonocephalus pulcher Peters.
Bothriopsis taeniata (Wagler, 1824)
Bothrops taeniatus Wagler, 1824:55. HOLOTYPE (lost or destroyed)
from “ad flumen Amazonum” (restricted by Vanzolini 1981, to the
stretch of river between the mouths of the Tajapuru and Negro
rivers). NEOTYPE (MNHN 1582, also type of Bothrops castelnaudi
Duméril, Bibron, and Duméril) [Designated by Hoogmoed and
Gruber 1983:337].
Diagnosis.—Bothriopsis taeniata is distinguished from all
congeners and from species of Bothrocophias and
Bothrops by the following combination of characters: (1)
lacunolabial present; (2) dorsal keels nontuberculate; (3)
infralabials and gulars mottled; (4) canthorostrals absent;
(5) subcaudals mostly entire (several divided subcaudals
present on distal one-third of tail in 70% of specimens we
examined; few divided proximal subcaudals present in
20%); (6) ventrals 203–254 and subcaudals 66–91
(Campbell and Lamar 1989); (7) loreal subtriangular; (8)
canthals moderate, separated by 2–4 scales in our sample;
(9) internasals in contact or separated by a single scale;
(10) postocular stripe black, about two temporals high, and
often stippled and obscure; (11) dorsum banded and with a
lichenose pattern of green, black, and yellow; prominent
row of yellow paraventral spots present; venter heavily
pigmented posteriorly.
Description.—The Bolivian sample consists of a small
female (LSUMZ 45976), 650 mm long, and an unsexed
juvenile (CBF 2304), 375 mm long. In addition, one of us
(LGA) obtained limited data from a captive specimen col-
lected in Pando. The tail is prehensile and 15% as long as
the head and body in the female and 18% as long in the
juvenile. The head accounts for 3–4% of snout-vent length
and is 70–90% as wide as long. The eye is large and
accounts for 18% of head length in the female and 30% in
the juvenile; its distance from the nostril accounts for 27%
of head length in the female and 39% in the juvenile.
The internasals are in contact medially, and canthoros-
trals are absent. The canthus is formed by thickened later-
al margins of the internasal, canthal, and upper preocular.
The canthals are relatively smooth and wide; 2–5 (3 ± 2, 3)
keeled scales separate them. The supraoculars are separat-
ed by 6–7 (7 ± 1, 3) keeled scales, one or two are “dou-
bled” and keeled. Flat platelike intersupraoculars, inter-
canthals, parietals, and occipitals are absent. The rostral is
82.0–89.7 (85.8 ± 5.4, 2) as wide as high. The loreal is
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9
entire and subtriangular; it is 80.0–162.5% (121.0 ± 58.0,
2) as high as long and in broad contact with the canthal.
One to three (3 ± 1, 2) prefoveals are present. The subla-
cunal is distinct from the middle preocular and no scale is
present at the juncture of the sublacunal, lacunolabial, and
third supralabial. A single elongate subocular is separated
from the supraocular by 1–2 postoculars and from the
supralabials by 1/1 keeled scale (=3/3 interoculabials). The
specimens have 7/7 supralabials, 9–12 (11 ± 1) infralabi-
als, and 2–4 infralabials contacting the chinshields. Five
gulars and 1–2 preventrals separate the chinshields from
231 ventrals and 73 subcaudals in the female and 230–233
ventrals and 78–81 subcaudals in the unsexed specimens.
All of the subcaudals are entire in the unsexed specimens,
and most are entire in the female, however the first
subcaudal and ten distal subcaudals are divided in this
specimen.
Every scale on the head bears at least some black
speckling. The dorsal head scales are bluish-green (=green
in life) with numerous black spots. The labials are paler
and have the highest concentrations of black speckling at
their edges giving the impression of poorly defined labial
bands. The postocular stripe is also heavily speckled and
thus obscure against the similar dorsal head patterning. It
is about two temporals in height and extends from the sub-
ocular and postoculars across part of the last three supral-
abials; it does not extend beyond the rictus. The gulars are
only diffusely pigmented and pale yellow. Although all
ventral are heavily pigmented, the pigmentation is most
diffuse anteriorly. Most of the venter is charcoal, and pairs
of circular yellowish spots on each ventral scale form rows
on the lateral margins.
The dorsum is lichenose and bears 27–30 (2) poorly
defined darker bands against a light green background.
Prominent yellow spots form a row along the paraventrals
and parasubcaudals. The spots are relatively large, each
covering the edge of a ventral, 1–3 paraventrals, and 1–2
scales on the second row of dorsals. The spots are separat-
ed by two scales and, in the female, number 69 on the body
and nine on the proximal portion of the tail. The dorsal and
ventral patterns on the proximal two-thirds of the tail are
the same as on the body; however, the last 37–43 (2) sub-
caudals are cream-colored ventrally. Dorsally, the distal
third of the tail has numerous alternating cream and brown
bands.
Distribution in Bolivia and Comparative Material.—
BOLIVIA. LA PAZ. Iturralde: “Camino entre
Tumupasa y San José de Uchupiamonas, Serranía Sadiri;
Area Natural de Manejo Integrado Madidi” (CBF 2304).
PANDO. Nicolás Suárez: “12 km by road S Cobija, ca. 8
km W on road to Mucden” (=Muckden; LSUMZ 45976);
Río Tahuamanu, San Sebastián (NK, live specimen).
ECUADOR: MORONA-SANTIAGO. USNM
283968. PASTAZA. USNM 165239, 165288, 165290–94.
PERU. HUÁNUCO: USNM 167625–26.
 10 ANNALS OF CARNEGIE MUSEUM
Fig. 6.—Top, Juvenile Bothrocophias microphthalmus (NK 1801, SVL
393 mm) from Pilón Lajas, Serranía Beu, La Paz (Photo by D. Rumiz).
Bottom, Bothrops andianus from Parque Nacional Amboró, Santa Cruz.
Remarks.—Fugler and De la Riva (1990) and Fugler et al.
(1995) report both “Bothriopsis taeniata” and “Bothrops
castelnaudi” (Fugler and De la Riva 1990, p. 38) or
“Bothriopsis castelnaudi” (Fugler et al. 1995, p. 58) from
Bolivia. We assume that the authors are referring to the
same species; the names are synonyms (Hoogmoed and
Gruber 1983). Campbell and Lamar (1989) included
Bolivia in the distribution of this species.
Bothrocophias Gutberlet and Campbell, 2001
Bothrocophias contains four species of small to moderate
terrestrial pitvipers with stout bodies and lacking a rattle or
prehensile tail. They share two synapomorphies: tubercu-
lar keels on the posterior dorsal scales and distinctive
white to cream spots on otherwise brownish infralabials
and gulars (Gutberlet and Campbell 2001; Gutberlet and
Harvey 2002). The snout is weakly to moderately elevat-
ed, canthorostrals are present in some species, and the
lacunolabial may be entire or divided. Gutberlet and
Campbell (2001) summarize characteristics that distin-
guish Bothrocophias from all other New World genera.
There are 124–177 ventrals; 38–64 mostly divided subcau-
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VOL. 74
dals (except in B. hyoprora where most or all subcaudals
are entire); 21–25 middorsal scale rows; 2–9 smooth or
keeled intersupraoculars; 7–8 supralabials; 8–11 infralabi-
als; 1–11 prefoveals; 1–3 canthals; 3–4 interoculabials;
4–5 palatine teeth; 12–15 pterygoid teeth; and 14–16 den-
tary teeth. In addition, the maxillary fang is approximate-
ly 1.5 times longer than the height of the maxilla; mesial
spines are present on the hemipenial lobes; there is a mod-
erate number (ca. 18–30 per lobe) of lateral spines on the
hemipenes; the hemipenial lobes are only slightly longer
than the organ’s base; the choanal process of the palatine
is attenuate distally; the ectopterygoid and base of the
pterygoid are approximately equal in length; the dorsal
surface of the frontal bones are predominately flat; and the
postorbital bones are large, contributing more to the dorsal
perimeter of the orbit than does the parietal. Species in this
genus are brown to reddish brown with darker bands.
Content.—Four species restricted to South America:
Bothrocophias campbelli (Freire-Lascano), B. hyoprora
(Amaral), B. microphthalmus (Cope), and B. myersi
Gutberlet and Campbell.
Bothrocophias microphthalmus (Cope, 1875)
Figure 6
Bothrops microphthalmus Cope, 1875 (preprint of 1876 article):182.
HOLOTYPE female from “between Balsa Puerto and Moyabamba,
Peru” (ANSP 11515) [examined]. Cope 1879:277 (tentative,
remarks); Orcés 1943:169; Prado and Hoge 1948:291; Fugler and De
la Riva 1990:38; Kuch and Freire 1995:81; David and Ineich
1999:231; McDiarmid et al. 1999:267.
Lachesis microphthalmus (Cope): Boulenger 1896:540.
Lachesis pleuroxanthus Boulenger, 1912:423. HOLOTYPE female from
“Alpayaca, Río Pastaza, E. Ecuador, 3600 feet” (BMNH
1946.1.19.88) [examined]. Amaral 1930a:60. [Added to synonymy
by Amaral 1930b:237].
Bothrops microphthalma (Cope): Amaral 1930b:237.
Trimeresurus microphthalmus (Cope): Schmidt and Walker 1943:295.
[The authors tentatively referred University of Arequipa specimen
67 to this species.]
Bothrops microphthalma microphthalma (Cope): Peters, 1960:510.
Bothrops microphthalmus microphthalmus (Cope): Rendahl and
Vestergren 1940:15; Hoge 1966:125; Peters and Orejas-Miranda
1970:49; Hoge and Romano-Hoge 1981:211; Carrillo de Espinoza
1983:16; Campbell and Lamar 1989:212; Carrillo de Espinoza and
Icochea 1995:21.
Porthidium microphthalmum (Cope): Schätti and Kramer 1993:266.
Fugler et al. 1995:59.
Bothrocophias microphthalmus (Cope): Gutberlet and Campbell 2001:4.
Diagnosis.—Bothrocophias microphthalmus is distin-
guished from all congeners and from species of Bothrops
and Bothriopsis by the following combination of charac-
ters: (1) lacunolabial absent; (2) dorsal keels tuberculate
on posterior of body; (3) infralabials and some gulars
brown with cream round or C-shaped marks; (4) can-
thorostrals usually present; (5) most subcaudals divided;
(6) ventrals fewer than 160; (7) loreal elongate; (8) can-
thals moderate and separated by 3–5 scales; (9) 1–3 scales
 2005 HARVEY ET AL.—BOLIVIAN PITVIPERS
usually present between internasals; (10) postocular stripe
distinct; (11) dorsum brown to charcoal, pattern of bands
with distinctive row of black spots on dorsal row 4 of neck
and cream stripe on anterior paraventrals.
Description.—Specimens in our sample have total lengths
ranging from 327–724 mm. The tail is 13–15% (14 ± 1, 4)
as long as snout-vent length in females and 19–20% (19 ±
1, 2) in males. Among the adult specimens, the head is
72.9–85.6% (80.5 ± 5.6, 4) as wide as long and accounts
for 5.2–5.6% (5.4 ± 0.2, 4) of snout-vent length in females
and 5.6–5.9% (5.7 ± 0.1, 2) in males. The eye is small, its
greatest diameter is 35.9–60.7% (50.4 ± 9.4, 6) as long as
eye-nostril distance, and its distance to the center of the
naris accounts for 21.4–27.3% (25.0 ± 2.0, 6) of head
length.
The rostral of our specimens is 73–96% (82 ± 10, 6) as
high as wide and 0 (25%, 12), 1 (75%, 12), or 2 (17%, 12)
canthorostrals lie between it and the internasals. The inter-
nasals are not in contact, being separated by 2 (60%, 5) or
3 (40%, 5) small scales. The canthals are narrow and sep-
arated by 4–5 keeled scales; 5–9 (7 ± 2, 7) smooth (57%,
7) or keeled (43%, 7) intersupraoculars are present. Two
specimens have a single doubled intersupraocular. In CBF
1899 and CBGR 24 (29%, 7), each supraocular is divided
into a pair of subequal scales.
The specimens have 4–8 (6 ± 1, 7) prefoveals. The lore-
al of this species is elongate being 73–96% (82 ± 10, 6) as
high as long; it is entire in our specimens and broadly con-
tacts the canthal and entire upper preocular. The middle
preocular and supralacunal are fused on both sides of NK
1814 (14%, 7), partialy fused on both sides of CBF 1899
(14%, 7), and discrete in the remaining specimens (71%,
7). The prelacunal is separated from the supralabials by a
complete row of subfoveals on one side of CBF 1899. On
the other side of CBF 1899 and in the remaining speci-
mens, the prelacunal contacts the second supralabial. All
specimens (100%, 7) have one or two subfoveals at the
juncture of the lacunals and the second and third supralabi-
als. On one side of NK 1814, the subfoveal contacts the
scale inside of the pit, thereby preventing the sublacunal
from contacting the prelacunal. The specimens have two
postoculars, 1–2 suboculars, and 3/3 (86%, 7) or 4/4 (14%,
7) interoculabials. Seven to eight supralabials and 8–10
infralabials are present, the first pair separating the mental
from the chinshields and usually (79%, 14) only the first
two contacting the chinshields; a third infralabial contacts
one or both chinshields in three specimens (21%, 14). The
chinshields are separated from the ventrals by 3–5 (4 ± 1,
7) gulars and 1–3 (2 ± 1, 7) preventrals. Females have
143–147 (145 ± 2, 4) ventrals and 42–45 (44 ± 1, 4) sub-
caudals, and males have 142–152 (147 ± 7, 2) ventrals and
50 (2) subcaudals. Four of the specimens (57%, 7) have
3–7 entire subcaudals, and all subcaudals were divided in
the remaining specimens (47%, 7). Interrictal counts
(26–31, 28 ± 2, 7) are 5–10 scales higher than counts at
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11
midbody. Twenty-one (71%, 7) or twenty-three (29%, 7)
dorsals at midbody reduce to 17–19 dorsals one head-
length in front of the vent.
All specimens in our sample are darkly pigmented. The
dorsal surfaces of the head are a paler brown than the inter-
spaces on the body. Labials and facial scales are brown
and speckled with melanophores. Cream, round, or
C-shaped spots edged in black are present on some infral-
abials and gulars. The dark brown postocular stripe over-
laps one and a half to two scales and crosses the rictus to
extend across about five rows of dorsals. A cream
supratemporal border about two scales wide edges the pos-
tocular stripe dorsally. On the juvenile and smaller adult
specimens, a continuous cream paraventral stripe is well
defined to about the level of the 15th ventral and broken
thereafter for the rest of the body (i.e., the dorsal halves of
the paraventrals are white and their ventral halves are
black. On the posterior part of the body, the white color is
restricted to the keel and anterior part of each paraventral).
In adult specimens the stripe is still visible but grayish in
color. On the anterior half of the flanks, a series of ten sin-
gle and entirely black scales on the fourth scale row are
each separated by 7–12 scales. This row of distinctive
black spots ends on the anterior one-third of the body.
Dorsal bands are clearly present anteriorly but difficult to
count posteriorly in some specimens; 16–18 (17 ± 1, 4)
dorsal bands about 4–7 vertebrals long are separated by
shorter (about 3–4 vertebrals) gray interspaces. The speci-
mens have a single dark charcoal band behind the vent fol-
lowed by a short gray interspace. The rest of the tail is dark
charcoal to black. Above, the distal one-half of the tail of
the juvenile is yellow. The gulars, preventrals, and anteri-
ormost ventrals are cream. Posteriorly, pigmentation of the
ventrals increases rapidly and most are heavily and uni-
formly pigmented. The ventral surface of the tail is heavi-
ly and uniformly pigmented.
Distribution and Comparative Material.—BOLIVIA.
BENI. Gral. José Ballivián: Rurrenabaque (NK 493).
COCHABAMBA. Carrasco: Cerro Leñe, 1580 m,
Parque Nacional Carrasco, 17° 23.367'S, 64°24.320'W
(CBF 1899); Arepucho, Parque Nacional Carrasco, 1000
m, 17°21'57"S, 65°14'30"W (CBG 24–26). LA PAZ. Nor
Yungas: Serranía Bellavista, 15°39'53"S, 67°27'23"W,
1300 m (CBF, uncatalogued). Sud Yungas: Pilón Lajas,
Serranía Chepete (NK 1814), Pilón Lajas, Serranía Beu
(NK 1801).
PERU. UNKNOWN. ANSP 11515 (holotype of
Bothrops microphthalmus).
ECUADOR. PASTAZA. BMNH 1946.1.19.88 (holo-
type of Lachesis pleuroxanthus).
Remarks.—Reports of this species’ presence in Bolivia
have been based solely on a pitviper collected by Orton
from the city of La Paz and identified as Bothrops
microphthlamus by Cope (1879). Now lost, the specimen’s
 12 ANNALS OF CARNEGIE MUSEUM
Fig. 7.—Distributions of Bothrocophias microphthalmus and Bothrops
andianus. Areas above 500 and 1000 m are shaded gray.
identification cannot be verified. Since no pitvipers occur
in the valley of La Paz, the specimens in Orton’s small col-
lection most likely came from lower elevations on the
adjacent humid slopes such as Coroico or the Valle de
Zongo. Cope’s description of the specimen could apply to
either Bothrops andianus or Bothrocophias microphthal-
mus: he noted that the prelacunal and second supralabial
are “partly cut off by suture on both sides.”
Presence of Bothrocophias microphthalmus in Bolivia
is confirmed herein, and the species’ distribution is extend-
ed as far south as Cochabamba (Fig. 7). It almost certain-
ly occurs in the Andean foothills of northern Santa Cruz.
Gutberlet and Campbell (2001) thought that presence
of smooth intersupraoculars was a defining characteristic
of Bothrocophias. Although true for most specimens,
intersupraoculars with low to well-developed keels are
present in 47% (12 specimens) of B. microphthalmus from
Bolivia and elsewhere in South America. Keeled inter-
supraoculars are slightly more frequent (60%, 5) among
B. hyoprora examined by us.
Bothrops Wagler, 1824
Definition.—Bothrops contains 31 species of small (about
500 mm) to large (exceeding 2500 mm) pitvipers with
stout to slender bodies and lacking a rattle. The rostral is
usually about as wide as high (the rostral is much higher
than wide in B. ammodytoides), canthorostrals are absent
except as very rare anomalies, a single entire loreal is pres-
ent, and a lacunolabial may be present or absent. In a few
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VOL. 74
species such as B. jonathani, the prelacunal and second
supralabial may be separated by a row of subfoveals. The
dorsal head plates are fragmented into small, keeled
scales. In most species, intersupraoculars and some scales
in the occipital region are occasionally fused into “dou-
bled” or plate-like scales. Campbell and Lamar (1989,
their tables 29 and 30) summarized variation in selected
meristic characters: species of Bothrops have 139–254
ventrals; 30–91 usually divided subcaudals (several proxi-
mal subcaudals are frequently entire; most or all subcau-
dals are entire in B. venezuelensis); 19–35 middorsal scale
rows; 1–14 intersupraoculars; 6–12 supralabials; 8–16
infralabials; and 2–4 interoculabials. Few generalities can
be made about coloration of Bothrops except that all of the
species are cryptically colored usually with a series of dor-
sal bands or subtriangular to X-shaped markings. A postoc-
ular stripe is usually present, although its width and color
are variable.
The skull of Bothrops is relatively narrow and elongate
(Werman 1992, 1999; Gutberlet 1998; Gutberlet and
Campbell 2001; Gutberlet and Harvey 2002). Osteology is
not known for most species; however, Werman (1992)
summarized variation for a diverse sample of nine species.
The following account based on his data summarizes char-
acters that distinguish Bothrops from other Neotropical
pitvipers. The lateral margin of the prefrontal is concave
and has a single lateral projection in dorsal view. The pos-
torbital is narrow or reduced and may or may not contact
the frontal. Dorsally, the frontals are flat, with elevated lat-
eral margins, and their width is less than or equal to that of
the basicranium. In dorsal view, the parietal is not triangu-
lar, narrowing markedly behind the postorbital. The medi-
al wall of the pit cavity is variable being absent to present
and well developed, and its anterolateral wall forms a
smooth arc or has a small rounded projection. The
ectopterygoid is longer than the base of the pterygoid. In
dorsal view, the anterior portion of the ectopterygoid is
T-shaped or has concave lateral margins. Where it articu-
lates with the pterygoid, the base of the ectopterygoid is
not spatulate and posseses a short, posteriorly directed,
finger-like projection. The shaft of the ectopterygoid is
narrow and somewhat cylindrical in cross section, and a
pair of pits is present on its postero-lateral surface where
the ectopterygoid retractor muscles attach.
Campbell and Lamar (1989) summarize variation in
hemipenial morphology of Bothrops. The hemipenes
extend 6–11 subcaudals, and the sulcus spermaticus bifur-
cates at the level of subcaudals 1–3 in most species.
However, the base of the organ is long in some species
such as B. alternatus so that the sulcus bifurcates between
subcaudals 7–8. Typically, large hooks at the base of the
lobes diminish in size distally, and the distal halves of the
lobes are calyculate. Number and size of the spines are
highly variable and useful in species identification.
Spinules or papillae cover the calyces proximally and are
absent distally.
 2005 HARVEY ET AL.—BOLIVIAN PITVIPERS
Fig. 8.—Male holotype of Bothrops andianus (MCZ 8832).
Content.—Thirty-one species restricted to South America
and the Panamanian Isthmus with the exception of
B. asper which reaches southern Tamaulipas, Mexico:
Bothrops alcatraz Marques, Martins, and Sazima; B. alter-
natus Duméril, Bibron, and Duméril; B. ammodytoides
Leybold; B. andianus Amaral; B. asper (Garman); B. atrox
(Linnaeus); B. barnetti Parker; B. brazili Hoge; B. carib-
baeus (Garman); B. colombianus (Rendahl and
Vestergren); B. cotiara (Gomes); B. erythromelas Amaral;
B. fonsecai Hoge and Belluomini; B. iglesiasi Amaral; B.
insularis Amaral; B. itapetiningae (Boulenger); B. jarara-
ca (Wied-Neuwied); B. jararacussu Lacerda; B. jonathani
Harvey; B. lanceolatus (Bonnaterre); B. leucurus Wagler;
B. lojanus Parker; B. marajoensis Hoge; B. moojeni Hoge;
B. muriciensis Ferrarezzi and Freire; B. neuwiedi Wagler;
B. osbornei Freire-Lascano; B. pictus (Tschudi); B. pirajai
(Amaral); B. sanctaecrucis Hoge; B. venezuelensis
Sandner-Montilla.
Remarks.—In an unpublished doctoral dissertation, Silva
(2000) elevated six subspecies of Bothrops neuwiedi and
recognized one undescribed species. His results have been
adopted by some researchers (e.g., Martins et al. 2002)
who are already referring to these taxa as full species. We
prefer to await publication of Silva’s results before using a
revised taxonomy of Bothrops.
Bothrops andianus (Amaral, 1923)
Figure 6
Lachesis lanceolatus: Barbour, 1913:507 (not of Bonnaterre). [added to
synonymy by Amaral, 1923]
Bothrops pictus: Barbour and Noble, 1920:620 (not of Tschudi). [USNM
60715 is now MCZ 12415; added to synonymy by Amaral, 1923]
Bothrops andiana Amaral, 1923:103. HOLOTYPE male from “Machu
Picchu, Department of Cuzco, Peru, about 9000-10,000 ft. altitude,
by Mr. G. F. Eaton (Yale Peruvian Expedition)” (MCZ 8832) [exam-
ined]. Amaral, 1930b:234.
Bothrops andianus (Amaral): Klemmer, 1963:403; Hoge, 1966: 113;
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13
Peters and Orejas-Miranda, 1970:43; Duellman, 1979:456; Hoge and
Romano-Hoge, 1981:201; Carrillo de Espinoza, 1983:7; Campbell
and Lamar, 1989:188; Carrillo de Espinoza and Icochea, 1995:20;
David and Ineich, 1999:220; McDiarmid et al., 1999:253.
Diagnosis.—Bothrops andianus is distinguished from all
congeners and from species of Bothrocophias and
Bothriopsis by the following combination of characters:
(1) lacunolabial present; (2) dorsal keels nontuberculate;
(3) infralabials and gulars immaculate in adults; (4) can-
thorostrals absent; (5) most subcaudals divided; (6) ven-
trals 166–179 and subcaudals 49–63; (7) loreal subrectan-
gular and elongate; (8) canthals moderate and separated by
3–5 scales; (9) internasals separated by 1–3 scales, rarely
(9%) in contact; (10) postocular stripe distinct and black;
(11) dorsum dull green to brown with pattern of subtrian-
gular blotches or French telephone-shaped marks; black
edging of dorsal blotches becoming 2–4 distinctive rows
of black blotches on posterior body and base of tail.
Redescription of the Holotype.—The holotype (Fig. 8) is a
male with an SVL of 514 mm; the tail length is approxi-
mately 15% of total length. The specimen has a triangular
head, strongly distinct from its neck. The head is longer
than wide, its width being approximately 77% of its
length. Dorsal head scales are keeled except for those
forming the canthus, the supraoculars, two large plate-like
scales in the interocular region, and a pair of plate-like
scales positioned just posterior and medial to the supra-
oculars. The internasals are in contact, five scales separate
the canthals, and three scales separate the supraoculars. A
large, central intersupraocular is about four times as large
as each of the scales flanking it. The supraoculars are 67%
as wide as long and roughly subtriangular. Twenty-six
interrictals are present.
Thickened, lateral edges of the internasal, canthal, and
upper preocular form a sharp and raised canthus rostralis.
