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Revision of Bolivian Apostolepis (Squamata: Colubridae)

Author(s): Michael B. Harvey


Source: Copeia, Vol. 1999, No. 2 (May 7, 1999), pp. 388-409
Published by: American Society of Ichthyologists and Herpetologists (ASIH)
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Copeia,1999(2), pp. 388-409

Revision of Bolivian Apostolepis(Squamata: Colubridae)

MICHAELB. HARVEY

Six species of Apostolepisare known to inhabit Bolivia. The holotypes of A. dor-


bignyiand A. nigroterminata are redescribed.Variationin these species and A. tenuis
and A. vittatais documented. Two new species are described:A. multicincta,from
1290 m in the BolivianAndes, is the first species of the genus ever found above
500 m; A. phiUipsiis a new species from northeasternSantaCruzdepartment.A key
to the Bolivianspecies and six species from adjacentcountriesis provided.

SNAKES of the genus Apostolepisare a speci- terminata Boulenger, A. tenuis Ruthven, and
ose group of small fossorial colubrids. They Apostolepisvittata (Cope). Except for NK (Museo
are closely related to Elapomorphusand Phalotris, de Historia Natural "Noel Kempff Mercado,"
and the three genera are often referred to as Santa Cruz, Bolivia), institutional abbreviations
"elapomorphines" (Savitzky, 1979; Lema, 1984; are those suggested by Leviton et al. (1985).
Ferrarezzi, 1993a, 1993b). The phylogenetic af- Measurements reported below include those
finities of the elapomorphines have been the taken with a meter stick to the nearest 1 mm,
subject of considerable interest yet remain un- snout-vent length (SVL) and tail length, or with
resolved (Cadle and Sarich, 1981; Cadle, 1982, an ocular micrometer to the nearest 0.01 mm:
1988). Most recently, Underwood and Kochva head length (caudal tip of last supralabial to tip
(1993) tested for a close relationship between of snout as seen in lateral view); head width (at
Apostolepis, Elapomorphus, and Atractaspis. Al- rictus; note this may not be the widest part of
though they concluded that the last is probably the head. In A. nigroterminata,for example, the
not closely related to elapomorphines, they not- head is widest at the level of the suture separat-
ed that the two groups nonetheless share some
ing the fifth and sixth supralabials); eye-nostril
special characteristics unknown in other snakes. distance (anterior edge of eye to center of nos-
Apostolepiscontains about 20 species distrib- tril); eye diameter; frontal length (frontal's
uted across cis-Andean South America. As sev-
greatest length, always along midline); suture
eral authors have pointed out (e.g., Cunha and between parietals; suture between prefrontals;
Nascimento, 1978; Vanzolini, 1986, unpubl.), suture between first pair of chinshields. Sex was
this group of snakes is rare in collections, and determined by subcaudal incision. Characteriza-
many species have been confused in the litera- tion of the hemipenis in the definition of the
ture. Until recently, three species had been re-
genus is based primarily on everted hemipenes
ported from Bolivia: Apostolepisnigroterminata(as of Apostolepisassimilis (MVZ 176333) and Apos-
Apostolepisborelli;Parker, 1928); Apostolepisdorbig- tolepisdimidiata (USNM 76369), a partially evert-
nyi (Peracca, 1897); and Apostolepistenuis (Ruth- ed organ of Apostolepismulticincta (NK 729), and
ven, 1927; Peters and Orejas-Miranda, 1972).
organs of A. nigroterminatadissected by earlier
Harvey (1998), examining material recently col- researchers in some specimens of A. nigrotermi-
lected by B. Phillips from the eastern border of
nata and examined in situ. A description of the
Bolivia, discovered a fourth species. As part of
an ongoing review of Bolivian snakes (Harvey, hemipenis of Apostolepisquinquelineata (Cunha
and Nascimento, 1978) was also considered.
1994), I here review the Bolivian Apostolepisand
comment on most species from adjacent coun- Terminology used in the description of hemi-
tries. Two new species are described, and the penes is that of Dowling and Savage (1960).
Scale counts from opposite sides of the same
types of A. dorbignyiand A. nigroterminataare re-
described. The identity of Apostolepislineata is specimen are separated by a slash (e.g., 6/7 in-
discussed. fralabials) as opposed to a dash which is used
to report ranges for more than one specimen.

MATERIALSAND METHODS
TAXONOMICCHARACTERS
This study is based on examination of six
specimens of two species of Elapomorphusand 64 In the diagnoses below, characters are pre-
specimens of 13 species of Apostolepis(Appen- sented in a numbered and standardized format
dix), including the holotypes or syntypes of A. corresponding to the following list of character-
dorbignyi(Schlegel), A. lineata (Cope), A. nigro- istics.
? 1999 by the American Society of Ichthyologists and Herpetologists

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HARVEY-BOLIVIAN APOSTOLEPIS 389

4. Temporalsand parietal-supralabial contact (Fig.


1).-Species of Apostolepistypically have no tem-
porals (0 + 0) or second temporals (0 + 1).
Number of supralabials contacting the parietals
showed considerable intraspecific variation and
was not used to diagnose species but is reported
in the descriptive sections of species accounts.

5. Numberof infralabialscontactingfirstpair of chin-


shields (Fig. 3).-Most species of Apostolepishave
four infralabials contacting the first pair of chin
shields, however a few species invariably have
A I 3mm only three.

6. Ventral and subcaudal ranges.-As in many


snakes, ventral and subcaudal counts are sexu-
ally dimorphic in some if not all species of Apos-
tolepis. When counting ventrals, the technique
of Dowling (1951) was used.

7. Light cephalic blotches(Fig. 2).-Many species


of Apostolepishave light, irregular blotches on
their cephalic plates. Often, these blotches are
especially prominent on the prefrontals where
they may fuse to form a continuous band across
the snout. To a certain degree, prominence and
distribution of light cephalic blotches varies in-
traspecifically. However, when present in a spe-
cies, they were noticeable in all specimens ex-
amined.

B L 3mm I 8. Light supralabial blotch (Fig. 2).-Most species


of Apostolepishave a light blotch below the eye.
Fig. 1. Diagnosticcharacteristicsof cephalic squa- Presence of a blotch is not polymorphic within
mation. (A) Contact between prefrontaland second a
species, although its extent (expressed as the
supralabial(both stippled) in Apostolepis tenuis(Ho-
number of supralabials at least partially covered
lotype, UMMZ 64436). (B) Five supralabials (stip-
pled) of Apostolepis
vittata(CM 2824). by the blotch) frequently is. In most species, the
blotch is subtriangular (e.g., Fig. 2A, C) and
covers part or all of supralabials 2-5; several spe-
1. Snout shape and extent of dorsal rostral exposure cies with pointed snouts (e.g., Apostolepisambi-
A. dimidiata, and Apostolepisphillipsi) have
(Figs. 1-2).-As a quantitative means of express- niger
of the snout, extent of the rostral an elongate blotch running along the ventral
ing projection
visible from above is compared to its distance edge of all their supralabials (Fig. 2B).
to the frontal.
9. Nuchal collars (Fig. 2).-A white nuchal collar
2. Numberof supralabials (Fig. 1).-In Apostolepis, of variable length may be present and may be
number of supralabials is fixed within and followed by a black nuchal collar. In some spe-
among groups of species except for some rare cies, the latter is nearly indistinct and is formed
and obvious instances of fusion. primarily through thickening of the lateral,
paravertebral, and vertebral stripes. Space be-
3. Nasalpreocular contact(Fig. 1).-The nasal and tween these stripes is filled by black pigment to
preocular may contact each other or be sepa- varying degrees in species with very narrow
rated by contact between the prefrontal and sec- black nuchal collars. I express length of these
ond supralabial. Polymorphism in this character collars as the number of dorsals they encom-
is rare and at low frequencies; loreals appear pass. The collars are usually irregular, and
only as apparent anomalies (e.g., on one side length was typically recorded as a range (e.g.,
only of FMNH 39646). 2-2.5) of dorsals for individual specimens.

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390 COPEIA, 1999, NO. 2

V.

(
A

~~4~~-~~E~~Bf~~

B L3mm I

((

C I 3mm I

Fig. 2. Nuchal and cephalic squamationand pigmentationof (A) Apostolepis (Paratype,NK878),


multicincta
phillipsi(Holotype, UTA 43940), and (C) Apostolepis
(B) Apostolepis (UTA 44687).
nigroterminata

10. Dorsal stripes (Fig. 4).-Species of Apostolepis ed. However, blunter tails appear to developed
may or may not have black or brown dorsal with age in some species.
stripes. Stripes are described using dorsal num-
ber as a landmark and to express width of 13. Chin and gular pigmentation.-Herein, refer-
stripes. ences to the "chin" refer to the area of the
mental, infralabials, and chin shields. The chin
11. Dorsal ground coloration.-Some species of and gular region show varying amounts of black
Apostolepismay be aposematic and have red dor- pigment among specimens of Apostolepis.Some
sa, whereas others have brown and tan dorsal specimens have a well-defined gular band. This
coloration. character exhibited considerable intraspecific
variation even within populations of some spe-
12. Extent of caudal band and color of the terminal cies such as A. nigroterminata.In other species
scale (Fig. 5).-Most species of Apostolepishave a such as A. assimilis, this character was relatively
black band completely surrounding the ends of conservative throughout their range.
their tails. The terminal scale may be either
partly or completely white or black. In a few GENUS APOSTOLEPIS
species with white terminal scales, one or more
rows of caudals proximal to the terminal scale Definition.-Small to medium fossorial snakes
are white. Intraspecific polymorphism in termi- with small heads not distinct from neck; snout
nal scale coloration was not observed in Apos- round to projecting; body long; tail short and
tolepis. Shape of the terminal scale varies intra- rhomboid (tail length 7-15% of SVL in Bolivian
specifically; it may be sharply pointed to round- species); eye small and round with round pupil;

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HARVEY-BOLIVIAN APOSTOLEPIS 391

Fig. 4. Stripingpattern at midbody of (A) Aposto-


2mm lepis phillipsi, (B) Apostolepisvittata, and (C) Apostolepis
B | I tenuis.

Fig. 3. Infralabialscontacting first pair of chin-


shields in (A) Apostolepis multicincta (Holotype, NK
729) and (B) Apostolepis phillipsi (Holotype, UTA (one exception, 17 + 17 + 17 in Apostolepispo-
43940). lylepis);high number of ventrals (200-266 in Bo-
livian species); anal plate divided; subcaudals di-
vided; sexual dimorphism in numbers of ven-
mouth reduced in size relative to most terres- trals and subcaudals.
trial colubrids; nasal entire; loreal absent; preo- Maxillary teeth increasing in size posteriorly.
cular and postocular single; preocular and nasal Anterior maxillary teeth separated from 2 large,
in contact or separated by contact between su- grooved fangs by short diastema.
pralabial and prefrontal; anterior temporal ab- Hemipenis semicapitate, subcylindrical, and
sent; posterior temporal present or absent; large weakly bilobed (i.e., only a very shallow cleft is
single scale occupying space between tips of last present distally); sulcus spermaticus bifurcating
supralabial and parietal; internasals absent; proximal to capitulum; base of hemipenis en-
paired prefrontals; frontal usually shorter than tirely spinulate; spines on asulcate side nearly 10
parietals; supralabials five or six; supralabials en- times as large as those on sulcate side; longest
tering eye, 2-3, rarely 4; infralabials 6-8, first 3 spines about two-thirds (A. assimilis) width to
or 4 contacting first pair of chinshields; scales equal (A. nigroterminata)to width of subcaudals;
smooth and lacking apical pits, 15 + 15 + 15 apex covered in calyces with papillate ridges;