Contribution to the canthus by the upper preocular is sub-
stantial and accounts for two-thirds of the length of this
scale. The rostral is 1.15 times higher than wide and about
as wide as the mental. The loreal is subrectangular, 66% as
high as long, and entire; it broadly contacts the canthal and
upper preocular. The foveal pit is bound by entire upper and
lower lacunals and the partially fused second supralabial
and prelacunal. The sublacunal is partially fused to the
lower preocular and broadly contacts the lacunolabial and
third supralabial. Scales are absent from the juncture of the
sublacunal and labials. The specimen has 3/3 prefoveals
and lacks canthorostrals. The diameter of the eye is a little
more than half as large as the distance between the anterior
border of the orbit and the center of the naris. The orbits are
bound by 1/1 supraoculars, 3/3 preoculars, 1/1 elongate
suboculars, and 2/2 postoculars. Each of the preoculars is
entire; the upper is subrectangular and nearly four times as
large as the oval and subequal middle and lower preoculars.
Each of the preoculars enters the orbit on both sides of the
specimen. The subocular is jagged and separated from the
 14 ANNALS OF CARNEGIE MUSEUM
Fig. 9.—Distributions of three Bolivian pitvipers. Open symbols repre-
sent published reports based on specimens not examined by us. Areas
above 500 and 1000m are shaded gray.
labials by one row of smooth scales (=3/3 interoculabials).
The specimen has 7/7 supralabials most of which are sub-
triangular. The foveal pit lies below an immaginary line
drawn between the eye and naris. A nasal pore is present.
The specimen has 9/9 infralabials. The chinshields are
2.6 times as long as wide, contact the first three infralabi-
als, and are separated from the mental by medial contact of
the first pair of infralabials. The mental groove is well
developed and flanked by the first infralabials, chin-
shields, and three rows of gulars. Five gulars and one pre-
ventral separate the chinshields from the ventrals.
Paraventrals are smooth adjacent to the anteriormost ven-
trals but develop increasingly heavy keels posteriorly;
parasubcaudal keels are as well developed as the keels on
the adjacent caudals. The specimen has 21 dorsals at mid-
body, 17 dorsals one head-length in front of the cloaca,
160 ventrals, and 55 subcaudals. All of the subcaudals are
divided. The terminal spine is elongate and slightly
upturned distally.
The original color of the holotype has faded consider-
ably, although pattern is like that of referred material dis-
cussed below. The dorsum and flanks are olive-gray with
dark brown (black in life?) and cream markings. Keels are
the same color as adjacent areas of the same scale. The
dorsal surface of the head is uniformly gray, but almost the
entire stratum corneum is missing. The face, labials, and
underside of the head are pale yellow with very diffuse
gray flecks. A brown (black in life?) postocular stripe
begins on the upper portion of the subocular and covers
one row of temporals, the upper edge of supralabial 5, one-
half of supralabial 6, and three-quarters of supralabial 7.
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VOL. 74
Beyond the rictus, it extends across seven vertical rows of
dorsals. The postocular stripe is edged ventrally by a dif-
fuse line of melanophores then a sharp, cream pinstripe.
Dorsal to the brown postocular stripe lies a cream postoc-
ular border that begins on the postoculars and covers 2.5–3
temporals.
The dorsal pattern consists of inverted U-shaped marks
on the flanks and staggered at the midline. The marks are
separated by pale areas and have irregular dark brown edg-
ing on the anterior portions of the body. Posteriorly, the
dark brown edging becomes more noticeable and develops
into a series of dark brown blotches in the cloacal region.
The dorsal pattern extends onto the tail as rectangular,
brown bands separated by short cream interspaces. The
first six bands are edged in black. Dorsally, the distal one-
sixth of the tail is pale yellow.
The venter is heavily pigmented. Anteriorly,
melanophores are concentrated on the lateral edges of ven-
trals giving the impression of a yellow, midventral stripe.
At the level of the twenty-sixth ventral, pigmentation is
distributed across the venter though not uniformly.
Staggered paler and darker areas form a pattern reminis-
cent of lowland Bothrops atrox. Pigment increases caudad
on the body, and the proximal subcaudals are almost
entirely black. The distal 28 subcaudals are pale yellow
and sharply contrast with the rest of the tail.
In situ, the hemipenis of the holotype extends for 9
subcaudals.
Measurements in mm: SVL 514, tail length 89, head
length 25.69, head width 19.7, eye diameter 3.62, eye-
nostril distance 6.36, loreal length 1.60, loreal height 1.06,
rostral height 4.34, rostral width 3.74, greatest mental
width 3.8, greatest supraocular length 5.26, greatest
supraocular width 3.54, greatest chinshield length 5.20,
greatest chinshield width 2.02.
Variation.—(Except where noted, the following applies to
the Bolivian sample.) As for the holotype, the rostral is
usually slightly higher than wide (rostral width/rostral
height = 0.70–1.04, 0.93 ± 0.9, 17). Contact between the
internasals is rare, occurring in one Bolivian neonate and
the Peruvian holotype (9%, 23). More frequently, speci-
mens from Bolivia and Peru have one (61%, 23), two
(26%, 23), or three (4%, 23) scales between the inter-
nasals. A single canthorostral is present on one side only in
CBG 15; all other specimens lacked this character. The
canthals are separated by 3–5 (4 ± 1, 14) short and keeled
intercanthals, and 4–10 (7 ± 2, 17) keeled scales separate
the supraoculars. Most (73%, 18) specimens have 1–2
fused intersupraoculars, and large, flat plates occupy the
interocular region of some specimens. Flat plate-like
scales and “doubled” keeled scales are also frequently
(about 50% of the specimens) observed in the parietal
region, although this characteristic was scored for few
specimens. The lacunal scales are separated from the nasal
by 2–5 (3 ± 1, 18) prefoveals. The loreal is subrectangular,
 2005 HARVEY ET AL.—BOLIVIAN PITVIPERS
broadly contacts the canthal and supralacunal, and is usu-
ally about one-half as high as long (0.43–0.77, 0.55 ± 0.09,
17). Divided loreal scales were not observed in this
species. The second supralabial and prelacunal are discrete
on both sides of one specimen (CBG 17), but usually are
partially or completely fused. The sublacunal contacts the
lacunolabial and the third supralabial. A small scale is
present at the juncture of the lacunolabial, third supralabi-
al, and sublacunal on both sides of one specimen and one
side only of a second (8%, 14). The sublacunal is entire in
all specimens; it partially fuses to the middle preocular in
one specimen and completely fuses on one side only in a
second specimen. Two specimens have two suboculars on
one side only (6%, 18); all others have 1/1 subocular
(94%, 18) and 2/2 postoculars. In all Bolivian specimens,
the subocular is separated from the supralabials by a row
of keeled scales (3 interoculabials, 100%, 18). However,
the subocular contacts a supralabial 50% (9) of the time in
the Peruvian sample.
In Bolivia, this species has seven supralabials and nine
(78%, 18) or ten (22%, 18) infralabials. Usually (77%, 18),
the first three infralabials contact the chinshields; two or
four infralabials contact a chinshield on one side only in
three specimens. The chinshields are separated from the
ventrals by 4–7 (5 ± 1, 18) gulars and 1–3 (2 ± 1, 18) pre-
ventrals. Males have 166–174 (171 ± 3, 8) ventrals and
52–63 (57 ± 4, 8) subcaudals, and females have 169–179
(174 ± 3, 8) ventrals and 49–58 (52 ± 3, 8) subcaudals.
Usually (58%, 17), the subcaudals are all divided with
42% (17) of the specimens having 2–8 entire subcaudals.
Interrictal counts (23–28, 26 ± 1, 17) were 2–5 scales high-
er than dorsal counts at midbody. In this species, 21 (21%,
18) or 23 (79%, 18) scale rows at midbody reduce to 19
(79%, 18), 18 (16%, 18), or 17 (5%, 18) scale rows one
head-length in front of the vent.
CBF 446 with a total length of 973 mm is the largest
specimen known to us, and 12 specimens from Bolivia
exceed published size records for this species (“464 to 700
mm,” as summarized by Campbell and Lamar 1989). The
tail is 16–19% (18 ± 1, 7) as long as snout-vent length in
males and 15–17% (16 ± 1, 8) as long in females. Head
length accounts for 4.4–5.0% (4.6 ± 0.2, 17) of snout-vent
length, and eye-nostril distance is 23.6–28.8% (26.0 ± 1.4,
16) of head-length.
Recently collected specimens allow us to provide a
more complete description of coloration in this species. In
life, most Bolivian specimens were dull green with some-
what brighter green (bluish in preservative; note that this
is also true for some Peruvian specimens such as USNM
346224) ground color ventrally. The yellow color of the
chin, gulars, and preventrals rapidly fades to green ventral
coloration. In life and in the more recently preserved
Bolivian specimens, the postocular stripe, edging of the
dorsal blotches, skin between the scales, and the tongue
are all black rather than brown as in the holotype. Some
specimens have markings on the dorsal surface of the head
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15
(e.g., a black bar extends across the snout in front of the
supraoculars in CBF 446, and irregular black marks lie on
the interocular and parietal regions of this specimen). The
face and upper edges of the supralabials are olive green,
fading to yellow on the lower portions of the supralabials.
In our sample, the amount of black pigment on the venter
varies from moderate (e.g., NK 2037, UTA 39104) to near-
ly entirely black (NK 2038) posteriorly. In adults, the par-
aventral pattern is usually devoid of yellow blotches; in
most Bolivian specimens it consists of black blotches cov-
ering one-half to two paraventrals and separated by two to
three green scales. Eighteen to twenty-five dorsal blotches
are present and usually resemble poorly defined triangles,
but some specimens (e.g., NK 2037, UTA 39107) have the
French telephone design. Near the tail, black edging of the
dorsal marks becomes broken into 2–4 rows of black
blotches. In some adult specimens, all of the subcaudals
are melanic, whereas in most adults and all neonates the
distalmost 14–37 subcaudals are yellow or yellowish
green (sky blue in preservative). The dorsal surface of the
tail bears 5–12 subrectangular blotches, which are edged
in black proximally but lack black edging distally. In
neonates and adults from a single population, tail tips may
be black (e.g., NK 2014, 2037) or yellowish green (NK
2038–40). The iris is red in Bolivian specimens (Campbell
and Lamar 1989 report that the iris is colored similarly to
the supralabials in Peruvian specimens).
As for most Bothrops, facial pattern changes during
ontogeny. The pale yellow pinstripe edging the postocular
stripe is obvious in juveniles, faded in subadults, and
absent in large adult specimens. Juveniles have heavily
speckled chins and labials, and may also have poorly
defined pale spots on their infralabials. The speckling may
be visible in smaller adults such as the holotype and many
Bolivian specimens, but the labials are immaculate in larg-
er specimens. Among Bolivian specimens, alternating
cream and black paraventral blotches are visible in juve-
niles but not in adults.
Teeth were counted on one side of CBF 446. The spec-
imen has 8 palatine teeth, 24 pterygoid teeth, and 17 den-
tary teeth. The length of the first palatine tooth is about six
times the diameter of its base. The first dentary tooth is
nearly vertical and about six times as long as the diameter
of its base. The remaining dentary teeth become strongly
recurved posteriorly. Measured from the pit cavity, the
fang is one-third as long as head-length, and about 6.5
times as long as its diameter at its base.
Distribution and Comparative Material.—BOLIVIA.
COCHABAMBA. Carrasco: Chaquisacha, Parque
Nacional Carrasco, 1500 m, 17°24'42"S, 65°15'07"W
(CBG 14, CBG 15, CBG 16, CBG 17); Khara Huasi (FML
1039). LA PAZ. Nor Yungas: Parque Nacional y Area
Natural de Manejo Integrado Cotapata (CBF 1927, 1930).
Franz Tamayo: Apolo, El Piñalito, 2290 m (CBF 221).
Sud Yungas: Irupana, 2135 m, 16°28' S, 67°28'W (CBF
 16 ANNALS OF CARNEGIE MUSEUM
446, CBF 467), 12 km SE of Chuspipata on the road to
Chulumani, between 1500 and 2200 m (UTA 39107).
SANTA CRUZ. Caballero: Palmasola (NK 1855, NK
2014) Florida: Parque Nacional Amboró, Río San Rafael
próximo a las Calaminas (NK 2037); Parque Nacional
Amboró, próximo a El Bibosi (NK 2038); Parque
Nacional Amboró, El Bibosi, (NK 2039–40); Santa Rosa
de Lima (NK 2039–46); Bermejo (NK 421); La Yunga,
18°05'30"S, 63°54'30"W (UTA 39104).
PERU. CUZCO. MCZ 8832 (Holotype), MCZ 12415
(Paratype), USNM 346224. PUNO. USNM 267835–39,
UTA 26719.
Remarks.—Amaral (1923) did not supply a separate
description of the holotype, choosing to describe the type
series together. He does not illustrate any of the specimens.
His description is accurate except on two points: First, the
nasals of the holotype and one paratype (MCZ 12415,
other paratypes not examined in this study) are not divid-
ed. However, each nasal bears a pair of short notches, one
above and just posterior to the naris, the other posterior
and below the center of the naris. Second, other than the
canthals and internasals, scales covering the dorsal surface
of the snout are weakly keeled, not smooth.
The distinctive rectangular loreal scale of Bothrops
andianus is relatively rare in Neotropical pitvipers (Table
1). Interestingly, we only found rectangular loreals in
B. andianus and B. lojanus. Known only from Loja and
Zamora-Chinchipe provinces, Ecuador, 2100–2300 m ele-
vation, B. lojanus is another small terrestrial species
restricted to montane regions of the Andes like B. andi-
anus. The rectangular loreal is clearly apomorphic and
suggests a close relationship between these two similar
montane species.
Bothrops atrox (Linnaeus, 1758)
Figure 3
[Coluber] atrox Linnaeus, 1758:824. Two SYNTYPES from “Asia” (in
error, corrected to “Surinam” by Schmidt and Walker, 1943:295)
(“Mus. Ad.Fr.” now NHRM 100–101).
Diagnosis.—With the possible exception of a few prob-
lematic species of the Bothrops atrox complex (see
Remarks), B. atrox is distinguished from all congeners and
from species of Bothrocophias and Bothriopsis by the fol-
lowing combination of characters: (1) lacunolabial pres-
ent; (2) dorsal keels nontuberculate; (3) infralabials and
gulars pigmented to varying degrees but lacking regular
series of cream round to C-shaped marks; (4) canthoros-
trals absent; (5) subcaudals divided; (6) ventrals 182–200
and subcaudals 54–72 [Campbell and Lamar 1989 report
169–212 ventrals and 47–86 based on literature records
from across the range of the species complex. Their wider
ranges are based on published accounts that may not apply
to the species occurring in Bolivia. Our definition is based
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VOL. 74
on specimens we examined]; (7) loreal subtriangular,
higher than long; (8) canthals narrow and separated by 3–7
scales; (9) internasals usually in contact, separated by one
scale in 22% of the sample; (10) postocular stripe distinct
and brown; two temporals high; (11) dorsum brown, pat-
tern of subtriangular blotches or French telephone-shaped
marks; venter uniformly and heavily speckled (popula-
tions on Andean slopes) or with alternating cream and
charcoal, square blotches.
Comparisons among B. atrox Group Species.—Bothrops
atrox, B. brazili, B. moojeni, and B. sanctaecrucis are
rather easily confused with one another (Table 2).
Bothrops sanctaecrucis occurs in sympatry with B. atrox
in Beni and B. moojeni occurs with B. atrox in eastern
Santa Cruz. Although not yet collected in Bolivia,
B. brazili almost certainly occurs with B. atrox in La Paz
and Pando. Of the four, B. atrox is the most easily recog-
nized. Unlike the other species, Bolivian B. atrox have
well-defined, high, dark brown postocular stripes. When
present, the postocular stripes of the other species are one
scale high (vs. 2–2.5 scales high in B. atrox). The venter of
B. atrox is rather heavily pigmented and consists of stag-
gered cream and gray to black (=melanophores) squares
(lowland populations) or is uniformly and heavily speck-
led (specimens from the Andean slopes in La Paz). The
venter of Bothrops brazili is also moderately to heavily
pigmented, but the other two species have cream venters
with only a very diffuse pattern not consisting of staggered
squares. Bothrops brazili and B. sanctaecrucis may further
be distinguished from B. atrox and B. moojeni by fewer
than four (2–4 in B. brazili, 0–3 in B. sanctaecrucis)
keeled scales between the canthals (vs. 2–7 in B. atrox and
B. moojeni). In addition, B. sanctaecrucis has a pale green
iris (pinkish copper to bronze in the other species) and
Bolivian B. moojeni have curving stripes on their necks.
In neonates and juveniles of these four species, the cau-
dal one- to two-thirds of the tail is yellow. The tip of the
tail darkens with age, but it is most distinct in adult
Bothrops sanctaecrucis where the distal tail is black dor-
sally and ventrally sharply contrasting with the base of the
tail. In the other species, the dorsal surface of the tail is
about the same color as the dorsal blotches and does not
sharply contrast with the dorsal pattern. Table 2 summa-
rizes distinguishing characteristics of these similar species.
Description.—Except for Lachesis muta, Bothrops atrox is
the longest pitviper in Bolivia. In our sample, females are
619–1541 mm long (total length, 8), and their tails are
10–21% (16 ± 3, 8) as long as their snout-vent lengths; males
are 307–1009 mm, and their tails are 13–19% (18 ± 2, 7) of
their snout-vent lengths. The head of this species is longer
than wide and accounts for 3.9–5.1% (4.3 ± 0.4, 8) of snout-
vent length in females and 3.5–5.5% (4.3 ± 0.8, 7) in males.
Distance from the anterior border of the eye to the center of
the nostril is 16.5–25.7% (22.9 ± 2.4, 16) of head-length.
 2005 HARVEY ET AL.—BOLIVIAN PITVIPERS
In Bothrops atrox canthorostrals are absent. The inter-
nasals contact one another in most specimens (78%, 9) or
are separated by a single small scale (22%, 9). The can-
thals are narrow and separated by 3–7 (4 ± 1, 19) keeled
scales. In our sample, specimens have 4–10 (7 ± 1, 19)
intersupraoculars, and these scales were usually single and
keeled. A flat, plate-like scale resulting from fusion of
three or four scales lies in the intersupraocular region of
one specimen (AMNH 4038) and one or two doubled
scales are present in six others (30%, 19). Interrictals num-
ber 26–37 (28 ± 3, 20) and are consistently higher than
dorsal counts at midbody.
The rostral is usually slightly higher than wide (rostral
width is 84.2–102.9%, 94.0 ± 6.3, 11, of rostral height).
Although the prelacunal and second supralabial may be
only partially fused (e.g., LSUMZ 45975), a lacunolabial
is invariably present (100%, 20). The sublacunal contacts
the third supralabial and lacunolabial, and a small scale is
not present at the juncture of these three scales. In this
species, the loreal contacts the canthal and is single, sub-
triangular, and 74.4–100.0 (87.2 ± 8.5, 13) as high as long.
The upper preocular contributes to the canthus and is
always entire; the middle preocular is distinct from the
supralacunal. Two postoculars and 1–3 (2 ± 1, 19) suboc-
ulars are separated from the supralabials by one scale in all
but one specimen (3 interoculabials, 97%, 20): a subocular
contacts a supralabial on one side only in AMNH 4038.
This species usually has seven supralabials (7–8, 7 ± 0,
20). Infralabial counts show more variation and range
from 8–11 (10 ± 1, 20), the first 1–4 (3 ± 1, 20) contacting
the chinshields. Three to five (4 ± 1, 20) gulars and 1–3 (1
± 1, 20) preventrals separate the chinshields from the ven-
trals. Females have 189–198 (195 ± 3, 9) ventrals and
54–66 (61 ± 4, 9) subcaudals, and males have 183–200
(192 ± 6, 7) ventrals and 64–72 (67 ± 5, 7) subcaudals. In
all specimens (100%, 19), every subcaudal is divided.
As for most congeners, pattern is boldest in juvenile
specimens of Bothrops atrox. In a typical juvenile from the
Andean population (UTA 38035, Fig. 3), the postocular
stripe is brown and a sharp white line edges the stripe dor-
sally and ventrally. At its widest, the dark brown post-
ocular stripe covers two temporals, and it extends across
some or all of the last three supralabials beyond the rictus
to cover about 11 scales on the neck. The dorsal ground
color of the head is gray. A few black blotches are present
on the head: one between the canthals, a pair forming
poorly defined supratemporal brown stripes. A pale gray
band (2 scales wide) separates the postocular stripe from
the poorly defined supratemporal stripe. In this juvenile
specimen, the rostral, supralabials, and facial scales are
uniformly speckled. Large black spots are in the center of
the rostral, along the posterior edge of the lacunolabial,
and at the juncture of supralabials 4 and 5. On the chin,
black pigmentation is extensive. The mental and anterior
six infralabials have heavy black edges, the mental itself is
almost entirely black, and the chinshields and gulars are
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17
heavily mottled. This specimen has the “French tele-
phone” design on its dorsum. The brown telephone-shaped
marks are edged first in cream then in black. The marks
begin on the fourth dorsal row and are 6–7 dorsals long.
Black paraventral spots overlap the lateral edges of ven-
trals and parts of two paraventrals. Anteriorly on the body,
these spots are separated by 1–2 scales, but they become
more extensive posteriorly where they are separated by
near vertical cream lines. The distal one-fourth of the tail
is pale yellow.
In adults, the labial speckling persists, but is more dif-
fuse and faint. The postocular stripe is always prominent
and crosses 1–2 scales at its widest. On the body, 14–20
(18 ± 2, 13) blotches are followed by 4–7 bands on the tail.
The banding pattern on the tail gives way to a uniformly
pigmented distal portion accounting for the last 28–39
subcaudals. All specimens from the Andean slopes have
uniformly speckled venters, whereas all specimens from
the lowlands have staggered, squarish gray to charcoal
blotches on a cream background.
Distribution and Comparative Material.—BOLIVIA.
BENI. Gral. José Ballivián: Yucumo (NK 488); Pilón
Lajas, Laguna Azul, 340 m (NK 1903). Vaca Díez:
Cachuela Esperanza (AMNH 22524), Tumi Chucua
(USNM 280423–25). Unknown: “Brazil/Bolivia: Río
Beni between Brazil & Bolivia” (AMNH 36190). LA
PAZ. Abel Iturralde: Ixiama-Río Manurimi, Barraca
Santa Rosa, 12°10'S, 67°30'W (NK 2048); Puerto
Moscoso (CBF 1726); Chalalán, Area Natural de Manejo
Integrado Madidi (CBF 1931); Alto Río Madidi, Moira
Camp, 13°35'S, 68°46'W, 270 m (USNM 336150).
Caranavi: “40 km de Caranavi hacia Palos Blancos, 1475
m” (CBF 641). Nor Yungas: Bajo Hornuni, Parque
Nacional Cotapata (CBF 1928–29); stream 4.5 road km W
of Yolosa, a tributary of the Río Elena, 16.20°S 67.78°W
(UTA 38035). Sud Yungas: Pilón Lajas, Serranía Beu,
1500 m; 14°53'15"S, 67°17'31"W (NK 1805). PANDO.
Gral. Federico Román: Río Negro (CBF 1848). Nicolás
Suárez: “12 km by road S Cobija, ca. 8 km W on road to
Mucden” (=Mukden, LSUMZ 45975). SANTA CRUZ.
Velasco: Santa Cruz: Lago Caimán, Parque Nacional
“Noel Kempff Mercado” (NK 934); Flor de Oro, Parque
Nacional “Noel Kempff Mercado” (NK 1295).
UNKNOWN. “interior of Bolivia” (AMNH 4038).
Specimens not examined: BOLIVIA. LA PAZ. Sud
Yungas. “Bellavista, 15°38'S, 67°02'W,” 1500 m (no field
number given, EBD, De la Riva et al. 1992).
UNKNOWN. “Madre de Dios” (we assume that this spec-
imen, identified as Lachesis lanceolatus in the BMNH by
Boulenger 1896, is B. atrox).
Remarks.—The Bothrops atrox complex is the sister group
of a clade composed of B. lanceolatus and B. caribbaeus
and contains populations of cis-Andean lanceheads vari-
ously referred to as B.atrox, B. colombiensis, B. isabelae,
 18 ANNALS OF CARNEGIE MUSEUM
B. leucurus, B. marajoensis, and B. moojeni (Wüster et al.
1996; Wüster et al. 1999; Salomão et al. 1997, 1999;
Wüster et al. 2002). Investigators of species boundaries
within this complex recognize a “moojeni” phenotype for
populations traditionally referred to as B. moojeni in
Argentina, central-southern Brazil, and Paraguay and an
“atrox” phenotype in all other populations (Wüster et al.