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392 COPEIA, 1999, NO. 2

lepis goiasensis Prado; ApostolepisintermediaKos-


lowsky; A. lineata (Cope); Apostolepislongicaudata
Gomes; ApostolepisniceforoiAmaral; A. nigroter-
minata Boulenger; A. polylepisAmaral; Apostolepis
pymi Boulenger; A. quinquelineata Boulenger;
Apostolepis rondoni Amaral; A. tenuis Ruthven;
S mm
and A. vittata (Cope).
A I __I5 The remaining eight names are synonyms of
three species: Apostolepiscearensis,A. dimidiata,
and A. nigroterminata. Perhaps because of its
wide distribution and considerable dorsal pat-
tern polymorphism (Lema, 1993), A. dimidiata
has the largest number of synonyms among spe-
cies in the genus: ElapomorphuserythronotusPe-
ters, ApostolepisnigricepsWerner, Apostolepisbar-
B 5mm | rioi Lema, Apostolepisvillaricae Lema, and Apos-
tolepis ventrimaculatus Lema. Apostolepisamarali
Werner and Apostolepissanctae-ritaeWerner are
junior synonyms of A. cearensis(Amaral, 1930b;
Ferrarezzi, 1993a). ApostolepisborelliPeracca is a
junior synonym of A. nigroterminata (Amaral,
1930a, 1930b; this study).
The status of three Amazonian species, A.
5mm pymi, A. quinquelineata,and A. rondoni, has been
C L the subject of debate and remains unresolved.
Fig. 5. Caudal pigmentation a]nd terminal scale Apostolepisquinquelineataand A. pymi were con-
morphologyof (A) Apostolepis )si(Holotype,UTA
phillip sidered to be synonyms by Amaral (1930b) who
multicincta
43940), (B) Apostolepis (Elolotype, NK 729), examined the holotypes of each, and synony-
assimilis(MVZ176333).
and (C) Apostolepis mization is also favored by other authors (e.g.,
Cunha and Nascimento, 1978; Cunha et al.,
1985). Amaral (1930c, 1935) and Peters and
smooth, narrow furrow exte nding between Orejas-Miranda (1970) considered A. pymi to be
large spines and terminatin g in a pocket synonymous with ElapomorphuscoronatusSauva-
formed by deep calyces at poi nt where spines ge, and Vanzolini (1986) remarked that he sus-
grade sharply to apical calyces (this pocket is pected A. pymi to be a valid species extensively
clearly visible in the everted orrgans of both A. distributed in Amazonia. However, Ferrarezzi
assimilis and A. dimidiata). (1993a, 1993b) pointed out that E. coronatusis
Head darkly pigmented dors;ally with variable not a species of Apostolepisbut rather a junior
amounts of blotching; light bllotches or band synonym of Elapomorphuslepidus.
frequently present on snout; uvhite supralabial Ferrarezzi (1993a) cited the following differ-
blotch of variable size usually p)resent; gular re- ences between A. quinquelineata and A. pymi
gion immaculate to heavily pi gmented; black- (characteristics of latter in parentheses): tem-
and-white nuchal bands presenlt or not; dorsum porals 0 + 0 (0 + 1); large white spot on snout
striped or uniform; dorsal gr(ound coloration and white spots on head (spot on snout reduced
red or tan; venter usually imma.culate; posterior or absent, no spots on head); paravertebral
third of tail with black band; tip usually white, stripes very dark and cohesive (light and dif-
black in some species. fuse. A "diffuse" pattern is presumably due to
pigment being in the centers of scales on the
Content.-Twenty-eight species and subspecies sixth scale row); ventral surface of tail immac-
of Apostolepishave been descri bed. Although I ulate (black pigment generally more extensive
raise several questions regardinig the validity of on tail covering ventral and dorsal surfaces; the
certain names, I considered thle following spe- tip of the tail is white in both species).
cies when diagnosing new sy>ecies described The Belem population of A. pymi did not vi-
herein: A. ambiniger(Peters); Ajpostolepisarenaria olate these characters, but a specimen from
Rodrigues; A. assimilis (Reinl hart); Apostolepis Amazonas (AMNH 101955) collected in sym-
cearensisGomes; A. dimidiata (J[an); A. dorbignyi patry with a specimen fitting Ferrarezzi's
ata (Dumeril, Bi-
(Schlegel); Apostolepisflavotorqu (1993a) definition of A. quinquelineata (AMNH
bron, and Dumeril); A. gaboi Rodrigues; Aposto- 101954) had all the characteristics of A. pymibut

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HARVEY-BOLIVIAN APOSTOLEPIS 393

also had a large spot on the snout and white characterized beyond couplet 8 of the key, be-
spots on the head. A specimen from Guyana cause I was unable to determine whether the
(FMNH 121828) possessed some characteristics snout was pointed or round in life. However, the
of A. quinquelineata (large white spots on snout, terminal scale of A. lineata is entirely white. In
ventral surface of tail immaculate) and some the other five-lined species below couplet 8, the
characteristics of A. pymi (temporals 0 + 1, para- terminal scale is black with a white ventro-pos-
vertebral stripes indistinct). The status of A. terior blotch. The following additional obser-
pymi certainly requires further consideration vations on the syntype of A. lineata were made:
when more specimens from the type locality of subcaudals 24; 1/1 preoculars; preocular con-
A. quinquelineatabecome available. tacting nasal; 1/1 postoculars; temporals 0 + 0/
Finally, Cunha and Nascimento (1978) and 0 + 0; 6/6 supralabials, 5-6/5-6 contacting pa-
more recently Da Silva (1993) have argued that rietal; infralabials 1-4 contacting first chin-
A. rondoniand A. quinquelineataare synonymous. shield, 4-5 contacting second (could be as-
As in A. quinquelineata, the nuchal collars are sessed on one side only); vertebral stripe narrow
vestigial, temporals are present, and the lateral and restricted to dorsal 8; paravertebral stripe
stripes begin on dorsal 2 in A. rondoni. For the on dorsal 6; lateral stripe present at least on
purposes of this study, A. pymi, A. quinquelineata, dorsals 2-4 (not known if stripe extends to dor-
and A. rondoniare collectively referred to as the sal 1); 5/4 subcaudals black.
A. quinquelineatacomplex.
la. Three infralabials contacting first pair of
chin-shields(Fig. 3) --.-- ....................--- 2
KEY TO THE SPECIES OF APOSTOLEPISFROM lb. Four infralabialscontactingfirstpairof chin-
BOLIVIA AND ADJACENT REGIONS shields . ...-...- .......- ... ................................ 6
2a. Dorsum uniformly red except for nuchal
Much of Bolivia remains unexplored, and sev- and caudal bands .......................................... 3
eral additional species are likely to occur within 2b. Dorsum striped .- ....-............................ 4
its borders. Apostolepisambinige;,A. assimilis, A. 3a. Malesshort and stockyand with few ventrals
dimidiata, A. fiavotorquata, A. intermedia,A. quin- (219-225); chin and gular region lightlypig-
quelineata,and A. rondonihave all been reported mented; supralabialblotch moderate, cov-
from areas adjacent to Bolivia in Brazil (Kos- ering parts of three supralabials(Fig. 2A) --
Apostolepis multicincta
lowsky, 1898; Amaral, 1925; Da Silva, 1993), Par- 3b. Males long and slender and with manyven-
aguay (Lema, 1993), and Argentina (Serie, trals (266); chin and gular region heavily
1915; Gustavo Scrocchi, pers. comm.). Below, pigmented; supralabialblotch narrow,near-
each of these species is included in the key, be-
ly restricted to single supralabial
cause they are likely to occur in eastern Bolivia. Apostolepis dorbignyi
In formulating the key, I relied on my own 4a. Six supralabials(Fig. 1) ................................... 5
observations and on published studies of intra- 4b. Five supralabials.......................Apostolepis vittata
specific variation (e.g., Cunha and Nascimento, 5a. Snout round, three broad dorsal stripes,
1978; Lema, 1993; Da Silva, 1993). The key was high number of ventrals (245-265) and sub-
tested on all specimens examined. I have ex- caudals (37-46) --------------..- Apostolepis tenuis
amined every species included in the key except 5b. Snout projecting, five dorsal stripes, low
number of ventrals (217) and subcaudals
A. intermedia. The holotype and only known
(37) ..- .
...-.................. Apostolepis intermedia
specimen of A. intermediais lost (Lema, 1993; J. 6a. Snout round to moderatelypointed, nuchal
Williams, pers. comm.); however, Koslowsky's bands present or not, dorsum stripedor not
(1898) description and illustrations are suffi- 7
ciently thorough to allow it to be unambiguous- 6b. Snout extremelypointed (i.e., as in Fig. 1B),
ly distinguished from all its congeners. dorsum uniform, white nuchal band absent,
Apostolepislineata (Cope) was excluded from neck scales black __-..............Apostolepis ambiniger
the key, because it could not be characterized 7a. Dorsum striped or not, tip of tail white ....... 8
adequately. This species was originally repre- 7b. Dorsumuniform except for long (3 or more
sented by two syntypes; however, one of these dorsals long) bands on neck and tail;white
nuchal band present, tip of tail black (Fig.
(ANSP 11212) was discarded by E. R. Dunn who
added a label "11212 rotten and discarded" to 5C) .......-......_....... Apostolepis assimilis(Fig. 6)
8a. Snout pointed, extent of rostralvisible dor-
the jar containing the remaining syntype. The
sally greater than one-half its distance to
remaining specimen is in very bad condition; frontal (Fig. 2B) .
..................... 9
many of its scales are barely visible; all pigment 8b. Snout rounded, extent of rostralvisibledor-
on the head and most of the body is lost. Only sallyless than one-half its distance to frontal
on the tail is the original color pattern visible, (Fig. 2C) ............................................... 10
albeit quite faded. The species could not be 9a. White nuchal band present, dorsal ground

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394 COPEIA, 1999, NO. 2

4!- \

Fig. 6. Apostolepisassimilis (MVZ 176333, SVL 390


mm).
Fig. 7. Apostolepisquinquelineata (KU 127256, SVL
331 mm, top) and Apostolepisdimidiata (MCZ 27661,
coloration brown, five stripes present (Fig. SVL 402 mm, bottom).
4); ventrals immaculate ---.-- Apostolepisphillipsi
9b. White nuchal band absent, dorsal ground
coloration red; dorsal stripes black and of
variable width (Lema, 1993), usually consist- mens examined by Cunha and Nascimento,
ing of wide pair of lateral bands and a nar- 1978; Da Silva, 1993; this study); white nu-
row paravertebral band; usually, ventrals chal collar usually vestigial; dorsum striped
heavily pigmented -....- Apostolepisquinquelineatacomplex (Fig. 7)
Apostolepisdimidiata (Fig. 7)
lb. Many ventrals (253-257 in this study); white
1Oa. Relatively heavy bodied species; red to nuchal collars distinct (3-4 dorsals long in
brown dorsal ground coloration, with or specimens examined); dorsum striped or
without 3-5 longitudinal stripes, the first be- not ..---....-....
..
...-. --- Apostolepisflavotorquata
ginning on dorsal 2; temporals usually pre-
sent; nuchal collars vestigial (Fig. 7) or not;
SPECIESACCOUNTS
ventrals 203-257 .......................... ..... 11
lOb. Relatively gracile species; dorsal ground col-
oration brown; dorsum with five longitudi- Apostolepis dorbignyi (Schlegel, 1837)
nal stripes, the first beginning on dorsal 1;
white nuchal collar distinct though often Calamaria d'Orbignyi Schlegel, 1837:30. Holo-
narrow (1-3 dorsals long); temporals rarely type: MNHN 3664; type locality "Chile"
present, few ventrals (200-231) (probably in error).
Apostolepisnigroterminata Calamaria d'Orbignyii: Guichenot in Gay, 1848:
lla. Few ventrals (203-236 based on 192 speci- 73.

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HARVEY-BOLIVIAN APOSTOLEPIS 395

ElapomorphusOrbignyi:Dumeril, Bibron, and Du- rected posterolaterally; 6/6 supralabials: first