1997).
As reflected in our descriptions, we had little difficulty
differentiating between B. atrox and B. moojeni within
Bolivia. Moreover, these two species appear to be morpho-
logically distinct where they occur in sympatry within
Parque Nacional “Noel Kempff Mercado” (Fig. 9).
Nonetheless, species boundaries between B. moojeni and
B. atrox remain problematic elsewhere in South America.
In a multivariate analysis, Wüster et al.(1996) suggested
that a hybrid zone exists where populations of moojeni and
atrox come in contact in Pará and Piauí, Brazil. They sug-
gested three possible explanations for the presence of
seemingly intermediate specimens where the ranges over-
lap: 1. Populations of Bothrops moojeni are southern
B. atrox. 2. The forms are distinct species with converging
adaptations for similar environmental conditions where
their ranges contact one another. 3. Intermediate speci-
mens are hybrids. The status of B. moojeni was further
complicated by results of a phylogenetic analysis using
mtDNA (Wüster et al. 1999). Specimens with the “moo-
jeni” phenotype fell into two clades each of which also
contained specimens clearly referable to B. atrox.
However, levels of divergence were low (2.6–3.6% among
more basal clades).
Fig. 10.—Distributions of Bothriopsis bilineata and Bothrops jonathani.
The open symbol represents a published report based on a specimen not
examined by us. Areas above 500 and 1000 m are shaded gray.
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VOL. 74
In Bolivia, the status of populations of Bothrops atrox
occurring at moderate to high elevations along the Andean
slopes should be reevaluated when more specimens
become available. These snakes have uniformly speckled
venters with no trace of the staggered cream and gray
blotches on the ventrals of specimens from the lowlands.
We could not find other characters to separate these popu-
lations and consider them to be conspecific for the time
being.
Bothrops jonathani Harvey, 1994
Bothrops jonathani Harvey, 1994:61. HOLOTYPE male from “approxi-
mately 35 km N (by road) of El Empalme, Provincia Carrasco,
Departamento de Cochabamba, approximately 2800 m (17°45'S,
65°00'W)” Bolivia (NK 1000) [examined]. Dirksen, et al. 1995:23;
David and Ineich, 1999:229; McDiarmid et al., 1999:267.
Diagnosis.—Bothrops jonathani is distinguished from all
congeners and from species of Bothrocophias and
Bothriopsis by the following combination of characters:
(1) lacunolabial absent; (2) dorsal keels nontuberculate;
(3) labials with distinctive pattern: white with tan or black
edges; (4) canthorostrals absent; (5) all subcaudals divid-
ed; (6) ventrals 154–172 and subcaudals 37–42; (7) loreal
subtriangular; (8) canthals narrow and separated by 4–6
scales; (9) internasals in contact; (10) postocular stripe 4.5
temporals high, brown edged in black; (11) dorsum pale
brown with dark brown X- or V-shaped marks; venter
heavily pigmented with charcoal blotches; (12) counts for
dorsals (27–33), supralabials (9–12), infralabials (13–16),
and interoculabials (4) unusually high for Bothrops.
Description.—The largest specimen of this relatively
small species is an 881 mm female (NK 1618). The largest
male specimen is the holotype (620 mm). The tail is
13–16% (15 ± 2, 2) of snout-vent length in males and
11–12% (11 ± 0, 2) in females. The head is 66.5–80.4%
(74.7 ± 6.8, 4) as wide as long, and its length accounts for
4.6–5.2% (5.3 ± 0.8, 5) of snout-vent length in adults, 7%
in a 285 mm juvenile. Eye-nostril distance accounts for
22.3–24.6% (23.4 ± 0.9, 6) of head-length.
This species lacks canthorostrals and the internasals are
in contact. The canthals are separated by 4–6 (5 ± 1, 6)
keeled scales. The supraoculars are narrow and separated
by 8–12 (10 ± 2, 7) keeled scales. “Doubled” intersupraoc-
ulars occur in three specimens, however large plate-like
scales do not occur between the supraoculars or in the pari-
etal region of this species. Interrictal counts (31–37, 33 ±
2, 6) were 0–8 scales more than dorsal counts (27–33, 30
± 2, 7) at midbody.
The rostral of this species is 85.0–103.2% (93.4 ± 8.2,
6) as wide as high. Four to eight (5 ± 1, 7) prefoveals are
present, and the lacunals are always (100%, 6) separated
from the supralabials by a complete row of subfoveals
(lacunolabial absent). The loreal is entire, subtriangular,
 2005 HARVEY ET AL.—BOLIVIAN PITVIPERS
69 64 59
10
Bothriopsis taeniata
Bothrops neuwiedi
15
20
200 km
69 64 59
Fig. 11.—Distributions of Bothriopsis taeniata and Bothrops neuwiedi.
Open symbols represent published reports based on specimens not exam-
ined by us. Areas above 500 and 1000 m are shaded gray.
and 75.0–107.4% (96.4 ± 19.6, 6) as high as long. It broad-
ly contacts the canthal and the supralacunal. Unlike other
Bolivian pitvipers, the upper preocular is usually (83%, 6)
divided into a small, square preocular and a much larger
scale that contributes to the canthus. The middle preocular
and the supralacunal are fused in the paratype, but discrete
in the other specimens. Usually (83%, 6), the sublacunal
and prelacunal are separated by one or two subfoveal
scales that enter the pit. Four to eight suboculars and pos-
toculars are separated from the supralabials by two rows of
smooth scales (=4 interoculabials, 100%, 6). This species
has 9–12 (11 ± 1, 7) supralabials and 13–16 (14 ± 1, 7)
infralabials, the first three contacting the chinshields. The
first pair of infralabials separate the mental from the chin-
shields. Five to nine (6 ± 1, 7) gulars and 1–3 (2 ± 1, 7)
preventrals separate the chinshields from the ventrals.
Males have 156–164 (159 ± 4, 3) ventrals and 39–42
(40 ± 2, 3) subcaudals, and females have 161–172 (167 ±
6, 3) ventrals and 37–39 (38 ± 1, 2) subcaudals. All sub-
caudals are divided in this species.
In several ways, coloration of Bothrops jonathani dif-
fers noticeably from other Bolivian pitvipers. The dorsum
is dark tan with dark brown blotches edged in cream. The
postocular stripe is brown and edged above and below in
black. It is higher than in most species, overlapping four
and one-half scales at its highest point. The labials have a
complex pattern, being white with dark tan or black edges.
The ventral surface of the head is white with black and
smoke-gray markings. Twenty-four to thirty (26 ± 2, 6) V-
to X-shaped marks are present on the dorsal body and 5–9
(8 ± 2, 6) are present on the tail. The distal portion of the
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19
tail does not have a distinct, paler pattern although it is
10
uniformly gray ventrally in some specimens. The ventrals
are white with a ventral pattern consisting of staggered
smoke-gray to dark charcoal bands.
A color photo of this species was published by Dirksen
et al. (1995, fig. 9 and 10), presumably of ZFMK 57523.
Distribution and Comparative Material.—BOLIVIA.
15 COCHABAMBA. Carrasco: “approximately 35 km N
(by road) of El Empalme,” 2800 m (NK 1000, holotype);
“on the old highway connecting Santa Cruz to
Cochabamba, 97 km S (by road) of Cochabamba, 3220 m”
(UTA 34564). SANTA CRUZ. Caballero: San Juan del
Potrero (NK 1618 and 2036). Florida: “La Yunga de
Mairana,” 1960 m (NK 2041); El Fuerte, 2000 m (ZFMK
20 57523 and 58160). Vallegrande: 5 km E of Santa Rosita
(NK 718).
In addition to these localities, the species has also been
collected near Samaipata, Florida, Santa Cruz (Dirksen et
al., 1995: ZFMK 60172). This species appears to be
restricted to dry valleys of the Andes between 1900–3300
m (Fig. 10).
Bothrops moojeni Hoge, 1966
Bothrops atrox: Serié, 1915:106, 1921:169; Abalos and Mischis, 1975:75
(not of Linnaeus)
Bothrops atrox atrox: Hoge, 1953:195. (not of Linnaeus)
Bothrops moojeni Hoge, 1966:126. HOLOTYPE from “Brasília, Distrito
Federal, Brasil” (IB 23397). Peters and Orejas-Miranda, 1970:49;
Hoge and Romano, 1973:137; Hoge et al., 1975; Hoge et al.,
1981:91; Hoge and Romano-Hoge, 1981:211; Nascimento et al.,
1988:57; Campbell and Lamar, 1989:212; Cei, 1993:722;
Strussmann and Sazima, 1993:161; Wüster et al., 1994:76. Jorge da
Silva and Sites, 1995:898; Aquino et al., 1996:376; Harvey, 1998:
148 and 353; Buongermini and Waller, 1999:53; Freitas, 1999:72;
David and Ineich, 1999:232; McDiarmid et al., 1999:267.
Botrops moojeni (Hoge): McDiarmid and Foster, 1987:7. [misspelling]
Bothrops atrox moojeni (Hoge): Itoh et al., 1987:3132.
Diagnosis.—Bothrops moojeni is distinguished from all
congeners and from species of Bothrocophias and
Bothriopsis by the following combination of characters: (1)
lacunolabial present; (2) dorsal keels nontuberculate; (3)
infralabials and gulars immaculate; (4) canthorostrals
absent; (5) subcaudals divided; (6) ventrals 179–210 and
subcaudals 44–70 (Campbell and Lamar 1989); (7) loreal
subtriangular; (8) canthals narrow and separated by 4–7
scales; (9) internasals in contact (33%, 9) or separated by
1–2 scales; (10) postocular stripe absent or dark brown to
gray and narrow: less than one temporal high; (11) dorsum
gray to brown, pattern of subtriangular blotches; venter
cream with very diffuse if any pigmentation except on
edges; distinctive wavy occipital stripe extending onto neck.
Description of Bolivian Specimens.—We are aware of
only three museum specimens of this species from Bolivia.
A male specimen (NK 609) has a damaged head, and some
 20 ANNALS OF CARNEGIE MUSEUM
head measurements were not taken for this specimen. Two
female specimens are in relatively good condition.
The largest specimen is an adult female with a snout-
vent length of 975 mm and tail length of 157 mm. The tail
is 16–17% as long as snout-vent length in females and 19%
as long in the male specimen. The head is longer than wide
(head width/head length = 0.63–0.67 in females) and
accounts for 3% of snout-vent length. The eye is 51–80%
(2) as long as eye-nostril distance. These specimens lack
canthorostrals, and the internasals are in contact medially.
The canthals are narrow and separated from one another by
4–5 rows of keeled scales. In two specimens, the canthus is
formed by the internasal, canthal, and part of the upper pre-
ocular; however, the last scale is excluded from the canthus
by broad canthal-supraocular contact in the male specimen.
Seven (NK 609) or 10 keeled scales separate the supraocu-
lars; plate-like or fused intersupraoculars are not present.
Interrictal counts were the same (NK 610) or one to two
scales more than number of dorsals at midbody.
Two to four prefoveals are present in front of the
lacunolabial. The sublacunal contacts the third supralabial,
and no scale is present at the juncture of the lacunolabial,
third supralabial, and sublacunal. The loreal is single, sub-
triangular, and about as long as high. The middle preocu-
lar is distinct from the supralacunal, and neither the upper
preocular nor the sublacunal is divided. An elongate sub-
ocular is present in two specimens, but this scale is divid-
ed into 2/3 scales in NK 609. Two postoculars are present
in all specimens. The specimens have 3/3 interoculabials,
7/7 supralabials, and 10/10 infralabials, the first three con-
tacting the chinshields. Four to five gulars and one preven-
tral separate the chinshields from 196 (two females) or 197
ventrals and 60–62 (females) or 65 (male) divided subcau-
dals. Twenty-five to 27 dorsals reduce to 21 dorsals one-
head length in front of the cloaca.
The three Bolivian specimens have a narrow postocular
stripe about one scale high and curving around the rictus
where the stripe is thickest. A pair of stripes begins just
above the rictus and extends along the neck to the level of
the 12–24th ventral. The dorsal pattern consists of 14–15
brown triangular blotches, edged in black lines that slight-
ly curve into the triangles ventrally. These blotches are 6–9
dorsals at their longest point on the flanks. As in most
species of Bothrops, the blotches meet dorsally or are stag-
gered along the midline. Two or three bands are visible
proximally on the tail, but the dorsal surface of the tail is
uniformly gray (large adult) or tan (subadults) for two-
thirds of its length. The labials and chin are nearly immac-
ulate. A few diffuse blotches are present on the first infral-
abials of NK 610. The ventrals are cream; adjacent to dif-
fuse paraventral spots, every other ventral has a diffusely
black pigmented edge. The subcaudals are very diffusely
pigmented to the tip of the tail.
Distribution and Comparative Material.—On the basis of
specimens examined in this study, Harvey (1998) first
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VOL. 74
reported Bothrops moojeni for Bolivia (Fig. 9). The range
depicted by Campbell and Lamar (1989) in their map 71
was extended northward and westward much closer to the
Bolivian border by Nascimento et al. (1988) who reported
the species from Chapado dos Guimarães and the Serra das
Araras in central Mato Grosso, Brazil. Wüster et al. (1994)
and Strussman and Sazima (1993) reported B. moojeni
from localities in the northern Pantanal of Mato Grosso.
Specimens Hoge (1953) identified as Bothrops atrox from
Chavantina on the Rio das Mortes in eastern Mato Grosso
are probably B. moojeni. Based on the recent finding of
Martínez and Martínez, Cei (1993) argued that reports of
B. atrox from Misiones, Argentina, are based on specimens
of B. moojeni. At present, the only museum specimens
from Bolivia have been collected on top of the Serranía de
Huanchaca, a table-top mountain straddling the border
between Bolivia and Brazil (Harvey 1998, 1999; Harvey
and Gutberlet 1998). We have seen photographs (with
some morphological data, P. Strimple, personal communi-
cation) of live Bothrops moojeni from El Tumbador near
Corumbá in the Bolivian Pantanal (Germán Busch
province).
BOLIVIA. SANTA CRUZ. Germán Busch: El
Tumbador (no museum number, photo only). Velasco:
Parque Nacional “Noel Kempff Mercado” (NK 156); Las
Gamas, Parque Nacional “Noel Kempff Mercado” (NK
609–610).
BRAZIL. MATO GROSSO DO SUL. USNM
100749. PARANÁ. FML 2506, 2513–14. SÃO PAULO.
USNM 165485.
PARAGUAY. AMAMBAY. USNM 253143–44,
342425–26.
Remarks.—McDiarmid et al. (1999) discuss the problem-
atic allocation of Bothrops leucostigma Wagler.
Hoogmoed and Gruber (1983) suspect that the type speci-
men may be B. moojeni rather than B. jararaca as thought
by other authors (Peters and Orejas-Miranda 1970;
Vanzolini 1981).
Bothrops neuwiedi (Wagler, 1824)
Bothrops Neuwiedi Wagler, 1824:56. Likely HOLOTYPE from “provin-
cia Bahiae” (Bahia province, Brazil, according to Vanzolini,
1981:xxiv) (ZSMH 2348/0). [Holotype was designated as a lectotype
by Hoogmoed and Gruber (1983)].
Diagnosis.—Bothrops neuwiedi is distinguished from all
congeners and from species of Bothrocophias and
Bothriopsis by the following combination of characters:
(1) lacunolabial absent; (2) dorsal keels nontuberculate;
(3) infralabials and gulars diffusely pigmented but lacking
distinct cream circular to C-shaped marks; (4) canthoros-
trals absent; (5) all subcaudals divided; rarely with few
entire subcaudals; (6) ventrals 165–186 and subcaudals
40–54 (Campbell and Lamar 1989, report 163–190 ven-
 2005 HARVEY ET AL.—BOLIVIAN PITVIPERS
trals and 37–54 subcaudals in this polytypic species); (7)
loreal subtriangular; (8) canthals narrow and separated by
3–6 scales; (9) internasals in contact; (10) postocular stripe
brown, edged in black, 1.5–3 temporals high; (11) dorsum
straw-colored to pale brown or gray with darker brown,
French telephone-shaped or subtriangular markings; large,
usually V-shaped dark brown blotches on dorsal surface of
head; ventral surface of head lacking pair of brown gular
stripes.
Description.—Bolivian Bothrops neuwiedi probably never
exceed a meter and a half in total length. In our sample,
females are 415–1032 mm long (total length), and their
tails are 12–16% (14 ± 1, 13) as long as their snout-vent
lengths; males are 318–830 mm and their tails are 15–19%
(17 ± 1, 24) of their snout-vent lengths. The head of this
species is longer than wide and accounts for 3.6–5.1%
(44.2 ± 0.4, 13) of snout-vent length in females and
4.1–5.9% (4.3 ± 0.4, 22) in males. Distance from the ante-
rior border of the eye to the center of the nostril is
22.3–31.1% (25.5 ± 2.2, 26) of head-length.
In Bothrops neuwiedi, canthorostrals are absent and the
internasals contact one another (100%, 21). The canthals
are narrow and separated by 2–6 (4 ± 1, 32) keeled scales.
In one specimen (CM 2730), two large scales between the
canthals appear to have formed through fusion of the first
and second rows of scales between the canthals and inter-
nasals. Specimens we examined had 5–10 (7 ± 1, 39) inter-
supraoculars, and these scales were usually single and
keeled. Thirty-five percent (39) of the specimens had 1–4
fused scales between the supraoculars, and these com-
pound scales were usually doubled and keeled. Interrictal
counts (23–30, 26 ± 2, 37) were 0–8 scales higher than
dorsals at midbody (23–27, 25 ± 1, 39).
In our sample, variation in scales associated with the pit
is considerable. Three to eight prefoveals (5 ± 1, 38) lie
between the postnasal and prelacunal. The prelacunal con-
tacts the second supralabial in all (97%, 32) but one spec-
imen (MCZ 28603) where these scales are separated by a
row of subfoveals. Condition of the sublacunal was scored
for a larger sample than other characters. Most frequently
(56%, 32), the sublacunal contacts both the prelacunal and
the third supralabial, and a single subfoveal is present at
the juncture of the sublacunal, prelacunal, and supralabi-
als. In several specimens (18%, 32), the subfoveal is
absent. Rarely, two subfoveals are present (4%, 32), or the
sublacunal does not contact the third supralabial (4%, 32).
On one side only of MCZ 20620, the sublacunal is divid-
ed into subequal anterior and posterior portions. About 6%
(32) of the time, the sublacunal and prelacunal are separat-
ed by contact between the scale inside the pit and the
subfoveal.
In this species, the loreal contacts the canthal and is sin-
gle, subtriangular, and 73.9–133.3% (99.9 ± 15.1, 24) as
high as long. The upper preocular contributes to the can-
thus and is entire. The middle preocular is distinct from the
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21
supralacunal; in one specimen (CM 2730), these two
scales are separated by a single square scale on both sides.
Two postoculars and 1–6 (2 ± 1, 38) suboculars are sepa-
rated from the supralabials by one scale in most specimens
(3 interoculabials, 92%, 39). Four interoculabials are pres-
ent on one side only in three specimens (8%, 39). This
species usually has eight supralabials (8–10, 8 ± 0, 38).
Infralabial counts varied more frequently and ranged from
10–13 (11 ± 1, 38), the first 2–4 (3 ± 0, 38) contacting the
chinshields. Two to five (4 ± 1, 38) gulars and 1–4 (2 ± 1,
39) preventrals separate the chinshields from the ventrals.
Females have 170–189 (177 ± 5, 15) ventrals and 40–50
(45 ± 3, 14) subcaudals, and males have 165–186 (172 ±
5, 24) ventrals and 44–54 (50 ± 3, 24) subcaudals. In all
but one specimen (97%, 39), every subcaudal was divided;
the third through fifth subcaudal were entire in CM 2729.
The dorsal surface of the head is gray to tan with brown
markings. Peñaranda et al. (1994, fig. 6) observed such
extreme variation in dorsal head pattern among 60
Bolivian neonates from four clutches that they were able
to identify individuals on the basis of head pattern alone
(Abalos and Baez 1963 document similar variation for
Argentinean populations). Specimens typically have a
large blotch (usually single) on the dorsal surface of the
snout. The markings on the head tend to be V-shaped,
although they are frequently broken into a series of blotch-
es extending from the interocular region obliquely onto the
neck. The postocular stripe is brown and sharply edged
ventrally in black. At its widest, it covers 1.5–3 temporals.
It extends across the last three to four supralabials and onto
about four scale rows beyond the rictus. The labials and
scales of the chin are diffusely pigmented, the
melanophores tending to be most concentrated near scale
margins.
Relative to most Bolivian pitvipers, the dorsal pattern
of Bothrops neuwiedi appears both bolder and more com-
plex. The 17–25 (20 ± 4, 31) dorsal marks are usually a
“broken” version of the “French telephone” pattern: the
marks are usually staggered and each is associated with a
pair of round spots on the flanks. The marks are narrowly
edged first in tan then in black and have dark brown cen-
ters. The dorsal ground coloration ranges from straw-
colored to pale brown to gray. Paraventral blotches are
poorly defined though darkly pigmented and overlap the
lateral edge of the ventrals and the first two rows of dor-
sals. Notably, the sides of the tail are nearly immaculate
pale brown; 3–7 subrectangular dark brown blotches are
restricted to the dorsal surface of the proximal two-thirds
of the tail. In neonates, the distal one-third of the tail is
pale yellow, however this distal portion becomes dark
brown (in specimens from Santa Cruz) or tan (e.g., UTA
38038 from Tarija) with age.
Natural History.—Peñaranda et al. (1994) present repro-
ductive and growth data for Bolivian Bothrops neuwiedi.
Gestation period was nine months in captive specimens
 22 ANNALS OF CARNEGIE MUSEUM
maintained in La Paz (Peñaranda et al. 1994) where ambi-
ent temperatures are considerably cooler than in the low-
lands where most populations of this species occur.
Although previously thought to occur only up to 600 m
(Campbell and Lamar 1989), some new material in this
study comes from the intermontane valleys and slopes of
the Andes in Santa Cruz and Chuquisaca. Bothrops
neuwiedi occurs in grassland, dry forest, and scrub forest
at least as high as 2000 m and is sympatric with B.
jonathani in the vicinity of the El Fuerte ruins near
Samaipata, Santa Cruz (Fig. 11).
Distribution and Comparative Material.—BOLIVIA.
BENI. Yacuma: Espíritu (CBF 358, 449). Gral. José
Ballivián: Yucumo, Estancia Porvenir, 47 km E of San
Borja (CBF 412, 460–61). Mamoré: “Estancia Yutiola, ca
20 km S San Joaquín” (AMNH 104578); “ca 2 km N San
Joaquín,” (AMNH 104579); San Joaquín (FMNH 140200,
FMNH 161561–63). CHUQUISACA. Luis Calvo: El
Salvador, Cantón Carandayti (FML 2388, 2396). Sud
Cinti: trail between El Palmar and Limón (UTA 39112).
SANTA CRUZ. Andrés Ibáñez: Santa Cruz de la Sierra
(CM 119–120, CM 122–123, FML 1033, NK 124, 168,
189, 197, 205, 330, 475, 556, 719, UTA 34559–60);
Terevinto, las pampas de Urubó (NK 178, 490); “Guafilo
cera de Cotoco” (NK 179); El Vallecito (NK 227);
Potrerillo de Wendá, Cantón Terebinto (NK 957–59,
1419); savana W Río Piray on road from Santa Cruz de la
Sierra to Comunidad Las Cruces, 17.75°S 63.25°W (UTA
38037). Caballero: San Juan del Potrero (NK 686).
Chiquitos: “Santiago”=Santiago de Chiquitos, 700 m
(FMNH 195872, FMNH 195900, FMNH 195904); San
José (CM 34828). Cordillera: Perforación (NK 1886–87);
San Antonio del Parapetí, 20°1'S, 63°13'W (AMNH
141425-141432); “La Brecha, ca 104–120 km NE
Charagua, Izozog Region,” Cordillera province (AMNH
141433–36). Florida: “Río Colorado” (CM 2856–59); dirt
road leading from highway near Samaipata to El Fuerte
ruins, ca. 0.5 km S of Quebrada El Fuerte (UTA 38036); La
Yunga near Mairana (UTA 38075); Mataral (NK 899, 1029,
1469, 1625, 1877, 1897); Los Negros (NK 1606);
Pampagrande (NK 717, 860–61, 902, 1266, 1271, 1273,
1269, 1475, 1509, 1519, 1720, 1856, 1870, 1984). Germán
Busch: Puerto Busch (NK 1187). Ichilo: Buena Vista
(AMNH 36008–09, CM 2712–13, CM 2724–27, CM
2729–30, CM 2802, CM 2814–15, CM 2819, CM 2896,
CM 2903–04, CM 2923–26, CM 2933–34, CM 2963, CM
2965, MCZ 20620–22); “Río Surutú, W of Buena Vista,”
(CM 2773, CM 2913). Ñuflo de Chávez: San Ramón (NK
503). Sara: no other data (MCZ 11857). Vallegrande:
Vallegrande, 2000 m (NK 847). Unknown: no other data
(MCZ 29229–31). TARIJA. Gran Chaco: Villamontes
(MCZ 28603); on road from Villamontes to Entre Ríos, 38.0
km W of the Hoterma on the Río Pilcomayo (UTA 38038).
Note: CM specimens in the list above from Buena Vista
are paratypes of Bothrops neuwiedii bolivianus Amaral.
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VOL. 74
Specimens not examined by us: BOLIVIA. BENI.
Gral. José Ballivián: “Estación Biológica Beni” (EBD,
no museum number given, Fugler and De la Riva 1990),
“50 km E de San Borja” 200 m (EBD no museum number,
De la Riva et al. 1992); Yacuma: Santa Ana de Movimas
(BMNH, Boulenger 1898. Now referred to as just “Santa
Ana,” the old name refers to the Movimas Indians native
to the area). SANTA CRUZ. Chiquitos: Cañada larga
(Peñaranda et al. 1994). Florida: Vicinity of Samaipata