meril, 1854:834. contacting rostral and nasal; second contacting
ElapomorphusD'Orbignyi:Jan, 1862:43. Strauch, nasal, prefrontal, and preocular; second and
1884 (1888):194. third/second through fourth entering orbit;
Apostolepisorbignyi:Cope, 1861:524. third through fifth/fourth and fifth contacting
Apostolepis dorbignyi:Boulenger, 1896:236. Per- single subrectangular postocular; fifth and sixth
acca, 1897:15; Peters and Orejas-Miranda, contacting parietal; temporals absent; four oc-
1970:22; Lema, 1978:21; Ferrarezzi, 1993a: cipitals; median occipital positioned between
234. caudal tips of parietals and same size as adjacent
dorsals; left lateral occipital enlarged and filling
Definition.-(1) Snout bluntly rounded, length space just caudal to contact of sixth supralabial
of rostral visible dorsally less than one-half its with parietal; same space filled by two occipitals
distance to frontal; (2) six supralabials; (3) nasal on right side; mental slightly longer than wide,
and preocular separated by prefrontal; (4) tem- contacting chinshields; strong mental groove;
porals absent; (5) three infralabials contacting length of suture between first pair of chin-
first pair of chinshields; (6) ventrals 264 in shields slightly longer than suture between pre-
males, 229 in females, subcaudals 38 in males, frontals; number of infralabials unknown due to
unknown in females; (7) blotches present on damage, first three contacting first chinshield,
parietals or not and broad cream band crossing third and fourth contacting second chinshield;
snout; (8) light supralabial blotch narrow about chinshields elongate and well differentiated
size of one supralabial; (9) white nuchal collar from gulars; chinshields separated from ventrals
3 dorsals long; black nuchal collar 4 dorsals by six gulars and one preventral; 264 ventrals;
long; (10) dorsum lacking stripes; (11) dorsum 38 subcaudals; anal and subcaudals divided;
cream in preservative (red in life?); (12) ter- dorsals smooth, 15-15-15; no apical pits; tail con-
minal scale entirely cream and bluntly rounded; ical; terminal scale bluntly rounded.
(13) chin and gular region heavily pigmented. Measurements: SVL 362, tail length 49; head
Apostolepisdorbignyiis likely to be confused length 5.81; head width 3.98; eye-nostril dis-
with A. assimilisand A. cearensis.All three species tance 1.48; eye diameter 0.91; frontal length
lack stripes and have long nuchal collars. Apos- 2.39; suture between parietals 2.16; suture be-
tolepis dorbignyidiffers from both species (char- tween prefrontals 1.14; suture between first pair
acteristics in parentheses) in lacking temporals of chinshields 1.48.
(temporals 0 + 1) and in having only three Color in preservative: Dorsal surface of head
(four) infralabials contacting the first pair of dark brown (black several years after being pre-
chinshields. In addition, A. dorbignyihas a cream served; Schlegel, 1837; Dumeril et al., 1854);
terminal scale (black in A. assimilis) and a single, elongate cream blotch on each parietal
rounded snout (pointed in A. cearensis).Below, paralleling interparietal suture; broad cream
A. dorbignyiis compared to a new species, A. mul- blotch across snout enclosing first supralabials,
ticincta. nasals, two-thirds of prefrontals, and part of ros-
tral; cream nuchal collar extending for 3.0-3.5
Redescriptionof the holotype.-A male; tail short, dorsals, followed by brown collar 4 dorsal long;
13.5% of total length; head short, 1.1% of SVL; chin and gular regions heavily pigmented; in-
head not distinct from neck, its width 69% of complete gular band present; cream supralabial
its length; pupil round; eye-nostril distance 25% blotch narrow, restricted to supralabial 4 and
of head length; snout round in dorsal and lat- caudal edge of 3; dorsal and ventral body uni-
eral view; rostral only slightly visible from above, formly cream (dorsum red and venter cream
its dorsal aspect smaller than mental; length of several years after being preserved; Schlegel,
rostral visible from above less than one-half 1837; Dumeril et al., 1854); caudal fifth of tail
(about 40%) its distance to frontal; prefrontal- black (6 subcaudals black); terminal scale and
rostral contact broadly separating nasals; suture 38th pair of subcaudals cream.
between prefrontals about half as long as hex- Variation: A female (MZUT 963) differed
agonal frontal; frontal about two-thirds as wide from the holotype in having 1 preventral, 229
as long; parietals about half as wide as long; ventrals, 2-3/2-3 supralabials entering eye, and
length of suture between parietals 90% length the first pair of infralabials contacting each oth-
of frontal; nasal divided ventral to nostril, sep- er. This specimen had lost part of its tail ex-
arated from preocular by broad contact be- plaining the subcaudal count of "13/13 + N"
tween prefrontal and second supralabial; preo- provided by Peracca (1897) and atypically low
cular rectangular, about one-fifth length of na- for the genus. The specimen's head is extremely
sal; naris centrally positioned in nasal and di- desiccated and measurements of it were not tak-

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396 COPEIA, 1999, NO. 2

Fig. 8. Distributionof Apostolepis


in Bolivia.Stip-
pled areas are above 1000 (darkergray) and 500 m.

en. Snout-vent length is 449 mm. Unlike the


holotype, the parietals are immaculate black. A
light band across the snout is evident though
discolored probably as an artifact of fixation
and/or desiccation.

Probable distribution and remarks.-The locality


for the holotype of A. dorbignyiis almost certain- Fig. 9. Dorsal and ventral views of Apostolepis
mul-
ticincta(Holotype, NK 729).
ly in error because no other specimens of the
genus are known from trans-Andean localities.
Koslowsky (1898) speculated that the type of the Museum of Comparative Zoology by A.
this species actually came from Mato Grosso or Amaral (MCZ 20763, 20765) are actually A. as-
eastern Bolivia. MZUT 963 was collected by Al- similis. Claims that the species occurs in Brazil
fredo Borelli from the Bolivian Chaco (Depar- and Paraguay (Amaral, 1930c; Peters and Ore-
tamento Tarija) at Aguairenda (Peracca, 1897; jas-Miranda, 1970; Lema, 1978) similarly are not
Fig. 8). based on museum specimens of A. dorbignyito
The key provided by Lema (1978) states that my knowledge.
A. dorbignyihas two longitudinal black lines (his
couplet, 20, p. 40). Tracing backward in his key, ApostolepismulticinctaNew Species
A. dorbignyiis reached by passing through cou-
Figure 9
plet 12. However, the information there is also
at odds with the only known specimens: the ho- Holotype.-Museo Noel Kempff Mercado (NK)
lotype and MZUT 963 have three not four in- 729 (field number IF 82), an adult male col-
fralabials contacting the first pair of chinshields lected in the vicinity of Pampagrande, Provincia
and lack longitudinal dorsal stripes. Florida, Departamento Santa Cruz, Bolivia by
Amaral (1930a, 1935) has remarked that A. native collectors in 1996 for Lucindo Gonzales
dorbignyiis relatively common in southern Brazil A. and Ingid Fernandez S.
begging the conclusion that he mistakenly was
referring to some other species. Specimens orig- Paratypes (2).-NK 878 and ZFMK 66375 both
inally identified as A. dorbignyiand deposited in juvenile males from Pampagrande and SanJuan

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HARVEY-BOLIVIAN APOSTOLEPIS 397

del Portrero, Provincia Florida, Departamento and slender. Head length of the holotype of A.
Santa Cruz, Bolivia, respectively. dorbignyi (5.81 mm) is the same or about the
same as that of the paratypes of A. multicincta.
Referredspecimens(5).-NK 1313, 1369, 1461-62, However, the paratypes are almost one-half the
1486 all collected by native collectors in the vi- length of the holotype of A. dorbignyi.The labial
cinity of Pampagrande. These specimens were blotch is more extensive in the new species
not included in the type series, because they (one-half of supralabial 3, all of 4, and one-third
were not made available to me. They were com- of 5) than in A. dorbignyi(one-third of 3, two-
pared to a description of the types by L. Gon- thirds of 4, and none of 5); the chin is lightly
zales, and noteworthy variation among the re- pigmented in the new species, but very dark in
ferred specimens is discussed below. A. dorbignyi;and the lightly pigmented band
across the snout is broader in the new species
Definition.-A small species of Apostolepis,SVL to where it occupies all of the prefrontals and the
333 mm, differing from all congeners by the fol- anterior one-third of the frontal compared to
lowing combination of characteristics: (1) snout one-half of the prefrontals and none of the
bluntly rounded, length of rostral visible dor- frontal in A. dorbignyi.
sally about equal to one-half its distance to fron- The new species might also be confused with
tal; (2) six supralabials; (3) nasal and preocular other species having uniformly red dorsa and
separated by prefrontal; (4) temporals absent; blunt snouts. Apostolepisassimilis differs from A.
(5) three infralabials contacting first pair of multicincta (characteristics in parentheses) in
chinshields; (6) ventrals 212-233, subcaudals having a black tail tip (terminal scale entirely
31-40; (7) dorsal head plates immaculate black white or entirely gray and white), four infrala-
except for broad gray band (white in juveniles) bials contacting the anterior chinshields
across snout; (8) light supralabial blotch mod- (three), and a posterior temporal (temporals
erate, occupying part or all of 3 supralabials; (9) absent). Apostolepisflavotorquata differs from A.
white nuchal collar 2.0-2.5 dorsals long edged multicincta (characteristics in parentheses) in
by narrow gray (white in juveniles) collar 0.5 having the black neck band irregular and to 2
dorsals long; black nuchal collar 4 dorsals long; scales long (4 scales long), preocular-nasal con-
(10) dorsum lacking stripes; (11) dorsum red; tact (scales separated by broad prefrontal-su-
(12) terminal scale mostly gray (white in juve- pralabial contact), and in having four infrala-
niles), bluntly pointed; (13) chin nearly immac- bials contacting the anterior chinshields
ulate; gular region not pigmented. (three). Apostolepiscearensishas temporals (tem-
Unlike any other species of Apostlepis,the ho- porals absent), four infralabials contacting the
lotype of A. multicincta has a gray band across chinshields (three), and a pointed snout
the snout, posterior to the white nuchal band, (round).
and across most of the proximal portion of the
terminal scale. However, these bands are not Descriptionof holotype.-An adult male; tail short,
present in the paratypes presumably due to ei- 15% of total length; head short, 2.3% of SVL;
ther polymorphism or ontogenetic variation. head not distinct from neck, its width 65% of
Apostolepismulticinctais most likely to be con- its length; pupil round; eye-nostril distance 24%
fused with A. dorbignyi,and the two species share of head length; snout round in dorsal and lat-
several apomorphic characteristics suggesting eral view; rostral clearly visible from above, its
that they are each other's closest relatives. Ex- dorsal aspect about equal to mental; length of
cept for the band crossing the snout of the ho- rostral visible from above just greater than one-
lotype, the gray bands and red dorsal color have half (about 70%) its distance to frontal; pre-
faded. It is unlikely that in life A. dorbignyihas frontal-rostral contact broadly separating nasals;
a gray band across its snout. Schlegel's (1837) suture between prefrontals 44% of the length
description indicates that the red color had not of hexagonal frontal; frontal about two-thirds as
faded at the time he described the holotype; he wide as long; parietals about half as wide as
relates that the band across the snout was pure long; length of suture between parietals 92% of
white. the length of frontal; nasal divided ventral to
The eight known specimens of A. multicincta naris, separated from preocular by broad con-
(the types are males, sex of referred specimens tact between prefrontal and second supralabial;
is unknown) have many fewer ventrals (212- preocular rectangular, about one-fifth length of
233, x = 225, SD = 6.98) than male A. dorbignyi nasal; naris centrally positioned in nasal and di-
(266), and these counts reflect the difference rected posterolaterally; 6/6 supralabials: first
in habitus between the species. The new species contacting rostral and nasal; second contacting
is stocky and short, whereas A. dorbignyiis long nasal, prefrontal, and preocular; second and