(Dirksen et al. 1995: ZFMK 60173). TARIJA. Arce:
“Quebrada de San Telmo–Cooperativa John Kennedy.
22°30'S, 64°22'W” (FML uncatalogued, collectors tag GM
2009 M, Lavilla and Scrocchi 1999). Gran Chaco: Caiza,
Aguairenda, San Francisco (Peracca 1897); “Tatarenda,
Caiza, Bolivian Chaco” (Lönnberg 1902); “Villa Montes”
(Mertens 1929).
Remarks.—Campbell and Lamar (1989, fig. 233) include
a photograph taken by P. Bettella of a snake from the mesa
of the Serranía de Huanchaca. Bettella and the authors
identified this snake as Bothrops neuwiedi. Extensive col-
lecting in this area has failed to produce specimens of
B. neuwiedi. The dorsal pattern of the specimen is unlike
Bolivian B. neuwiedi but closely resembles B. moojeni
from the Serranía. We cannot locate Bettella’s specimen
and suspect that he misidentified it or that data sent to
Campbell and Lamar was in error. We do not include the
Serranía de Huanchaca in the distribution of B. neuwiedi.
Although the type series of Bothrops neuwiedi boli-
vianus is enormous (71 specimens), Amaral (1927) does
not describe the series and provides only limited data for
the holotype. We examined most of the type series, and
Griffin (1916) tabulated meristic and mensural data for it.
Bothrops sanctaecrucis Hoge, 1966
Lachesis lanceolatus: Griffin, 1916:222 (in part: CM 313 now MCZ
17699; CM 43 and CM 121 now lost) (not of Bonnaterre).
Bothrops atrox: Amaral, 1926:320 (in part: CM 313 now MCZ 17699);
Amaral, 1927:6 (not of Linnaeus)
Bothrops jararacussu Lacerda: Amaral, 1925:43, 1926:320 (in part, not
of Lacerda).
Bothrops sanctaecrucis Hoge, 1966:133. HOLOTYPE from “Oromomo,
Río Sécure, upper Beni, Bolivia” (IB 24575). Peters and Orejas-
Miranda, 1970:54; Hoge and Romano-Hoge, 1981:219; Fugler,
1986:48; Campbell and Lamar, 1989:223; Fugler and De la Riva,
1990:38; Fugler et al., 1995:59; David and Ineich, 1999:236;
McDiarmid et al., 1999:271; Middendorf and Reynolds, 2000:162.
Diagnosis.—Bothrops sanctaecrucis is distinguished from
all congeners and from species of Bothrocophias and
Bothriopsis by the following combination of characters:
(1) lacunolabial present; (2) dorsal keels nontuberculate;
(3) infralabials and gulars immaculate; (4) canthorostrals
absent; (5) subcaudals divided; (6) ventrals 175–191 and
subcaudals 55–65; (7) loreal subtriangular; (8) canthals
wide, in contact or separated by 1–3 scales; (9) internasals
in contact or rarely separated by 1 small scale; (10) post-
 2005 HARVEY ET AL.—BOLIVIAN PITVIPERS
ocular stripe absent or dark gray to black and narrow: less
than one temporal high; (11) dorsum tan to pale gray with
brown X-shaped or “French telephone”-shaped marks first
edged in cream then in black; venter, labials, and chin
immaculate cream (diffusely pigmented in juveniles); dis-
tal one-half of tail black in adults.
Description.—Bothrops sanctaecrucis is a large terrestrial
pitviper attaining a total length of at least 1303 mm (CBF
1023, female, snout-vent length 1151, tail length 152). The
tail is 13–17% (15 ± 1, 8) as long as snout-vent length in
females and 16–21% (19 ± 2, 6) as long in males. The head
is 45.4–79.5% (61.9 ± 11.4, 11) as wide as long and
accounts for 4.1–5.1% (4.4 ± 0.2, 5) of snout-vent length
in females and 4.0–4.8% (4.4 ± 0.3, 5) in males. Eye-
nostril distance accounts for 20.2–25.7% (23.9 ± 1.8, 8) of
head length. In B. sanctaecrucis, canthorostrals are absent;
9% (32, includes data from 19 neonates) of the specimens
had a small scale between the internasals. The canthals are
broad and usually separated from one another by 1–3 (36,
includes data from 19 neonates) keeled scales. In two of a
series of 19 neonates and in one adult female (AMNH
6771), the canthals broadly contact one another medially.
The canthus is formed by the internasal, canthal, and part
of the upper preocular. Six to eight (7 ± 1, 18) usually
keeled scales separate the supraoculars. The intersupraoc-
ulars are occasionally (31%, 18) fused to form doubled,
keeled scales; one specimen (NK 226) had a large platelike
intersupraocular about three times as large as adjacent
scales. Interrictal counts (25–29, 27 ± 1, 17) were 1–5
scales more than the number of dorsals at midbody
(21–27, 24 ± 2, 17).
One to three (2 ± 1, 16) prefoveals are present in
front of the lacunolabial. In all specimens, the sublacu-
nal contacts the third supralabial and subfoveals are usu-
ally absent: a subfoveal is present at the juncture of the
lacunolabial, third supralabial, and sublacunal on one
side of a single specimen (CBF 673; 3%, 17). The lore-
al is single, subtriangular, and about as long as high. It
usually contacts the canthal; however, in one specimen
(MCZ 17699), a rectangular scale separates the other-
wise normal (i.e., subtriangular) loreal and canthal on
both sides of the head. The middle preocular is distinct
from the supralacunal, and neither the upper preocular
nor the sublacunal is divided. An elongate subocular and
two postoculars are present in most specimens; the sub-
ocular is divided into two scales 12.5% (18) of the time.
The specimens have 3/3 interoculabials, usually eight
supralabials (nine supralabials are present on one side
only of two specimens), and 9–11 (usually 10) infralabi-
als, the first three contacting the chinshields. Three to
five (4 ± 0, 18) gulars and 1–3 (1 ± 1, 18) preventrals are
present. Female specimens have 178–191 (184 ± 3, 9)
ventrals and 55–61 (58 ± 3, 9) subcaudals, whereas
males have 175–180 (177 ± 2, 6) ventrals and 59–65 (63
± 2, 6) subcaudals.
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23
The iris of Bothrops sanctaecrucis is uniformly pale
green with few black specks. The head and body are tan to
pale gray with 17–21 (18 ± 2, 12) X-shaped or “French
telephone”-shaped marks. The dorsal marks are edged first
in cream, then in black; their centers are brown. In two
well-preserved specimens (CBF 673, 1009), a pair of
oblique, cream lines two scales wide edge the supraoculars
and extend obliquely across the supratemporal region to
the level of the rictus. We could not tell if these lines are
present in most of our specimens because they are faded;
however, some well-preserved specimens clearly lack the
cream lines. Three (17%, 18) of our adult and juvenile
specimens have a narrow, dark brown postocular stripe
about one to one-half of a temporal wide. The postocular
stripe is faint over the postoculars and temporals but dark-
ens where it crosses the tops of the sixth and seventh
supralabials to extend across the posterior one-half of the
last supralabial and the scale just behind it. In young
adults, the labials may be finely speckled, although the
labials become immaculate cream with age. The chin
and ventrals are nearly immaculate cream or they may be
diffusely pigmented with scattered melanophores.
Paraventral spots are poorly defined; each occupies two
paraventrals and one scale on the adjacent row of dorsals.
The paraventral spots are separated from one another by
one scale and from the dorsal blotches by three scales. The
subcaudals become increasingly pigmented distally; the
distal one-half of the tail is black in adults.
The distal third of the tail of neonates and young
juveniles is yellow. Like most Bothrops, neonates of
B. sanctaecrucis have heavily mottled chins and labials;
circular white spots are not present. None of the series of
19 neonates has a dark postocular stripe.
Distribution and Comparative Material.—BOLIVIA.
BENI. Moxos: Oromomo, “orillas del Río Sécure,” 250 m
(CBF 1009, 1023). COCHABAMBA. Carrasco:
Campamento Yuquis, “cerca al Río Chimore,” 16°47'OO"
S, 64°56'50" W (CBF 673, 776). Unknown: specific local-
ity unknown (AMNH 6771–72). SANTA CRUZ. Andrés
Ibáñez: Santa Cruz de la Sierra, province (MCZ 17699).
Ichilo: Buena Vista (MCZ 20618-19); San Carlos (NK
226); Yapacaní (NK 248, 358); El Condor-Moile (NK
359); El Potrerillo (NK 518, 633, 839); Río Maroñucú,
Parque Nacional Amboró (NK 619); Río Cheyo, camino al
campamento La Chonta (NK 763).
Not examined: IB 24576 (from Oromomo), UMMZ
(26 paratypes from Buena Vista), UMMZ and CM (15
paratypes from Río Surutú). Partially based on Miranda
et al. (1991), Middendorf and Reynolds (2000) report
B. sanctaecrucis from the Beni Biological Station
Biosphere Reserve (Gral. José Ballivián and Yacuma
provinces, 14°40'S, 66°30'W, 220 m) (Fig. 9).
Remarks.—Campbell and Lamar (1989) observed that
“this species remains an enigma; there are few literature
 24 ANNALS OF CARNEGIE MUSEUM
references to it, and, in spite of a large type series, varia-
tion is unreported.” At that time, even an approximation of
maximum size was unavailable (Campbell and Lamar
1989 reported a total length of 665 mm for a female
paratype, UTA 18689).
Specimen NK 518 gave birth to one undeveloped and
still-born embryo and 19 live neonates having snout-vent
lengths of 177–240 mm (229 ± 14, 19).
Crotalus Linnaeus, 1758
Snakes of the genus Crotalus are small to large viperids
characterized by presence of a rattle, a moderate (e.g.,
Crotalus stejnegeri) to extremely short tail, absence of a
nasal pore, position of the facial pit below the naso-ocular
line, and entire subcaudals (Burger 1971; Gutberlet and
Harvey 2002). The dorsal head plates are heavily fractured
in all species except the most basal member of the clade
(Murphy et al. 2002), Crotalus ravus, which retains the
colubrid arrangement of head plates.
Content.—Crotalus contains at least 27 species (Murphy
et al. 2002), all occurring in North America except
C. durissus.
Crotalus durissus Linnaeus, 1758
[Crotalus] Durissus Linnaeus, 1758:214. HOLOTYPE (apparently lost;
originally in the Claudius Grill Collection and later sent to the
ZMUU according to Klauber 1941) from “America” (restricted to
“Jalapa, Veracruz, Mexico” by Smith and Taylor 1950).
Description.—Our sample contains relatively few adults,
and we do not report sexual differences in morphometric
characteristics. Excluding the rattle, the tail is 7–10% (8 ±
1, 12) as long as snout-vent length. The head is 64–92%
(78.4 ± 8.7, 14) as wide as long and accounts for 3.1–5.4%
(4.8 ± 0.7, 12) of snout-vent length. Eye diameter accounts
for 15.6–21.1% (17.9 ± 1.8, 15) and eye-nostril distance
accounts for 21.8–24.8% (23.7 ± 1.0, 15) of head length.
On the snout, the internasals are in contact medially
and, in specimens we examined, scales were never present
between them or between the internasals and rostral. The
canthals are in contact medially and usually followed by
two prefrontals and 2–3 intersupraoculars (four prefrontals
and four intersupraoculars are present in AMNH 104580).
Behind each supraocular is a large postsupraocular plate
(rarely divided). Unlike other Bolivian pitvipers, the upper
preocular is entirely below the canthus so that only the
internasals and canthals form the canthus. Two loreals are
usually present, and both vary in shape moreso than the
loreal in Bothrops. The upper loreal is usually oval and
smaller than the lower loreal; it separates the postnasal
from the upper preocular and is fused to the canthal about
one-third of the time. Rarely (9%, 17), the upper loreal is
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VOL. 74
divided vertically so that two scales separate the nasal and
upper preocular. The lower loreal resembles the loreal of
most Bothrops in being subtriangular and as tall as long or
taller than long. Relative to other Bolivian pitvipers, the
prelacunal is large and contributes more than one-half of
the ventral margin of the pit. This scale is separated from
the second supralabial by one scale, although a portion of
it separates the subfoveals and contacts the third supralabi-
al about one-half of the time (46%, 20). A lacunolabial is
not present. The sublacunal is entire and does not contact
the supralabials in any specimen. Not counting the scale
between the prelacunal and supralabials, this species has
1–5 (3 ± 1, 20) prefoveals. The prenasal contacts the first
supralabial in all specimens. Elongate suboculars were
absent in most specimens (MCZ 7491 being the only
exception); instead 1–5 (3 ± 1, 20) suboculars and two
postoculars are present, and all the oculars below and
behind the eye tend to be subequal. The row of suboculars
is separated from the supralabials by 1–3 scales [=3–5, 4 ±
1 (20) interoculabials]. This species has 11–16 (14 ± 1, 20)
supralabials and 14–18 (16 ± 1, 20) infralabials; more
infralabials (3–5, 4 ± 1, 20) contact the genials than is
common in other Bolivian pitvipers. Medial contact of the
first pair of infralabials separates the mental from the chin-
shields. In three specimens (15%, 20), the first pair of
infralabials are divided to form a second pair of chin-
shields. The chinshields are separated from the ventrals by
4–8 (6 ± 1, 20) gulars and 1–3 (2 ± 1, 20) preventrals.
Males have 171–181 (176 ± 5, 3) ventrals and 23–30 (26
± 4, 3) subcaudals, and females have 169–184 (177 ± 6, 5)
ventrals and 20–29 (24 ± 4, 5) subcaudals. All but the 0–3
most distal subcaudals are divided. In our sample, 27–31
(29 ± 1, 16) dorsals at midbody reduce to 19–22 (21 ± 1,
16) one head-length in front of the vent. This species has
27–34 (31 ± 2, 17) interrictals.
In our sample, the dorsal surface of the head bears a
well-defined black band across the raised posterior mar-
gins of the prefrontals, the anterior one-half of the frontals,
and the anterior one-third of the supraoculars. The rostral
is brown with a distinct cream margin that contrasts
sharply against the brown nasal and internasal scales. Two
stripes are present on the face. A brown postlacunal stripe
extends from the sublacunal to the dorsal edges of, at least,
supralabials 3–4. The postocular stripe is brown and
1.5–3.5 temporals high. It becomes broken into one or two
irregular blotches after crossing the rictus. Except for scat-
tered, diffuse blotches and except where they are over-
lapped by facial stripes, the supralabials, infralabials, chin-
shields and gulars are cream. The mental is pale brown,
and this pigment usually extends onto the flanking first
pair of infralabials.
Three sets of stripes are present on the neck. The par-
avertebral stripes are three scales wide, the two outer
scales being a darker brown than the middle row of scales.
These stripes begin medial and just behind the supraocu-
lars, are always more than twice as long as the head, and
 2005 HARVEY ET AL.—BOLIVIAN PITVIPERS
Fig. 12.—Distribution of Crotalus durissus in Bolivia. Areas above 500
and 1000 m are shaded gray.
extend to the level of the 21–36th ventral. In occasional
specimens, a dark brown supraocular band connects the
anterior edges of the paravertebral stripes. Pale brown
accessory stripes are separated from the paravertebral
stripes by about three dorsals. These stripes are broken and
indistinct. Ventrolateral stripes are pale brown and two
scales wide; they do not include the paraventrals which are
cream. The ventrolateral stripes extend to the level of the
17–25 ventral and are shorter than the paravertebral
stripes.
Twenty-one to twenty-seven (24.0 ± 1.6, 16) diamond-
shaped blotches are present on the body, the last two or
three becoming increasingly “band-shaped” and poorly
defined. Except for these last two or three, the vertebral
diamonds are edged by a row of silver scales, then by a
row of dark brown scales. The scales in the center of the
diamonds are pale brown and about the same as the ground
color on the flanks. The paraventral diamonds are less dis-
tinct, because they lack the dark brown scales present in
the vertebral pattern. In some neonates (e.g., UTA
34566–67), five bands are visible on the tail; however, the
caudal two-thirds of the tail are usually much darker (black
in adults) than the base so that 2–3 bands are usually all
that are visible.
Distribution.—BOLIVIA. BENI. Gral. José Ballivián:
Estancia Alianza, “sabana entre Río Matos y Río
Curiraba” (CBF 389); El Totaizal (CBF 752). Mamoré:
“Río Mamoré, Puerto Siles” (AMNH 101911); “Estancia
Yutiole, ca 20 km S San Joaquín,”=Estancia Yutiola
(AMNH 104580–82). Moxos: Bocorondo, "siguiendo el
curso del Río Mamoré, 5 km Río abajo de Exaltación”
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25
(CBF 894). Yacuma: Espíritu, “bosque de galería entre
Arroyo Capibara y Dolores” (CBF 258 and CBF 362).
Unknown: “Río Mamoré, ca 23 km W San Javier,” bank
of river not specified, Cercado or Moxos province
(AMNH 101910). CHUQUISACA. Luis Calvo: “30 km
SE Carandaiti”=Carandayti (LACM 37669). LA PAZ.
Sud Yungas: Yanacachi, approximately 1800 m (CBF
504, 819); Taca, Laguna Chacajahuira (CBF 1699).
SANTA CRUZ. Andrés Ibáñez: Santa Cruz de la Sierra
(CM 118, 124); San Lorenzo “a 15 km de Cotoca” (NK
443). Chiquitos: 135 km N Pozo del Tigre, Empresa
ABFA, 17°34'48"S 61°71'33"W (NK 726); Finca Dos
Milanos (UTA 34566–70, 38040–44); Serranía de
Santiago, 800 m (FMNH 195925). Cordillera: Serranía
Parabanó (NK 974); “La Brecha, ca 104–120 km NE
Charagua, Izozog Region,” (AMNH 141446). Florida:
Pampagrande (NK 561, NK 626, 1267–68, 1957); Mairana
(NK 951), Mataral (NK 2150). Germán Busch: Estancia
Arco Iris cerca Laguna Cáceres (CBF 2115, NK 1148–49);
“Puerto Suárez, Laguna Cáceres, Tamarinero” (NK 1190).
Ichilo: Buena Vista (CM 2924); San Carlos (NK 246).
Sara: specific locality unknown (CM 30, 63).
UNKNOWN. MCZ 7491 (“Beni River Valley” (MCZ
7491)
Specimens not examined: BOLIVIA. LA PAZ.
“Campolican” (BMNH, Boulenger 1896. This locality
probably is a misspelling of Caupolican, the former name
of the province now known as Abel Iturralde). SANTA
CRUZ. Velasco: “Comunidad ‘La Florida,’” 14°35'S,
60°50'W (EBD, no museum number, De la Riva et al.
1992.) (Fig. 12).
Remarks.—In number of described subspecies, Crotalus
durissus rivals Bothrops neuwiedi, and, as for the latter
species, we suspect that many of these names will not sur-
vive careful scrutiny. Nonetheless, the status and mecha-
nisms of differentiation among the forms currently
referred to C. durissus remain some of the most interesting
problems in Neotropical herpetology (Vanzolini and
Calleffo 2002). Twelve to fourteen subspecies have been
recognized recently (Hoge 1966; Campbell and Lamar
1989; Harris and Simmons 1972a,b; McCranie 1993).
Most subspecies have been defined on the basis of color
pattern with little attention being paid to variation in squa-
mation (but see McCranie 1993).
Bolivian specimens have been referred to Crotalus
durissus terrificus (Harris and Simmons 1972a; Peters and
Orejas-Miranda 1970), although Hoge (1966) thought that
C. d. collilineatus probably occurs in Bolivia. In adjacent
Brazilian states, Nascimento et al. (1988) referred speci-
mens from the Chapada dos Guimarães, Mato Grosso, to
C. d. collilineatus, whereas Jorge da Silva (1993) referred
specimens from Vilhena (Rondônia) and Barracão
Queimado (Mato Grosso) to C. d. terrificus. Recent revi-
sors (Harris and Simmons 1972a; McCranie 1993) show a
gap between these subspecies along the border between
 26 ANNALS OF CARNEGIE MUSEUM
Brazil and Bolivia, however, Jorge da Silva (1993)
remarks that the two subspecies are sympatric over a large
area of central Brazil. Populations in the Serranía de
Huanchaca (Harvey 1998) and a C. durissus in the EBD
(De la Riva et al. 1992) from La Florida bridge published
gaps between the two subspecies in eastern Bolivia.
Specimens examined by us key (Peters and Orejas-
Miranda 1970; Harris and Simmons 1972 b) to
C. d. collilineatus rather than C. d. terrificus because (1)
scales in the centers of their rhomboid blotches are pale
brown, similar to the ground color and conspicuously con-
trasting with the dark brown scales edging the blotches
[vs. “centers of rhomboid blotches ‘not light,’ similar to
ground color, only slightly lighter than rest of blotch” in
C. d. terrificus (Harris and Simmons 1972a, p. 34)] and (2)
three sets of distinct stripes are present on the neck, the
accessory stripes being short, broken, and indistinct [vs.
“dorsal head pattern indistinct” in C. d. terrificus (Harris
and Simmons 1972a, p. 34)].
Assignment of the Bolivian sample to subspecies is
problematic. We note that ventral and subcaudal counts for
a few specimens in our study fall outside published ranges
(McCranie 1993) for both Crotalus durissus collilineatus
and C. d. terrificus. Nonetheless, our data fall within
ranges for the species as a whole.
Lachesis Daudin, 1803
Description.—The bushmasters are defined by a suite of
morphological synapomorphies including unusually high
scale counts (e.g., ventrals and dorsals), extensively divid-
ed distal caudals, oviparity, diamond-shaped dorsal blotch-
es, and tuberculate dorsals. Defining characteristics of this
genus were summarized by Campbell and Lamar (1989),
Werman (1992, 1999), and Gutberlet and Harvey (2002).
The rostral is usually wider than high and canthorostrals
are generally absent. A lacunolabial may be present or
absent. One or more small scales usually separate the
internasals and most of the dorsal head is covered in small,
smooth or obtusely keeled scales. Nine to fifteen scales
separate the supraoculars. Two loreals are present, the
supralacunal is generally discrete from the middle preocu-
lar, and the upper preocular is entire. The loreal pit is
crossed by the naso-orbital line and a nasal pore is present.
Bushmasters have 8–10 supralabials, 12–17 infralabials,
31–39 dorsal scale rows at midbody, 200–247 ventrals,
and 43–62 mostly divided subcaudals. The distal caudals
are finely divided into small spine-like scales.
Bushmasters are tan dorsally with dark brown
diamond-shaped dorsal blotches. The head is black or tan
with black markings. A dark postocular stripe extends to
the rictus or below the angle of the jaw. The venter is
immaculate cream.
Campbell and Lamar (1989) provide a recent descrip-
tion of the hemipenes of bushmasters: The hemipenes
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VOL. 74
extend about 9 subcaudals and are divided for about two-
thirds of their length. The sulcus spermaticus bifurcates at
the level of subcaudals 2–3. The proximal two-thirds to
three-fourths of each lobe is covered in moderate-sized
spines. Distally, the lobes are attenuate and calyculate.
Ridges of the calyces are smooth.
Werman (1992) and Gutberlet and Harvey (2002) list
characters of the skull of Lachesis. The following account
based on Werman’s data summarizes characters that distin-
guish Lachesis from other Neotropical pitvipers, especial-
ly Bothrops. The lateral margin of the prefrontal is convex
and smooth in dorsal view. The postorbital is large and
contacts the frontal. Dorsally, the frontals are flat without
elevated margins, and their width is greater than that of the
basicranium. In dorsal view, the parietal is triangular, with
concave postero-lateral margins. The medial wall of the pit
cavity is strongly developed. The ectopterygoid is longer
than the base of the pterygoid. In dorsal view, the anterior
portion of the ectopterygoid is broad or truncate. Where it
articulates with the pterygoid, the base of the ectoptery-
goid is not spatulate and possesses a short, posteriorly
directed, finger-like projection. The shaft of the ectoptery-
goid is flat, broad, and not tapering posteriorly; a single pit
is present on its postero-lateral surface where the
ectopterygoid retractor muscles attach.
Content.—Until recently, bushmasters were thought to
comprise a single, widespread polytypic species (Peters
and Orejas-Miranda 1970; Hoge and Romano-Hoge 1978;
Campbell and Lamar 1989). Using morphological, behav-
ioral, and biochemical evidence, Zamudio and Greene
(1997) recognized three species: Lachesis melanocephala
(Solórzano and Cerdas) and L. stenophrys (Cope) in
Central America and Northwestern South America and
L. muta (Linnaeus) in South America.
Remarks.—Lachesis has been characterized as having a
lacunolabial by some recent authors (e.g., Campbell and
Lamar 1989; Werman 1992) when, in fact, fusion of a
supralabial and the prelacunal is polymorphic (at high fre-
quencies in some populations) in both L. stenophrys
(Gutberlet and Harvey 2002) and L. muta (this study).
Lachesis muta Linnaeus, 1766
[Crotalus] mutus Linnaeus, 1766:373. HOLOTYPE from “Surinami”
(NHRM according to Andersson 1899).
Description.—Lachesis muta is the largest pitviper in
Bolivia. The largest specimen is a male (CBF, tag lost)
with a total length of 2319 mm (snout-vent length 2102
mm). Compared to Bothrops, the tail of bushmasters is rel-
atively short, being 10–12% (11 ± 1, 4) as long as snout-
vent length in males and 9–11% (10 ± 1, 2) as long in
females. The head is 68.6–74.4% (71.9 ± 2.5, 5) as wide as
long and its length accounts for 2.9–5.5% (3.5 ± 9.7, 5) of
 2005 HARVEY ET AL.—BOLIVIAN PITVIPERS 27