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398 COPEIA, 1999, NO. 2

third entering orbit; third and fourth contact- three (NK 878) or none are edged in black. The
ing single subrectangular postocular; fourth tip of the tail, the nuchal collar, and the pre-
through sixth contacting parietal. Temporals frontal band are white rather than gray.
absent; three occipitals; median occipital posi- The referred specimens have 212-233 ven-
tioned between caudal tips of parietals and trals and 31-37 subcaudals. The prefrontal
same size as adjacent dorsals; lateral occipitals band overlaps the prefrontals and anterior part
enlarged, each filling space just caudal to con- of the frontal; the supralabial blotch covers part
tact of sixth supralabial with parietal; mental or all of supralabials 3-5. The gular region and
slightly longer than wide, narrowly contacting chin are nearly immaculate except for black pig-
chinshields; strong mental groove; length of su- ment bordering the chinshields on the first
ture between first pair of chinshields nearly three infralabials. The tip of the tail is white in
twice as long as suture between prefrontals; 6/ all specimens (in NK 1486, the tip of the tail is
6 infralabials, first three contacting first chin- missing); 3-4 subcaudals are black. NK 1486 has
shield, third and fourth contacting second chin- 6/7 infralabials resulting from apparent fractur-
shield; chinshields elongate and well differen- ing of the first supralabial.
tiated from gulars; chinshields separated from Measurements of the holotype followed by
ventrals by five gulars and two preventrals; 219 those of the paratypes (NK 878 then ZFMK
ventrals; 40 subcaudals; anal and subcaudals di- 66375) in parentheses: SVL 333 (200, 188), tail
vided; dorsals smooth, 15-15-15; no apical pits; length 51 (22, 21); head length 7.51 (5.81, 5.7);
tail conical; terminal scale bluntly pointed. head width 4.89 (4.44, 4.5); eye-nostril distance
Hemipenis only partly everted (proximal one- 1.82 (1.37, 1.25); eye diameter 0.91 (0.68, 0.7);
third visible); base of hemipenis covered with frontal length 2.84 (2.16, 1.9); suture between
small spines on both sides; larger spines distal parietals 2.62 (2.28, 1.9); suture between pre-
on asulcate side. frontals 1.25 (1.02, 0.9); suture between first
Color in life: Dorsal surface of head immac- pair of chinshields 2.16 (1.59, 1.7).
ulate black except for wide gray band across
snout enclosing first supralabials, nasals, pre- Etymology.-The specific epithet multicinctais de-
frontals, and part of rostral and frontal; trian- rived from the Latin noun cinctum meaning
gular black blotch in center of rostral; white nu- band or belt and multus meaning many.
chal collar extending for 2.0-2.5 dorsals, fol-
lowed by gray nuchal collar 0.5 dorsals long, Distribution.-Most specimens of A. multicincta
then by black collar 4-5 dorsals long; chin and were obtained from native collectors in the vi-
gular regions immaculate except for black pig- cinity of Pampagrande (Bolivia: Santa Cruz:
ment along medial and anterior edges of in- Florida; 18?5'S; 64?6'W) at an elevation of ap-
fralabials; gular band absent; white supralabial proximately 1290 m (Fig. 8). One paratype
blotch moderate, enclosing supralabial 4 and (ZFMK 66375) came from the nearby town of
covering part of supralabials 3 and 5 and postoc- San Juan del Portrero (17?53'S; 64?17'W), a site
ular; dorsum red; venter cream; caudal fifth of located at about the same elevation and with
tail black dorsally; six subcaudals edged in black similar climate and vegetational composition.
laterally; terminal scale gray. Navarro (1994) pointed out the vegetational
similarity of this area to that of the Chaco. Data
Variation.-The paratypes (NK 878 followed by from the nearby meteorological station in Mair-
ZFMK 66375) have 225 and 233 ventrals, 32 and ana (45 km east of Pampagrande and at 1350
33 subcaudals; all other scale counts are the m) underscore the subhumid conditions in this
same. Their rostrals are less prominent dorsally: intermontane valley: average annual tempera-
rostral length is equal to one-half its distance ture is 20.9 C and average annual precipitation
from the frontal. The head of the holotype may 575.4 mm. Much of the natural habitat around
have been partially depressed upon capture, Pampagrande has been altered by agricultural
and the extent of that specimen's rostral expo- practices.
sure may not be representative of the species.
The paratypes' mentals partially separate the Remarks.-In the comparisons section, we em-
first infralabials but fall short of the chinshields. phasize the similarity of A. multicincta to the
The paratypes are nearly identical in colora- Chacoan species A. dorbignyi.These two species
tion to the holotype. Black pigment of the chin share several derived characteristics (based on
and gular regions is restricted to the anterior outgroup comparisons with other elapomor-
edges of the second supralabials in NK 878 and phines) that suggest they are each other's clos-
the medial edges of supralabials 1-3 in ZFMK est relatives. No other species of Apostolepishas
66375. Three pairs of subcaudals are black, and been collected above 500 m. However, an Apos-

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HARVEY-BOLIVIAN APOSTOLEPIS 399

tolepis closely related to a Chacoan congener


might have been predicted to occur within the
high, intermontane valleys of Santa Cruz. Sev-
eral reptiles and amphibians from the Chaco
also occur in this region. These populations are
effectively cut off from Chacoan populations by
the forested, eastern slopes of the Andes. Many
of these populations appear to have diverged
little, presumably because ecological barriers do
not affect them as significantly. However, several
lineages appear to have only recently diverged
into sister species. Several such pairs of species,
the first endemic to these Andean valleys and
the second occurring in the Chaco can be iden-
tified: Apostolepis multicincta and A. dorbignyi,
Bothropsjonathani and Bothropsalternatus (Har-
vey, 1994), Philodryas levisquamusand Philodryas
aestivus (Thomas, 1976; Thomas et al., 1999),
Fig. 10. A specimen of Apostolepis nigroterminata
Amphisbaena cegei and Amphisbaena darwini (UTA 44687) from Santa Cruz, Bolivia.
(Montero et al., 1997). In each of these pairs,
the species have only differentiated slightly,
strongly suggesting a common history to the head and neck, but not over the rest of the
faunas of the Chaco and these high Andean val- body, occur in other colubrids such as Scaphio-
leys. dontophisannulatus dugandi (Henderson, 1984).
In addition to A. multicincta, several conge-
ners such as A. assimilis and A. dorbignyihave
ApostolepisnigroterminataBoulenger, 1896
apparently aposematic coloration. Species of Figure 10
Apostolepisare rear-fanged colubrids with large
Duvernoy's glands. Although bites from Aposto- Apostolepis nigroterminata Boulenger, 1896:235.
lepis are unknown, some species such as A. as- Holotype BMNH 1946.1.9.77 (originally
similis feed on amphisbaenians and snakes (Sav- BMNH 81.5.13.76); type locality "Cayaria" (=
itsky, 1979), and they are likely venomous. Tha- Callaria, Ucayali, Peru). Amaral, 1930a:109;
les de Lema nearly died from a bite by another 1930b:48; 1930c:226; 1935:149; Peters and
elapomorphine, Phalotris lemniscatus(R. Fernan- Orejas-Miranda, 1970:23; Lema, 1978:30; Fug-
des, pers. comm.). ler, 1986:46.
Nuchal and caudal banding in some Aposto- Apostolepis borelli Peracca, 1904:9. Holotype
lepis is reminiscent of the banding in sympatric MZUT 962; type locality Urucum, Mato Gros-
coral snakes. Some other colubrids such as Ni- so, Brazil. Amaral 1925:8; Parker, 1928:98;
nia sebaeand juveniles of several species of Clelia Ferrarezzi 1993a:237.
have nuchal banding and immaculate red dorsa
(Campbell and Lamar, 1989) yet are often pre- Justification of synonymy.-Boulenger (1896) de-
sumed to be mimics of sympatric coral snakes. scribed A. nigroterminatafrom "Cayaria" in what
The head of A. multicinctaclosely resembles the is now the department of Ucayali, Peru. Amaral
only species of coral snake known from Pam- (1930b, 1930c) examined the types of A. nigro-
pagrande, Micrurus frontifasciatus (Campbell terminataBoulenger and A. borelliPeracca as well
and Lamar, 1989, provide a color photo of this as specimens identified as A. borellifrom Buena
species). Presence of a prefrontal band varies Vista, Bolivia (Parker, 1928) and found no con-
among species in both Apostolepisand Micrurus, vincing difference to separate them. His deci-
sympatric A. multicincta and M. frontifasciatus sion was followed by Lema (1978) who com-
both have such a band, and the dorsal surfaces pared the holotype of A. borellito specimens also
of their heads are otherwise immaculate black. examined in this study (AMNH 87942, FMNH
Interestingly, the terminal scale of A. multicincta 39646, and UMMZ 60773, 67962-63). Ferrarezzi
is entirely white or gray and white, and this "ex- (1993a) did not follow this synonymization and
tra" band makes the tail of this species appear cited the mental-chinshield contact as a differ-
more similar to that of coral snakes. In most ence between the taxa. He also cited differences
congeners, the terminal scale is mostly black in coloration: black tail tip in A. nigroterminata,
and the white portion is ventral and terminal. white in A. borelli;white labial blotch restricted
Similar patterns of banding on the tail and the to 4th supralabial in A. borellibut on third and