snout-vent length. The eye is moderate, its diameter being 69 64 59

39.2–51.1% (43.0 ± 4.8, 5) as large as eye-nostril distance. 10 10

The distance from the anterior corner of the eye to the cen-
ter of the nostril accounts for 27.2–30.7% (29.1 ± 1.1, 6)
of head length.
Unlike most specimens of Bothrops, the rostral is usual-
ly shorter than wide in this species. Greatest rostral height
is 85.4–100.0% (90.6 ± 5.7, 5) of rostral width.
Canthorostrals are absent in all except one specimen (8.3%, 15 15

22); AMNH 101912 has a single canthorostral on one side.

The internasals are smooth, small, and oval-shaped scales
widely separated by 2–3 flat scales behind the rostral. The
internasals do not contribute to the canthus as is the case in
Bothrops. The canthus is formed by the postnasal and three
or four smooth canthals separating the postnasal from the
supraocular. The anteriormost canthals are separated by 20 20

6–12 (9 ± 1, 13) flat, rounded scales. Intersupraocular

fusions or “plate-like scales” are absent and the smooth
supraoculars are separated by 10–13 (12 ± 1, 14) smooth,
juxtaposed and rounded or polygonal scales. 200 km

This species has 1–6 (4 ± 1, 9) prefoveals. Two loreals

69 64 59
are present in all specimens and the lower loreal is
82.6–111.5% (91.8 ± 10.7, 7) as high as long. Shape of the Fig. 13.—Distribution of Lachesis muta in Bolivia. Open symbols repre-
sent published reports based on specimens not examined by us. Areas
lower loreal is irregular being square to subtriangular to above 500 and 1000 m are shaded gray.
tear-drop shaped. The upper preocular does not contribute
to the canthus or extend onto the dorsal surface of the
dals project away from the axis of the tail and give it a
head. In our sample, this scale is not divided and the mid-
bristly appearance. In contrast to Bothrops, interrictal
dle preocular does not fuse to the supralacunal. The lower
counts (27–31, 28.4 ± 1.3, 10) were 3–6 scales lower
preocular has been lost or is fused to the sublacunal which
rather than higher than counts at midbody. Thirty-two to
enters the orbit. The sublacunal region is quite variable in
thirty-five (32.7 ± 1.5, 10) dorsals at midbody reduce to
our sample. Lacunolabials are present in 50% and the
23–25 (24.4 ± 0.9, 9) one head-length in front of the vent.
prelacunal and supralabials are in contact in 50% of the
In Bolivian specimens, the dorsal ground color is tan
specimens. When the prelacunal and supralabials are in
with dark brown to black markings. The rostral, facial
contact, the second supralabial is frequently divided verti-
scales, scales below the eye, labials, chin, and most ven-
cally. An uncatalogued female from La Paz is unique in
trals are immaculate. The internasals, canthals, and
lacking subfoveals on one side. In the other specimens, the
supraoculars are also immaculate; however, the dorsal sur-
sublacunal contacts the third or fourth supralabial and one
face of the head between these scales bears irregular black
or two subfoveals are at the juncture of the sublacunal,
blotches. The dark brown to black postocular stripe is nar-
supralabials and prelacunal (65%, 26), or the sublacunal is
row (1–1.5 temporals wide) and does not cross any of the
completely separated from the supralabials by one row of
supralabials although it overlaps the scales directly above
subfoveals (35%, 26).
the rictus and extends obliquely for as many as seven scale
AMNH 6775 is the only specimen (9%, 22) with a sin-
rows behind the rictus. Two rows of temporals above the
gle subocular on both sides; the other specimens (91%, 22)
postocular stripe are immaculate. A pair of large dark
have 2–3 suboculars and 2 postoculars. Our specimens
brown to black blotches on the neck are followed by 25–33
have four (42%, 24) or five (58%, 24) interoculabials,
(27 ± 3, 11) diamond-shaped blotches on the body. The
eight (18%, 22), nine (73%, 22), ten (5%, 22), or eleven
blotches fail to reach the paraventrals anteriorly but do so
(5%, 22) supralabials, and 13–16 (14 ± 1, 24) infralabials.
near the vent and may extend onto the edges of the most
The first two (79%, 24) or first three (21%, 24) infralabi-
posterior ventral scales. Nearing the vent, paler brown
als contact the chinshields. Five to nine (7 ± 1, 12) gulars
blotches fill most of the centers of the interspaces on the
and 2–6 (3 ± 1, 12) preventrals separate the chinshields
flanks, but these marks are absent anteriorly. Short tan
from the ventrals. Males have 215–230 (223 ± 6, 6) ven-
interspaces usually one to two scales long separate 5–7 (6
trals and 49–52 (49 ± 2, 6) subcaudals, and females have
± 1, 10) dark brown to black bands on the tail. These bands
218–225 (221 ± 4, 3) ventrals and 47–62 (53 ± 8, 3) sub-
extend ventrally onto the subcaudals, but are darkest dor-
caudals. Thirty-three percent (9) of the specimens had one
sally. Distally and ventrally, three quarters of the elongate
or two entire subcaudals near the vent; the last 13–20 sub-
terminal spine are yellow and sharply contrast with the
caudals are doubly divided. The doubly divided subcau-
brown base of the scale and the adjacent subcaudals.

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 28 ANNALS OF CARNEGIE MUSEUM
Distribution and Comparative Material.—Although this
species was first reported from Santa Cruz by Griffin
(1916), maps in recent publications (Campbell and Lamar
1989; Zamudio and Greene 1997) have shown the species
only in northern provinces of the country. Our studies and
research by Visinoni (1995) reveal a much broader distri-
bution of bushmasters in Bolivia than previously thought
(Fig. 13). Based on several reliable reports, Harvey (1998)
included this species in the fauna of Parque Noel Kempff
Mercado, Velasco, Santa Cruz.
BOLIVIA. BENI. Gral. José Ballivián: Bosque
Chimanes (CBF 1118); “El Totaizal, orilla del Río Matos,
40 km E. San Borja (CBF, uncatalogued).” Marban:
Estancia Totaí, 15°32'40"S, 64°17'05"W (NK 1958). Vaca
Díez: “Río Mamoré, Guayaramerín,” (AMNH 101912).
Unknown: specific locality unknown (AMNH 6775).
COCHABAMBA. Chapare: Villa Tunari (FML 2403).
LA PAZ. Abel Iturralde: Tumupasa (CBF 1934); Alto
Madidi (CBF 1936); “Camino a Tumupasa, propiedad la
Magdalena a 11 km de San Buena Ventura” (CBF 740);
“Entre aserradero Moira e Ixiamas” (CBF 742). SANTA
CRUZ: Andrés Ibáñez: Santa Cruz de la Sierra (CM 125,
2847, FML 642). Ichilo: Buena Vista (AMNH 36010).
Ñuflo de Chávez: La Trancas (NK 628).
Specimens not examined: BOLIVIA. BENI. Gral.
José Ballivián: “48 km E de San Borja” (EBD, no muse-
um number given, De la Riva et al. 1992). SANTA CRUZ.
Sara: “Estancia ‘La Ponderosa,’ Santa Rosa de Sara, Prov.
Gutierrez” (EBD, no museum number given, De la Riva et
al., 1992). UNKNOWN. “Bolivia” (BMNH, Boulenger
1896).
A. Visinoni (1995) reported locality information for 33
specimens of Lachesis muta in the NK, BMNH, the Museo
Universidad Técnica del Beni, Trinidad, Beni, Bolivia
(MUTB), and the Estación Biológica Doñana, Sevilla,
Spain (EBD, specimens also mentioned by Fugler and De
la Riva 1990), the author’s private collection, and unspec-
ified Bolivian collections. Although we probably exam-
ined the same specimens as Visinoni in the NK and CBF,
he did not provide museum numbers and we cannot be
sure that our studies are based on some of the same mate-
rial. His specimens come from the following localities:
BOLIVIA. BENI. Cercado: Casarabe, 14°50'S, 64°28'W.
Gral. José Ballivián: Estación Biológica del Beni, 14°49'S,
66°51'W (EDB). Iténez: El Alba, Río Blanco, 12°50'S,
64°00'W. Moxos: Tío Tijamuchi, 14°50'S, 65°12'W
(MUTB). COCHABAMBA. Carrasco: Bulo Bulo,
17°13'S, 64°21'W; Yvirgasama, 17°02'S, 64°52'W.
Chapare: Aroma, 16°35'S, 65°47'W; Chipiriri, 16°50'S,
65°20'W; Puerto San Francisco, 16°45'S, 65°20'W. LA
PAZ. Iturralde: San Buenaventura, 14°30'S, 67°35'W
(CBF). PANDO. Nicolás Suárez: Cobija, 11°02'S
68°44'W. SANTA CRUZ. Andrés Ibáñez: El Tomo,
18°00'S, 63°30'W; Terevinto, 17°43'S, 63°23'W; Potrerillo
del Guenda, 17°50'S, 63°36'W. Guarayos: Puerto
Almacén, 15°46'S, 62°15'W (EBD); Yotaú, 16°10'S,
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VOL. 74
62°50'W. Ichilo: Caranda, 17°33'S, 62°32'W (NK); “3 Km
antes del puente río Ichilo,” 17°20'S, 64°20'W; “Puente río
Palometillas,” 17°14'S, 64°20'W; San Germán, 17°20'S,
64°00'W; S. Salvador (Yapacaní), 17°25'S, 63°49'W; San
Rafael de Amboró, 17°36'S, 63°36'W; Buena Vista,
17°27'S, 63°40'W (BMNH). Ñuflo de Chávez:
Perseverancia, 14°45'S, 62°35'W. Obispo Santistevan:
Chane, 17°05'S, 63°10'W. Sara: Santa Rosa del Sara,
17°07'S, 63°35'W. Velasco: "35 Kms Oeste de S.
Ignacio,"16°30'S, 61°05'W (EBD).
Species Not Known from Bolivia
Presumably based on Kempff-Mercado (1975), Campbell
and Lamar (1989) reported Bothrocophias hyoprora from
Bolivia (see also Fugler and De la Riva 1990). This species
may occur in Pando or along the humid Andean foothills;
however, voucher specimens from Bolivia do not exist.
Kempff-Mercado’s report may have been based on juve-
nile B. microphthalmus, which are very similar to
B. hyoprora.
Bothrops brazili was reported for Bolivia by Kempff-
Mercado (1975) and is included in checklists for the coun-
try (Campbell and Lamar 1989; Fugler and De la Riva
1990; Fugler et al. 1995). It was not, however, reported for
Bolivia in its original description (Hoge 1954). This
species’ presence in Bolivia needs to be confirmed. The
species is known from Madre de Dios, southern Peru
(Morales and McDiarmid 1996) and Rondônia, Brazil
(Vanzolini 1986) and almost certainly occurs in the rain-
forest of northeastern Santa Cruz, Pando, and La Paz.
Nonetheless, we are aware of no vouchered material from
within the country’s borders. USNM 123975 appears to be
a specimen of Bothrops brazili and was collected by
Raymond Gilmore from “Bolivia” without other data.
However, a typed note was placed in the jar of this speci-
men by Alphonse R. Hoge dated 4 April 1968 and states
that the locality should read Brazil.
A snake originally identified as Bothrops sanctaecrucis
was collected in the vicinity of Porto Velho, Rondônia,
Brazil. This specimen (USNM 48931) is in poor condition
and represented by a head only. Like other B. brazili and
unlike all but one B. sanctaecrucis we examined, it pos-
sesses three scales between the canthals and has 7/7
supralabials (vs. 8–9 infralabials in B. sanctaecrucis). We
suspect that it is actually a specimen of B. brazili from yet
another locality near the Bolivian border.
A single specimen of Bothrops jararacussu is currently
alive at the zoo in Santa Cruz de la Sierra, Santa Cruz,
Bolivia. Unfortunately, the zoo staff has no precise locali-
ty information for the specimen, although they suspect that
it came from Brazil, possibly brought by P. Bettella, who
visited the Instituto Butantan. On the other hand, in corre-
spondence with W.W. Lamar, Bettella claimed that he had
a Bolivian specimen of B. jararacussu; again no precise
 2005 HARVEY ET AL.—BOLIVIAN PITVIPERS
data were given although poor photographs of the speci-
men were sent to Lamar. Without question, the snake in the
photo is B. jararacussu. Bettella routinely deposited
snakes from his personal collection in the Museo Noel
Kempff Mercado. However, this museum does not and
never has had a specimen of B. jararacussu.
Two specimens in the Carnegie Museum were identi-
fied as Bothrops jararacussu by Amaral (1925 and 1926)
after Griffin (1916) originally referred them to B. lanceo-
latus. Unfortunately, both specimens along with two
paratypes of B. sanctaecrucis were taken to Brazil by
Alfonse Hoge in the 1960s. Although R. Crombie subse-
quently was able to retrieve some of these specimens,
CM 43 and CM 121 are still (as of June 2000) missing
from the collection (S. Rogers, personal communication).
At the same time Griffin (1916) examined CM 43 and CM
121, he also examined CM 313 and also referred it to
B. lanceolatus. Amaral (1926) referred CM 313 to
B. atrox; however, CM 313 (now MCZ 17699) is Bothrops
sanctaecrucis. Counts of scale rows at midbody (23),
number of ventrals (181, 185), number of subcaudals (60,
66), number of supralabials (8), number of intersupraocu-
lars (7, 6) and scales between the subocular and labials (1,
1) recorded by Griffin (1916) for the lost specimens fall
within the known ranges of both B. jararacussu and
B. sanctaecrucis. These specimens are almost certainly
B. sanctaecrucis because they were collected near Santa
Cruz de la Sierra (CM 43 from “Provincia del Sara” and
CM 121 from “Santa Cruz de la Sierra”). Especially in
recent years, this area of Bolivia has been heavily collect-
ed, and it is inconceivable to us that B. jararacussu would
have escaped detection. To our knowledge, no Bolivian
specimens of B. jararacussu exist in museums. The
species’ occurrence in Bolivia has been reported by
Fig. 14.—Male lectotype (BMNH 1946.1.19.51) of Bothriopsis
chloromelas from Huancabamba, Peru.
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29
Amaral (1930b), Hoge (1966), Hoge and Romano-Hoge
(1981), Fugler and De la Riva (1990), Campbell
and Lamar (1989), Cei (1993), and Fugler et al. (1995).
During the course of this study, we found specimens of
B. sanctaecrucis misidentified as B. jararacussu in sever-
al major U.S. museums (e.g., AMNH, MCZ, CM).
Above, we clarified the identity of Bothriopsis oligolepis.
During the course of our investigation into the status
of this species, we examined all specimens of Bothriopsis
chloromelas in the United States. We take this opportunity
to redescribe this poorly known species and designate a
lectotype.
Bothriopsis chloromelas (Boulenger, 1912)
Figure 14
Lachesis chloromelas Boulenger, 1912:423. 3 SYNTYPES (BMNH
1946.1.17.66 [formerly 1911.12.13.63], 1946.1.19.51, and
1946.1.19.52) from “Huancabamba, E. Peru, above 3000 ft.”
Lectotype [designated herein] male (BMNH 1946.1.19.51, formerly
1912.11.1.10) [examined].
Lachesis lanceolatus: Griffin, 1916:83 (not of Lacépede)
Lachesis peruvianus: Griffin, 1916:226 (not of Boulenger).
Bothrops chloromelas (Boulenger): Amaral, 1926:320; Amaral,
1930b:235.
Bothrops chrysomelas (Boulenger): Amaral, 1926:320 [misspelling].
Lachesis peruvianus: Werner, 1927:256 (not of Boulenger).
Lachesis chloromelas Boulenger: Amaral, 1930a:58 [Amaral thought that
this species may be a “variety” of B. peruviana].
Bothrops chloromelas (Boulenger): Amaral, 1930b:235; Pifano and
Römer, 1949:294; Meneses, 1974:70.
Bothrops oligolepis (Werner): Parker, 1934:273; Dunn, 1946:19 (part);
Hoge, 1966:131 (part); Peters and Orejas-Miranda, 1970:53 (part);
Duellman, 1979:456 (part); Hoge and Romano-Hoge, 1981:217
(part); Carrillo de Espinoza, 1983:17; David and Ineich, 1999:233
(part). [These and subsequent authors thought that Werner’s name
oligolepis properly applied to B. chloromelas and not B. peruvianus].
Bothriopsis oligolepis (Werner): Campbell and Lamar, 1989:174 (part);
Carrillo de Espinoza and Icochea, 1995:20 (part); McDiarmid et al.,
1999:249 (part).
Bothriechis oligolepis (Werner): Schätti and Kramer, 1993:244 (part).
[These authors considered B. chloromelas, B. oligolepis, and B. pul-
chra to be a single species.].
Bothriechis oligolepis oligolepis (Werner): Golay et al., 1993:37 (in part,
also includes B. oligolepis).
Diagnosis.—Bothriopsis chloromelas is distinguished
from all congeners and from species of Bothrocophias and
Bothrops by the following combination of characters: (1)
lacunolabial present; (2) dorsal keels nontuberculate; (3)
infralabials yellow with small black spots; (4) canthoros-
trals absent; (5) anteriormost 5–25 subcaudals entire; dis-
tal subcaudals divided; (6) ventrals 183–204 and subcau-
dals 49–64; (7) loreal subtriangular; (8) canthals narrow;
(9) postocular stripe black and 2–3 temporals high; (11)
head black with yellow lines forming circular patterns or
head yellow with heavy black stippling and marking;
supralabials green, heavily speckled in black; dorsum
banded and lichenose.
 30 ANNALS OF CARNEGIE MUSEUM
Description.—The largest specimen of Bothriopsis
chloromelas is a female (LSUMZ 41037) with a total
length of 948 mm; the largest male specimen is probably
FMNH 59205. Although its snout-vent length cannot be
measured because only the skin of the body is available,
the FMNH specimen appears to be about the same size as
the LSUMZ specimen. The tail is 15–19% (16 ± 2, 3) of
snout-vent length in females and 16% of snout-vent length
in a male specimen. The species’ head is triangular and
strongly distinct from its neck. The head is longer than
wide, its width being 70–79% (74 ± 4, 4) of its length, and
it accounts for 4.0–4.6% (4.3 ± 0.3, 4) of snout-vent
length. Larger scales on the dorsal surface of the head are
smooth. All but one specimen have large, flat plates
appearing to have resulted from the fusion of three to five
scales and interspersed with ordinary keeled scales
between the supraoculars and in the parietal region. In
MCZ 156347 only a single “doubled” scale lies between
the supraoculars and it is doubly keeled rather than flat.
The dorsal surfaces of the internasals, canthals, and upper
preoculars are smooth. The supraoculars are rugose, about
half as wide as long (45–51%, 48.4 ± 2.5, 5), roughly rec-
tangular, and separated medially by 5–8 (6 ± 2, 6) scales.
Two specimens (BMNH 1946.1.19.51 and MCZ 156347)
have a single, greatly thickened keel running the length of
each of their supraoculars. Scales just behind the inter-
nasals are large and may be flat or keeled. Twenty-six to
twenty-eight interrictals (27 ± 1, 6) are present.
Thickened, central portions of the internasal, canthal,
and upper preocular form a sharp and raised canthus ros-
tralis. The rostral is about as high as wide and about as
wide as the mental. The loreal is 73.7–94.1% (80.6 ± 8.1,
5) as high as long; it broadly contacts the canthal and pre-
ocular. The foveal pit is bound by entire upper and lower
lacunals and the fused (not fused but in contact on one side
only in LSUMZ 41037) second supralabial and prelacunal.
This species has 1–4 (3 ± 1, 6) prefoveals and lacks small
scales between the anterior nasal and rostral. The eye-
nostril distance is 24.4–27.9% (25.8 ± 1.5, 5) of head
length. Suboculars are separated from the supralabials by
a row of keeled scales (=3 interoculabials, 100%, 6), and
the subocular is elongate and jagged. In MCZ 156347, the
subocular is fused to the middle preocular on both sides of
the head. Three preoculars are present, the upper subrec-
tangular and about four times as large as the square to rec-
tangular middle preocular; the small lower preocular is
excluded from the orbit by the suboculars in all specimens.
On both sides of the type specimen, the middle preocular
is fused to the lower lacunal. All specimens have two small
postoculars. Most supralabials are subtriangular and high;
all specimens have seven on both sides of the head (100%,
6). The foveal pit lies below an immaginary line drawn
between the eye and naris. A nasal pore is present.
Infralabials number 9–11 (10 ± 1, 6), the first 2–3 (3 ±
1, 5) contacting the chinshields. The chinshields are about
twice as long as wide and are separated from the mental by
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VOL. 74
medial contact of the first pair of infralabials. The mental
groove is shallow. Four to six (5 ± 1, 5) gulars and 1–3 (2
± 1, 5) preventrals separate the chinshields from the ven-
trals. The paraventrals and parasubcaudals are keeled,
although less heavily than the dorsals. Female Bothriopsis
chloromelas have 185–204 (193 ± 10, 3) ventrals, 49–64
(56 ± 8, 3) subcaudals, and two males have 183–184
ventrals and 54–62 subcaudals. Five to twenty-five (12 ±
8, 6) proximally located subcaudals are entire. Usually
(83%, 6), 23 dorsals reduce to 19 one head-length in front
of the vent; FMNH 59205 has 25 reducing to 21 dorsals.
The terminal spine is blunt, elongate, and slightly upturned
distally.
Bothriopsis chloromelas bears what may arguably be
the most ornate color pattern of any pitviper. The dorsal
surface of the head and upper sides of the snout may be
almost entirely black (e.g., in BMNH 1946.1.19.51 and
LSUMZ 41037) with many larger scales being marked by
yellow lines forming hollow circular patterns. On some
specimens (e.g., FMNH 59205), wide black stripes form a
V-shaped mark edging the supraoculars; considerably
more dorsal head scales are yellow, although all are heav-
ily speckled and blotched in black. The postocular stripe is
immaculate black and about two and a half temporals
wide. It occupies the upper portions of the last three
supralabials and usually curves around the corner of the
mouth onto the gular scales. Both the supralabials and
infralabials are heavily speckled and blotched in black.
The supralabials are green, the infralabials yellow or yel-
low and green anteriorly and near the rictus. The gulars
and chinshields are immaculate yellow except in FMNH
59205, which bears some black specks on the chinshields.
The ventrals are yellow anteriorly and grade to green pos-
teriorly. Both the ventrals and subcaudals are heavily
speckled and blotched. Pale paraventral spots like those in
most forest pitvipers are just discernible, most prominent-
ly in AMNH 104298. The dorsal pattern consists of three
types of irregular bands. Two of these types are mostly
black: one band is shorter (2–4 dorsals long) and bears
many immaculate scales and few scales lightly marked in
yellow; the scales of the longer black band (3–6 dorsals
long) bear a complex yellow reticulation. These two types
of bands alternate along the flanks of this species and they
are separated by short (1–2 dorsals long) bands of green
scales heavily speckled and blotched with black. Both
types of black bands begin on the third dorsal row; they
may extend across the back or be staggered along the mid-
line. This same dorsal pattern extends onto the tail. The
posterior one-third to one-half of the tail is pink to yellow
in preservative (at least ventrally as is the case in MCZ
156347 and FMNH 59205).
Campbell and Lamar (1989, fig. 153) provided color
photographs of LSUMZ 41037 and noted that, in life, the
iris is black with faint paler smudges. Skin between the
dorsals is black.
 2005 HARVEY ET AL.—BOLIVIAN PITVIPERS
Specimens Examined and Distribution.—This species is
known from the Andes of northern and central Peru. One
specimen is known from the Serranía de Sira. In addition
to the departments for which we examined specimens,
Carrillo de Espinoza and Icochea (1995) report
Bothriopsis oligolepis from several additional depart-
ments. Some of these records may properly apply to
B. oligolepis (e.g., reports from Apurimac, Cuzco, and
Puno), whereas others (Junín) fill in gaps in the distribu-
tion we report here for Bothriopsis chloromelas.
PERU. HUÁNUCO: “Cerros de Sira, 1550 m”
(AMNH 104298), “prov. Leoncio Prado, Dtto. Valdizan,
La Divisoria” (MCZ 156347). JUNIN: “Chanchamayo,
Pulcalpa, 1300 m” (FMNH 59205. LORETO: “Tarma,
elev. 6,000 ft.” (CM 373), “Fundo Cinchoa, 72 km E of
Tingo María, 4600–5000 ft.” (USNM 119020). PASCO:
“Santa Cruz, ca. 9 km SSE Oxapampa” (LSUMZ 41037).
PIURA: “Huancabamba, E. Peru, above 3000 ft.”
(BMNH 1946.1.19.51, formerly 1912.11.1.10, lectotype).
Remarks.—Griffin (1916) misidentified CM 373 as
Lachesis lanceolatus on p. 83, then as Lachesis peruvianus
on p. 226. We confirm Amaral’s (1926) synonymization of
these names. This juvenile specimen is unquestionably
Bothriopsis chloromelas and not the superficially similar
B. pulchra.
ACKNOWLEDGMENTS
For loan of specimens under their care we thank
N. Gilmore (ANSP), L. Ford (AMNH), A.G. Kluge
(UMMZ), C.J. McCarthy (BMNH), H. Voris (FMNH),
R. Aguayo V. (Universidad Mayor de San Simon,
Cochabamba), R. Reynolds (USNM), D. Kizirian
(LACM), L. Trueb (KU), D.A. Rossman (LSUMZ),
J. Wiens (CM), J. Hanken (MCZ), and W. Böhme
(ZFMK). We were provided with working space at the
AMNH by D.R. Frost and L.S. Ford, at the CM by
J.J. Wiens, at the FML by J.G. Scrocchi, at the MCZ by
J.E. Cadle, at the USNM by R.P. Reynolds, and at UTA by
J.A. Campbell. I. De la Riva, D. Rumiz, and E.N. Smith
kindly donated photographs for this publication.
P. Strimple supplied us with photos and data from a speci-
men of B. moojeni. This research was partially funded by
research and travel grants from the ETSU College of Arts
and Sciences, Office of International Programs, and Office
of Research and Sponsored Programs. J.A. Campbell,
W.W. Lamar, I. De la Riva, and R.L. Gutberlet, Jr., read
earlier drafts of the manuscript, encouraged our research,
and provided many helpful suggestions.
LITERATURE CITED
ABALOS, J.W., AND E.C. BAEZ. 1963. Variaciones del diseño en Bothrops
neuwiedii meridionalis de Santiago del Estero. Acta Zoologica
Lilloana (Argentina), 19:479–486.
Downloaded From: https://bioone.org/journals/Annals-of-Carnegie-Museum on 09 Sep 2020
Terms of Use: https://bioone.org/terms-of-use Access provided by Pontifica Universidade Catolica do Rio Grande do Sul (PUCRS)
31
ABALOS, J.W., AND C.C. MISCHIS. 1975. Elenco sistemático de los ofidios
argentinos. Boletín de la Academia Nacional de Ciencias, Córdoba
(Argentina), 51:55–76.
AMARAL, A. DO. 1923. New genera and species of snakes. Proceedings of
the New England Zoological Club, 7:85–105.
———. 1925. On the differentiation of the species Bothrops atrox
(Linné, 1758), B. jararaca Wied, 1824), and B. jararacussu
Lacerda, 1884. Contributions from the Harvard Institute for Tropical
Biology and Medicine, 2:22–43.
———. 1926. Ophidia from South America in the Carnegie Museum: a
critique of Dr. L.E. Griffin’s “Catalog of the Ophidia from South
America at present (June, 1916) contained in the Carnegie
Museum.” Annals of Carnegie Museum, 16:319–323.
———. 1927. Studies of Neotropical Ophidia IV. A new form of
Crotalidae from Bolivia. Bulletin of the Antivenin Institute of
America, 1:5–6.
———. 1930a. (dated 1929). Estudos sobre ophidios neotropicos. XVII.
Valor systematico de varias formas de ophidios neotropicos.
Memórias do Instituto Butantan (Brazil), 4:1–68.
———. 1930b. (dated 1929). Estudos sobre ophidios neotropicos. XVIII.
Lista remissiva dos ophidios da região neotropica. Memórias do
Instituto Butantan (Brazil), 4:126–271.
ANDERSON, L.G. 1899. Catalogue of Linnean type-specimens of snakes in
the Royal Museum in Stockholm. Bihang till Kongliga Svenska
Vetenskaps-Akademiens. Handlingar (Sweden), 24:1–35.
AQUINO, A.L., N.J. SCOTT, AND M. MOTTE. 1996. Lista de anfibios y rep-
tiles del Museo Nacional de Historia Natural del Paraguay (marzo,
1980–setiembre, 1995). Pp. 331–400, in Colecciones de Flora y
Fauna del Museo Nacional de Historia Natural del Paraguay (O.R.
Martínez, ed.). Museo Nacional de Historia Natural del Paraguay,
San Lorenzo, Paraguay.
BARBOUR, T. 1913. Reptiles collected by the Yale Peruvian Expedition of
1912. Proceedings of the Academy of Natural Sciences of
Philadelphia, 65:505–507.
BARBOUR, T., AND G.K. NOBLE. 1920. Amphibians and reptiles from
southern Peru collected by the Peruvian expedition of 1914–1915
under the auspices of Yale University and the National Geographic
Society. Proceedings of the United States National Museum,
58:609–620.
BOULENGER, G.A. 1896. Catalogue of the Snakes in the British Museum
(Natural History). Vol. 3. Taylor and Francis, London, United
Kingdom.
———. 1905b. Notice of some new fossil reptiles from the Karroo beds
of South Africa. Records of the Albany Museum (Grahamstown,
South Africa), 1:331–337.
———. 1898. A list of the reptiles and batrachians collected by the late
prof. L. Balzan in Bolivia. Annali dell Museo Civico di Storia
Naturale di Genova (Italy), ser. 2, 19:128–133.
———. 1903. Description of new snakes in the collection of the British
Museum. Annals and Magazine of Natural History, ser. 7,
12:350–354.
———. 1912. Description of new reptiles from the Andes of South
America, preserved in the British Museum. Annals and Magazine of
Natural History, ser. 8, 10:420–424.
BUONGERMINI, P., E., AND T. WALLER. 1999. Geographic distribution:
Bothrops moojeni (Brazilian lancehead). Herpetological Review,
30:53.
BURGER, W.L. 1971. Genera of pitvipers. Unpublished Ph.D Dissert.,
University of Kansas, Lawrence, Kansas.
CAMPBELL, J.A., AND W.W. LAMAR. 1989. The Venomous Reptiles of
Latin America. Cornell University Press, Ithaca, New York.
———. 1992. Taxonomic status of miscellaneous Neotropical viperids,
with the description of a new genus. Occasional Papers The
Museum Texas Tech University, 153:1–31.
CARRILLO DE ESPINOZA, N. 1983. Contribución al conocimiento de las
serpientes venenosas del Perú de las familias Viperidae, Elapidae, e
Hydrophiidae. Publicaciones del Museo de Historia Natural “Javier
Prado” (Perú), 30:1–55.
CARRILLO DE ESPINOZA, N., AND J. ICOCHEA. 1995. Lista taxonómica pre-
 32 ANNALS OF CARNEGIE MUSEUM
liminar de los reptiles vivientes del Perú. Publicaciones del Museo
de Historia Natural, Universidad Nacional Mayor de San Marcos
(Perú), 49:1–27.
CEI, J.M. 1993. Reptiles del noroeste, nordeste y este de la Argentina.
Herpetofauna de las selvas subtropicales, puna y pampas. Museo
regionale di Scienze naturali Torino, Monografia (Italy), 14:1–949.
COPE, E.D. 1875 [preprint of 1876 article]. Report on the reptiles brought
by professor James Orton from the middle and upper Amazon, and
western Peru. Journal of the Academy of Natural Sciences of
Philadelphia, 8:159–87.
———. 1879. Eleventh contribution to the herpetology of Tropical
America. Proceedings of the American Philosophical Society,
18:261–277.
DAVID, P., AND I. INEICH. 1999. Les serpentes venimeux du monde: systé-
matique et repartition. Dumerilia, 3:3–499.
DE LA RIVA, I., J. CASTROVIEJO B., AND J. CABOT N. 1992. Pseustes sul-
phureus (Wagler, 1824) (Serpentes: Colubridae) nueva especie para
Bolivia y datos sobre la herpetofauna boliviana. Acta Zoologica
Lilloana, 41:215–218.
DIRKSON, L., P.L. IBISCH, J. KOHLER, AND W. BOHME. 1995. Zur
Herpetofauna der semihumiden Samaipata-Region, Bolivien. II.
Reptilien. Herpetofauna (Germany), 17:15–28.
DOWLING, H.G. 1951. A proposed standard system of counting ventrals in
snakes. British Journal of Herpetology, 1:97–99.
DUELLMAN, W.E. 1979. The herpetofauna of the Andes: patterns, distri-
bution, origin, differentiation and present communities.
Pp. 371–459, in The South American Herpetofauna: Its Origin,
Evolution, and Dispersal (W.E. Duellman, ed.). Monograph of the
Museum of Natural History, The University of Kansas, No. 7.
DUNN, E.R. 1946. A small collection from eastern Peru. Proceedings of
the Biological Society of Washington, 59:17–20.
FREITAS, M.A. DE. 1999. Serpentes da Bahia e do Brasil. Editora Dall,
Bahia, Brazil.
FUGLER, C.M. 1986. Una lista preliminar de las serpientes de Bolivia.
Ecología en Bolivia (Bolivia), 8:45–72.
FUGLER, C.M., AND I. DE LA RIVA. 1990. Herpetologica Boliviana: Lista
provisional de las serpientes conocidas en el país. Museo Nacional
de Historia Natural (Bolivia) Comunicación, 9:22–53.
FUGLER, C.M., I. DE LA RIVA, AND J. CABOT. 1995. Herpetologica
Boliviana: una lista comentada de las serpientes de Bolivia con
datos sobre su distribución. Ecología en Bolivia, 1995:40–87.
GOLAY, P., H.M. SMITH, D.G. BROADLEY, J.R. DIXON, C. MCCARTHY, J.-C.
RAGE, B. SCHATTI, AND M. TORIBA. 1993. Endoglyphs and Other
Major Venomous Snakes of the World. A Checklist. Azemiops,
Switzerland.
GRIFFIN, L.E. 1916. A catalogue of the ophidia from South America at
present (June, 1916) contained in the Carnegie Museum with
descriptions of some new species. Memoirs of the Carnegie
Museum, 7:163–228.
GUTBERLET, R.L., JR. 1998. The phylogenetic position of the Mexican
blacktailed pitviper (Squamata: Viperidae: Crotalinae).
Herpetologica, 54:184–206.
GUTBERLET, R.L., JR., 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.
GUTBERLET, R.L., JR., AND M.B. HARVEY. 1998. Comments on the pro-
posed conservation of the specific and subspecific names of
Trigonocephalus pulcher Porter, 1862 and Bothrops albocarinatus
Shreve, 1934 (Reptilia, Serpentes) by the designation of a neotype
for T. pulcher. Bulletin of Zoological Nomenclature, 55:29–32.
———. 2002. Phylogenetic relationships of New World pitvipers as
inferred from anatomical evidence. Pp. 51–68, in Biology of the
Vipers (G.W. Schuett, M. Höggren, M.E. Douglas, and H.W. Green,
eds.). Eagle Mountain Publishing, Eagle Mountain, Utah, U.S.A.
HARRIS, H.S., AND R.S. SIMMONS. 1972a. A checklist of the rattlesnakes
(Crotalus durissus group) of South America. Bulletin of the
Maryland Herpetological Society, 8:27–32.
———. 1972b. Keys to the Neotropical species and subspecies of the
Downloaded From: https://bioone.org/journals/Annals-of-Carnegie-Museum on 09 Sep 2020
Terms of Use: https://bioone.org/terms-of-use Access provided by Pontifica Universidade Catolica do Rio Grande do Sul (PUCRS)
VOL. 74
Crotalus durissus group. Bulletin of the Maryland Herpetological
Society, 8:33–40.
HARVEY, M.B. 1994. A new species of montane pitviper (Serpentes:
Viperidae: Bothrops) from Cochabamba, Bolivia. Proceedings of the
Biological Society of Washington, 107:60–66.
———. 1998. Reptiles and amphibians of Parque Nacional Noel Kempff
Mercado. Pp. 144–166, 348–355, in A Biological Assessment of
Parque Noel Kempff Mercado, Bolivia (T.J. Killeen and T.S.
Schulenberg, eds.). Conservation International, Washington, D.C.
———. 1999. Revision of Bolivian Apostolepis (Squamata: Colubridae).
Copeia, 1999:388–409.
HARVEY, M.B., J. APARICIO E., AND L. GONZALES A. 2003. Revision of the
venomous snakes of Bolivia: Part 1. The coralsnakes (Elapidae:
Micrurus). Annals of Carnegie Museum, 72:1–52.
HARVEY, M.B., AND R.L. GUTBERLET, JR. 1998. Lizards of the genus
Tropidurus (Iguania: Tropiduridae) from the Serranía de
Huanchaca, Bolivia: new species, natural history, and a key to the
genus. Herpetologica, 54:493–520.
KITCHING, J.W. 1977. Distribution of the Karroo vertebrate fauna. Memoirs
of the Bernard Price Institute for Palaeontological Research, 1:1–131.
HENLE, K., AND A. ERHL. 1991. Zur Reptilienfauna Perus nebst
Beschreibung eines neue Anolis (Iguanidae) und zweier neuer
Schlangen (Colubridae). Bonner Zoologische Beiträge (Germany),
42:143–180.
HOGE, A.R. 1953 (dated 1952). Notas erpetológicas 2a. Contribuição ao
conhecimento dos ofídios do Brasil central. Memórias do Instituto
Butantan (Brazil), 24:179–214.
———. 1954. A new Bothrops from Brazil. Bothrops brazili, sp. nov.
Memórias do Instituto Butantan (Brazil), 25:15–22.
———. 1966 (dated 1965). Preliminary account of Neotropical Crotalinae
(Serpentes Viperidae). Memórias do Instituto Butantan (Brazil),
32:109–184.
HOGE, A., AND S.A. ROMANO. 1973 (dated 1972). Sinopse das serpentes
peçonhentas do Brasil. Memórias do Instituto Butantan (Brazil),
36:109–208.
HOGE, A., S.A. ROMANO, P.A. FEDERSONI, JR., AND C.L.S. CORDEIRO. 1975
(dated 1974). Lista das espécies de serpentes coletadas na região da
usina hidroelétrica de Ilha Solteira. Brasil. Memórias do Instituto
Butantan (Brazil), 38:167–179.
HOGE, A., AND S.A. ROMANO-HOGE. 1981 (dated 1978/79). Poisonous
snakes of the world. Part 1. Checklist of the pit vipers Viperoidea,
Viperidae, Crotalinae. Memórias do Instituto Butantan (Brazil),
42/43:179–310.
HOGE, A. R., C.R. RUSSO, M.C. SANTOS, AND M.F.D. FURTADO. 1981 (dated
1978/79). Snakes collected by “Projeto Rondon XXII” to Piauí,
Brazil. Memórias do Instituto Butantan (Brazil), 42/43:87–94.
HOOGMOED, M.S., AND U. GRUBER. 1983. Spix and Wagler type specimens
of reptiles and amphibians in the Natural History Musea in Munich
(Germany) and Leiden (The Netherlands). Spixiana, supplement,
9:319–415.
ITOH, N., N. TANAKA, S. MIHASHI, AND I. YAMASHINA. 1987. Molecular
cloning and sequence analysis of cDNA for batroxobin, a thrombin-
like snake venom enzyme. Journal of Biological Chemistry,
262:3132–3135.
JORGE DA SILVA, N., JR. 1993. The snakes from Samuel Hydroelectric
power plant and vicinity, Rondônia, Brazil. Herpetological Natural
History, 1:37–86.
JORGE DA SILVA, N., JR., AND J.W. SITES. 1995. Patterns of diversity of
Neotropical squamate reptile species with emphasis on the Brazilian
Amazon and the conservation potential of indigenous reserves.
Conservation Biology, 9:873–901.
KEMPFF-MERCADO, N. 1975. Ofidios de Bolivia. Academia Nacional de
Ciencias, La Paz, Bolivia.
KLAUBER, L.M. 1941. A new species of rattlesnake from Venezuela.
Transactions of the San Diego Society of Natural History,
30:333–336.
———. 1972. Rattlesnakes: Their Habits, Life Histories, and Influence on
Mankind. 2nd edition. 2 volumes. University of California Press,
Berkeley, California.
 2005 HARVEY ET AL.—BOLIVIAN PITVIPERS
KLEMMER, K. 1963. Liste der rezenten Giftschlangen: Elapidae,
Hydrophiidae, Viperidae und Crotalidae. Pp. 255–264 in Die
Giftschlangen der Erde. N.G. Elwert Universitäts-und-Verlags-
Buchhandlung, Marburg, Germany.
KRAUS, F., D.G. MINK, AND W.M. BROWN. 1996. Crotaline intergeneric
relationships based on mitochondrial DNA sequence data. Copeia,
1996:763–773.
KUCH, U. 1997. Comments on the proposed conservation of the specific
and subspecific names of Trigonocephalus pulcher Peters, 1862 and
Bothrops albocarinatus Shreve, 1934 (Reptilia, Serpentes) by the des-
ignation of a neotype for T. pulcher. Bulletin of Zoological
Nomenclature, 54:245–249.
KUCH, U., AND A. FREIRE. 1995. Notes on morphology, reproduction and
medical importance of the poorly known Small-eyed Lancehead,
Bothrops microphthalmus Cope, 1876, in Ecuador. Herpetozoa
(Germany), 8:81–83.
LAVILLA, E.O., AND G. SCROCCHI. 1999. Anfibios y reptiles de Tariquía.
Pp. 83–92, in Relevamiento de la biodiversidad de la Reserva
Nacional de Flora y Fauna Tariquía (Tarija, Bolivia) (J.A. González,
G.J. Scrocchi, and E.O. Lavilla, eds.). Fundación Miguel Lillo, San
Miguel de Tucumán, Argentina.
LAZELL, J.D., JR. 1964. The Lesser Antillean representatives of Bothrops
and Constrictor. Bulletin of the Museum of Comparative Zoology at
Harvard College, 132:245–273.
LEVITON, A.E., R.H. GIBBS, JR., E. HEAL, AND C.E. DAWSON. 1985.
Standards in herpetology and ichthyology. Part I. Standard symbolic
codes for institutional resource collections in herpetology and ichthy-
ology. Copeia, 1985:802–832.
LINNAEUS, C.L. 1758. Systema Naturae per Regna Tria Naturae, Secundum
Classes, Ordines, Genera, Species, cum Characteribus, Differentiis,
Synonymis, Locis, Editio Decima. Volume 1. Laurenti Salvi, Holmiae
(=Stockholm).
———. 1766. Systema Naturae per Regna Tria Naturae, Secundum
Classes, Ordines, Genera, Species, cum Characteribus, Differentiis,
Synonymis, Locis, Editio Duodecima. Volume 1. Laurenti Salvi,
Holmiae (=Stockholm).
LÖNNBERG, E. 1902. On a collection of snakes from north-western
Argentina and Bolivia containing new species. The Annals and
Magazine of Natural History, 7:457–462.
MARTINS, M., O.A.V. MARQUES, AND I. SAZIMA. 2002. Ecological and phy-
logenetic correlates of feeding habits in Neotropical pitvipers of the
genus Bothrops. Pp. 307–328, in Biology of the Vipers (G.W. Schuett,
M. Höggren, M.E. Douglas, and H.W. Green, eds.). Eagle Mountain
Publishing, Eagle Mountain, Utah, U.S.A.
MASLIN, T.P. 1942. Evidence for the separation of the crotalid genera
Trimeresurus and Bothrops, with a key to the genus Trimeresurus.
Copeia, 1942:18–24.
MCCRANIE, J.M. 1993. Crotalus durissus Linnaeus. Neotropical
Rattlesnake. Catalogue of American Amphibians and Reptiles,
577:1–11.
MCDIARMID, R.M., J.A. CAMPBELL, AND T.A. TOURÉ. 1999. Snake Species
of the World. A Taxonomic and Geographical Reference. Volume 1.
The Herpetologists’ League, Washington, D.C.
MCDIARMID, R.M., AND M.S. FOSTER. 1987. Additions to the reptile fauna
of Paraguay with notes on a small herpetological collection from
Amambay. Studies on Neotropical Fauna and Environment, 22:1–9.
MENESES, O. 1974. Ofidios y ofidismo en el Peru. I. Las serpientes
venenosas del Perú. Revista del Instituto Zoonosis e Investigación
Pecuaria 2:69–77.
MERTENS, V.R. 1929. Über einige Amphibien und Reptilien aus Süd-
Bolivien. Zoologischer Anzeiger, 86:57–62.
MIDDENDORF, G., AND R. REYNOLDS. 2000. Herpetofauna of the Beni
Biological Station Biosphere Reserve, Amazonian Bolivia: additional
information, and current knowledge in context. Pp. 151–169 in
Biodiversity, Conservation and Management in the Region of the
Beni Biological Station Biosphere Reserve, Bolivia (O. Herrera-
MacBryde, F. Dallmeier, B. MacBryde, J.A. Comiskey, and C.
Miranda, eds.). Smithsonian Institution, Washington, D.C.
MIRANDA, C., M. O. RIBERA A., J. SARMIENTO T., E. SALINAS, AND C. NAVIA
Downloaded From: https://bioone.org/journals/Annals-of-Carnegie-Museum on 09 Sep 2020
Terms of Use: https://bioone.org/terms-of-use Access provided by Pontifica Universidade Catolica do Rio Grande do Sul (PUCRS)
33
R., EDS. 1991. Plan de Manejo de la Reserva de la Biosfera Estación
Biológica del Beni 1991. Academia Nacional de Ciencias de Bolivia,
Estación Biológica del Beni and LIDEMA, La Paz (Bolivia).
MONTERO, R., G. SCROCCHI, M.E. MONTAÑO, AND I.M. FERNÁNDEZ S. 1995.
Nuevas citas de saurios, anfisbénidos y ofidios para Bolivia.
Cuadernos de Herpetología (Argentina), 9:7–13.
MORALES, V.R., AND R.W. MCDIARMID. 1996. Annotated checklist of the
amphibians and reptiles of Pakitza, Manu National Park Reserve
Zone, with comments on the herpetofauna of Madre de Dios, Peru.
Pp. 503–522, in Manu. The Biodiversity of Southeastern Peru (D.E.
Wilson and A. Sandoval, eds.). National Museum of Natural History,
Washington, D.C.
MURPHY, R.W., J. FU, A. LATHROP, J.V. FELTHAM, AND V. KOVAC. 2002.
Phylogeny of the rattlesnakes (Crotalus and Sistrurus) inferred from
sequences of five mitochondrial DNA genes. Pp. 69–92, in Biology of
the Vipers (G.W. Schuett, M. Höggren, M.E. Douglas, and H.W.
Green, eds.). Eagle Mountain Publishing, Eagle Mountain, Utah,
U.S.A.
MYERS, C.W., AND J.A. CAMPBELL. 1981. A new genus and species of col-
ubrid snake from the Sierra Madre del Sur of Guerrero, Mexico.
American Museum Novitates, 2708:1–20.
NASCIMENTO, F.P. DO, T.C.S. DE ÁVILA-PIRES, O.R. DA CUNHA. 1988.
Répteis Squamata de Rondônia e Mato Grosso colectados através do
Programa Polonoroeste. Boletim Museu Paraense Emílio Goeldi,
Série Zoologia (Brazil), 4:21–66.
OBST, F.-J. 1977. Die herpetologische Sammlung des Staatlichen Museums
für Tierkunde Dresden und ihre Typusexemplare. Zoologische
Abhandlungen. Staatlichen Museum für Tierkunde in Dresden
(Germany), 34:171–186.
ORCÉS V., G. 1943. Ofidios Venenosos del Ecuador (Conclusión). Flora
(Ecuador), 3:165–170.
PARKER, H.W. 1934. Reptiles and amphibians from Southern Ecuador.
Annals and Magazine of Natural History, 10:264–273.
PARKINSON, C.L., J.A. CAMPBELL, AND P.T. CHIPPINDALE. 2002. Multigene
phylogenetic analysis of pitvipers, with comments on their biogeogra-
phy. Pp. 93–110, in Biology of the Vipers (G.W. Schuett, M. Höggren,
M.E. Douglas, and H.W. Green, eds.). Eagle Mountain Publishing,
Eagle Mountain, Utah, U.S.A.
PEÑARANDA B., E., A. CALVO C., AND J. PEÑARANDA. 1994. Biología repro-
ductiva de Bothrops neuwiedi (Serpentes, Viperidae) en cautiverio:
resultados preliminares. Ecología en Bolivia (Bolivia), 23:57–70.
PERACCA, M.G. 1897. Viaggio del Dott. Alfredo Borrelli nel Chaco boli-
viano e nella Repubblica Argentina. Bolletino del Musei di Zoologia
ed Anatomia Comparata, 12:1–19.
PETERS, J.A. 1960 The snakes of Ecuador. A checklist and key. Bulletin of
the Museum of Comparative Zoology at Harvard College,
122:491–541.
PETERS, J.A., AND B. OREJAS-MIRANDA. 1970. Catalogue of the Neotropical
Squamata. Part I. Snakes. United States National Museum Bulletin,
297:i–viii, 1–347.
PIFANO C., F., AND M. RÖMER. 1949. Ofidios ponzoñosos de Venezuela. I.
Nueva comprobación en Venezuela del Bothrops medusa (Sternfeld,
1920) Amaral, 1929 y redescripción de la especie. Archivos
Venezolanos de Patología Tropical y Parasitología Médica
(Venezuela), 1:277–300.
PRADO, A, AND A. HOGE. 1948 [dated 1947]. Notas ofiológicas 21.
Observações sobre serpentes do Perú. Memórias do Instituto Butantan
(Brazil), 20:283–-296.
RENDAHL, H., AND G. VESTERGREN. 1940. Notes on Colombian snakes.
Arkiv För Zoologi (Sweden), 33:1–16.
SALOMÃO, M.G., W. WÜSTER, R.S. THORPE, AND BBBSP. 1999. MtDNA
phylogeny of Neotropical pitvipers of the genus Bothrops (Squamata:
Serpentes: Viperidae). Kaupia, 8:127–134.
SCHMIDT, K.P., AND W.F. WALKER, JR. 1943. Peruvian snakes from the
University of Arequipa. Fieldiana, 24:279–296.
SCHÄTTI, B., AND E. KRAMER. 1993. Ecuadorianische Grubenottern der
Gattungen Bothriechis, Bothrops und Porthidium (Serpentes:
Viperidae). Revue suisse de Zoologie (Switzerland), 100:235–278.
SERIÉ, P. 1915. Suplemento a la fauna erpetológica argentina. Anales del
 34 ANNALS OF CARNEGIE MUSEUM
Museo Nacional de Historia Natural de Buenos Aires, 27:83–100.
———. 1921. Catálogo de los ofidios argentinos. Anales Sociedad
Ciencias de Argentina, 92:145–172.
SILVA, V.X. 2000. Revisão sistemática do complexo Bothrops neuwiedi
(Serpentes, Viperidae, Crotalinae). Unpublished Ph.D. Dissert.
Universidade de São Paulo, São Paulo, Brazil.
SMITH, H.M., AND E.H. TAYLOR. 1950. Type localities of Mexican reptiles
and amphibians. University of Kansas Science Bulletin, 33:313–380.
STRUSSMANN, C. AND I. SAZIMA. 1993. The snake assemblage of the
Pantanal at Poconé, western Brazil: faunal composition and ecologi-
cal summary. Studies on Neotropical Fauna and Environment,
28:157–168.
VANZOLINI, P.E. 1981. The scientific and political contexts of the Bavarian
expedition to Brasil. (Introduction) Pp. ix–xxix in J.B. von Spix and
J.G. Wagler: Herpetology of Brazil. SSAR reprint (K. Adler, ed.).
Society for the Study of Amphibians and Reptiles, Lawrence, Kansas.
———. 1986. Levantamento herpetológico da área do estado de Rondônia
sob a influência da rodovia BR 364. Programa Polonoroeste,
Relatório de Pesquisa (Brazil), 1:1–50.
VANZOLINI, P.E., AND M.E.V. CALLEFFO. 2002. A taxonomic bibliography of
the South American snakes of the Crotalus durissus complex
(Serpentes, Viperidae). Anais da Academia Brasileira de Ciências
(Brazil), 74:37–83.
VISINONI, A. 1995. Contribuciones para el conocimiento de la herpetofau-
na boliviana: Lachesis muta muta (Linnaeus, 1766) (Serpentes:
Viperidae) en Bolivia. Ecología en Bolivia, 24:103–112.
WAGLER, J. 1824. Serpentum brasiliensium species novae ou Histoire
naturelle des espèces nouvelles de serpents, recueillies et observées
pendant le voyage dans l'intérieur du Brésil dans les années 1817,
1818, 1819, 1820, exécuté par ordre de Sa Majesté le Roi de Bavière,
publiée par Jean de Spix, ..., écrite d'après les notes du voyageur par
Jean Wagler... F.S. Hübschmann, Munich (Germany).
WERMAN, S.D. 1992. Phylogenetic relationships of Central and South
American pitvipers of the genus Bothrops (sensu lato): cladistic analy-
sis of biochemical and anatomical characters. Pp. 21–40, in Biology
of the Pitvipers (J.A. Campbell and E.D. Brodie, Jr., eds.). Selva,
Tyler, Texas, USA.
———. 1999. Molecular phylogenetics and morphological evolution in
Neotropical pitvipers: an evaluation of mitochondrial DNA sequence
information and the comparative morphology of the cranium and
palatomaxillary arch. Kaupia, 8:113–126.
WERNER, F. 1901. Reptilien und Batrachier aus Peru und Bolivien.
Abhandlungen und Berichte des Königlichen Zoologischen
Downloaded From: https://bioone.org/journals/Annals-of-Carnegie-Museum on 09 Sep 2020
Terms of Use: https://bioone.org/terms-of-use Access provided by Pontifica Universidade Catolica do Rio Grande do Sul (PUCRS)
VOL. 74
und Anthropologisch-Ethnographischen Museums zu Dresden
(Germany), 2:1–13.
———. 1922. Synopsis der Schlangen-familien der Amblycephalidae und
Viperidae nebst Übersicht über die kleineren Familien und die
Colubriden der Acrochordinengruppe. Auf Grund des
Boulengerschen Schlangenkatalogs (1893–1896). Archiv für
Naturgeschichte (Germany), 88:185–244.
———. 1927. Neue oder wenig bekannte Schlangen aus dem Wiener
naturhistorischen Staatsmuseum (III. Teil). Sitzungsberichte der
Akademie der Wissenschaften in Wien. Mathematisch-
Naturwissenschaftliche Klasse (Austria), 135:243–257.
WÜSTER, W. 1998. Comments on the proposed conservation of the specif-
ic and subspecific names of Trigonocephalus pulcher Peters, 1862
and Bothrops albocarinatus Shreve, 1934 (Reptilia, Serpentes) by the
designation of a neotype for T. pulcher. Bulletin of Zoological
Nomenclature, 55:34–36.
WÜSTER, W., S. PIERINI, AND G. PUORTO. 1994. Bothrops moojeni.
Herpetological Review (Geographic Distribution) 25:166 (originally
2:76 with printing errors.).
WÜSTER, W., R.S. THORPE, G. PUORTO, AND BBBSP. 1996. Systematics of
the Bothrops atrox complex (Reptilia: Serpentes: Viperidae) in Brazil:
A multivariate analysis. Herpetologica, 52:263–271.
WÜSTER, W, M.G. SALOMÃ, R.S. THORPE, G. PUORTO, M.F.D. FURTADO,
S.A. HOGE, R.D.G. THEAKSTON, AND D.A.WARRELL. 1997.
Systematics of the Bothrops atrox complex: new insights from multi-
variate analysis and mitochondrial DNA sequence information. Pp.
93–113, in Venomous Snakes: Ecology, Evolution, and Snakebite (R.
S. Thorpe, W. Wüster, and A. Malhotra, eds.). Oxford University
Press, Oxford, United Kingdom.
WÜSTER, W., M.G. SALOMÃO, G.J. DUCKETT, R.S. THORPE, AND BBBSP.
1999. Mitochondrial DNA phylogeny of the Bothrops atrox species
complex (Squamata: Serpentes: Viperidae). Kaupia, 8:135–144.
WÜSTER, W., M.G. SALOMÃO, J.A. QUIJADA-MASCAREÑAS, R.S. THORPE,
AND BBBSP. 2002. Origins and evolution of the South American
pitviper fauna: evidence from mitochondrial DNA sequence analysis.
Pp. 111–128, in Biology of the Vipers (G.W. Schuett, M. Höggren,
M.E. Douglas, and H.W. Green, eds.). Eagle Mountain Publishing,
Eagle Mountain, Utah, U.S.A.
ZAMUDIO, K.R., AND H.W. GREENE. 1997. Phylogeography of the bushmas-
ter (Lachesis muta: Viperidae): implications for Neotropical biogeog-
raphy, systematics, and conservation. Biological Journal of the
Linnean Society, 62:421–442.
 2005

TABLE 1. Selected diagnostic characters for some South American pitvipers. Data in the table were collected from specimens examined during the course of this study.

Species Rectangular Canthorostrals Two or Fewer Scale(s) More than Subocular-

(sample size) Loreal Intercanthals Between Three Supralabial
Internasals Prefoveals Contact

Bothriopsis bilineata (20) Absent Absent 20% (2–5) 0% 51% (2–20) 15%
B. pulchra (7) Absent Absent 0% (3–5) 14% 17% (2–4) 67%
B. punctata (8) Absent Absent 25% (2–4) 62% 0% (1–3) 6%
B. taeniata (11) Absent Absent 45% (2–4) 18% 0% (2–3) 36%
Bothrops alternatus Absent Absent 0% (4–7) 0% 11% (3–10) 0%
B. andianus (23) Present Absent 0% (3–5) 91% 17% (1–5) 0%
B. atrox (Bolivia) (17) Absent Absent 0% (3–7) 22% 6% (1–5) 0%
B. brazili (6) Absent Absent 0% (3–4) 67% 33% (1–6) 0%
B. caribbaeus (5) Absent Absent 40% (2–3) 0% 0% (0–2) 0%

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B. itapentiningae (6) Absent Absent 0% (5–9) 0% 58% (2–7) 67%
B. jararaca (20) Absent Absent 5% (2–6) 0% 10% (1–6) 0%
B. jararacussu (15) Absent Absent 100% (0–2) 0% 0% (1–3) 3%
B. jonathani (6) Absent Absent 0% (4–6) 0% 100% (4–8) 0%
B. lanceolatus (5) Absent Absent 0% (5–7) 40% 0% (2–3)
B. lojanus Present Absent 0% (3–5) 0% 0% (1–2)
HARVEY ET AL.—BOLIVIAN PITVIPERS

B. moojeni (12) Absent Absent 0% (4–7) 67% 8% (1–4) 0%

B. neuwiedi (39) Absent Absent 3% (2–6) 0% 17% (3–8) 0%

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B. sanctaecrucis (16) Absent Absent 3% (0–3) 9% 0% (1-3) 0%
B. venezuelensis (8) Absent Absent 0% (3–5) 0% 6% 0%
Bothrocophias hyoprora (6) Absent 100% 0% (3–6) 100% 100% (5–9) 0%
B. microphthalmus (12) Present 75% (1–2) 0% (4–5) 100% 100% (4–8) 0%
Crotalus durissus (16) Absent Absent 100% (0) 0% 87% (2–5) 0%
Lachesis muta (8) Absent 8% (1) 0% (8–12) 100% 67% (2–6) 0%
35
 36 ANNALS OF CARNEGIE MUSEUM VOL. 74

TABLE 2. Characteristics of four easily confused Bothrops occurring or potentially occurring in Bolivia.

Characteristic B. atrox B. moojeni B. sanctaecrucis B. brazili

Postocular stripe
Ventral pigmentation
2–2.5 temporals high 1 temporal high 1 temporal high 1 temporal high
or absent or absent
Heavy, usually alternating Diffuse Diffuse Moderate to heavy
cream and gray square
blotches

Scales between canthals 2–6t 4–7 0–3 2–4

Stripe on neck Absent To level of ventral 12–24 Absent Absent

Distal tail of adults Not sharply Not sharply Black; sharply Not sharply
contrasting contrasting contrasting contrasting
with dorsum with dorsum with dorsum with dorsum

Iris Bronze Bronze Pale green Copper to pinkish copper

Supralabials 7–8 (Bolivia) 7 8–9 7–9

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 2005 HARVEY ET AL.—BOLIVIAN PITVIPERS 37

APPENDIX
Additional Comparative Material

Bothriopsis medusa—VENEZUELA. Aragua: USNM 129585.
Bothriposis pulchra—COLOMBIA. UTA 39066. ECUADOR.
FHGO 263, 884, 1216, 2019, KU 121347–48. PERU. ZFMK 41480.
Bothriopsis punctata—COLOMBIA. Cauca: USNM 124258,
151706. Chocó: USNM 72355. ECUADOR. Chimborazo: USNM
310822. Esmeraldas: USNM 20629–30. Pinchincha USNM
165286, 232521. Boothrocophias campbelli—ECUADOR:
Pinchincha: USNM 165322, 165340. Bothrocophias myersi—
COLOMBIA. Cauca: USNM 154051, 151708. Bothrops
alternatus—ARGENTINA. Formosa: USNM 63499. BRAZIL.
Paraná: USNM 100706, 100722. Rio Grande do Sul: USNM
287508. São Paulo: USNM 38182–83, 69288–89, 69292, 69294–95,
55862. PARAGUAY. Presidente Hayes: USNM 342418–20,
342422, 342424. URUGUAY. Cerro Largo: USNM 65611.
Bothrops ammodytoides—ARGENTINA. La Rioja: USNM 73421.
Bothrops brazilli—ECUADOR. Pastaza: USNM 165339, 321137.
PERU. Amazonas: USNM 316670–71. Madre de Dios: USNM
345184. Bothrops caribbaeus—SANTA LUCIA: USNM 11312,
13841, 15083, 15082, 222953. Bothrops cotiara—BRAZIL. Santa
Catarina: USNM 76317, 100695, 100750. Bothrops itapetingae—
BRAZIL. São Paulo: USNM 38187, 39059, 76320, 165514–16.
Bothrops jararaca—BRAZIL. Rio de Janeiro: USNM 98628–35,
207676. Santa Catarina: USNM 217841. São Paulo: USNM
38177–79, 100698, 100670–71, 100679, 208138. Bothrops
jararacussu—BRAZIL. Santa Catarina: USNM 217842. Paranã:
USNM 110720, 165510. Rio de Janeiro: USNM 98647, 100713. São
Paulo: USNM 38180, 39054, 69300–02, 71338. PARAGUAY.
Itapua: USNM 253589–92. Bothrops lanceolatus—MARTINIQUE:
St. Pierre: USNM 11317–18; Unknown: USNM 11308. TOBAGO
[probably in error: These specimens almost certainly came from
Martinique (Lazell 1964)]: USNM 10116, 10122. Bothrops lojanus—
EQUADOR. Loja: USNM 98927, 98935, 232519. Bothrops pictus—
PERU. Lima: USNM 49992. Bothrops venezuelensis—
VENEZUELA. Aragua: USNM 129580–82, 129584, 134405,
142397, 259174–75.

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