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400 COPEIA, 1999, NO. 2

fourth in A. nigroterminata;longitudinal lines oc- nigroterminata.They also have no, or vestigial


cupying dorsals 2-4 in A. nigroterminatabut only white nuchal collars, whereas the collar is dis-
4 and part of 3 in A. borelli.Contra the descrip- tinct although often narrow in A. nigroterminata.
tion in Ferrarezzi (1993a), the terminal scale of At midbody, the lateral stripe begins on the up-
the holotype of A. nigroterminatais mostly white per half of dorsal 1 in A. nigroterminatabut on
with a band of melanophores dorsally and prox- the upper half of dorsal 2 among snakes in the
imally (Boulenger, 1896, states "end of tail A. quinquelineatacomplex. Finally, snakes of the
black; lower parts white"). North of Buena Vis- A. quinquelineata complex have a more robust
ta, size of the labial blotch appears to decrease habitus very similar to Elapomorphusand Phalo-
clinally in populations of this species (see be- tris. Apostolepisnigroterminataand the vast major-
low). Width of the lateral band appears to be ity of Apostolepisare more gracile. Rarely, A. fla-
polymorphic in this species. A specimen from votorquata has dorsal stripes (e.g., AMNH
Mato Grosso (AMNH 87942) had the lateral 93559), however this species has a higher num-
band on dorsal 4 and part of 3, whereas the ber of ventrals (253-257 in specimens examined
lateral band covers dorsals 1-4 in the holotype in this study) than A. nigroterminataas well as
and all other specimens examined. Ferrarezzi many of the same characteristics that serve to
(1993a) did not examine the holotype of A. dor- distinguish the A. quinquelineatacomplex from
bignyi and was apparently unaware that it also A. nigroterminata:a robust habitus, temporals
has the first pair of infralabials separated by the usually present.
mental. Polymorphism in this character occurs Apostolepisniceforoiwas described from La Ped-
in A. multicincta;in NK 878 the mental is sepa- rera, Bajo Caqueti, Colombia, and its range
rated from the chinshields, in NK 729 and might overlap that of A. nigroterminata.The ho-
ZFMK 66375 (Fig. 3A), the mental contacts the lotype and only known specimen of A. niceforoi
chinshields. Complete separation of the first in- differs from A. nigroterminata(characteristics in
fralabials also appears rarely among specimens parentheses) in having 248 ventrals (200-231)
of Micrurus, especially Micrurus mipartitus and 0 + 1 temporals (usually 0 + 0; Amaral,
(Amaral, 1926, 1930b; Roze, 1996). Based on 1935). Amaral (1935) reports seven stripes
these observations, no reliable differences exist (five), the first reportedly beginning on row 2
to justify recognition of A. borellias a valid spe- (row 1) as in snakes of the A. quinquelineata
cies. complex. A complete nuchal collar is absent
from his illustration of the holotype of A. nice-
Definition.-A small species of Apostolepis,SVL to foroi.
370 mm, differing from all its congeners by the
following combination of characteristics: (1) Redescriptionof the holotype.-The holotype of A.
snout blunt, extent of rostral visible from above nigroterminatabears a faded, albeit embossed,
less than one-half its distance to the frontal; (2) tag from the British Museum of Natural History:
usually six supralabials; (3) nasal usually con- 1946.1.9.77. The specimen was collected by W.
tacting preocular; (4) temporals usually absent; Davis from "Cayaria."
(5) four infralabials contacting first pair of chin- A subadult male; tail short, 8.3% of total
shields; (6) ventrals 200-231, subcaudals 23-33; length; head not distinct from neck, its width
(7) light blotches present on dorsal surface of 90% of its length; pupil round; eye-nostril dis-
head; (8) light supralabial blotch present; (9) tance 21% of head length; snout round in dor-
white nuchal collar 1-3 dorsals long followed by sal and lateral view; rostral only slightly visible
narrower (0.5-2.0 dorsals) black nuchal collar; from above, its dorsal aspect smaller than men-
(10) five dorsal stripes: lateral stripe widest and tal; prefrontal-rostral contact broadly separating
occupying scale rows 1-4, rarely ventrals to row nasals; suture between prefrontals about half as
4 or restricted to rows 3-4; narrow paravertebral long as hexagonal frontal; frontal about half as
stripe on rows 5 and 6; narrow vertebral stripe wide as long; parietals about half as wide as
on row 8; (11) terminal scale black with a cream long; length of suture between parietals 88% of
tip; (12) dorsum brown and tan; (13) extent of the length of frontal; nasal entire, broadly con-
chin and gular pigmentation variable. tacting single preocular; preocular rectangular,
Apostolepisnigroterminatais most likely to be about one-third length of nasal; naris centrally
confused with the Amazonian snakes of the A. positioned in anterior half of nasal, directed
quinquelineata complex. These species usually slightly lateral of anterior; 6/6 supralabials: first
have temporals (as in a series near the Bolivian contacting rostral and nasal; second contacting
border in Rondonia; Da Silva, 1993) or the tem- nasal and preocular; second and third entering
poral fused with the last supralabial, whereas orbit; third, fourth, and fifth contacting single
temporals are absent in most specimens of A. round postocular; fifth and sixth contacting pa-

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HARVEY-BOLIVIAN APOSTOLEPIS 401

rietal. Temporals absent; three occipitals; me- as the fifth tooth. Bolivian and Peruvian speci-
dian occipital positioned between caudal tips of mens have spines covering the basal portions of
parietals and same size as adjacent dorsals; lat- the hemipenes (examined in situ); the largest
eral occipitals enlarged and filling space just spines on the asulcate side are equal in length
caudal to contact of sixth supralabial with pari- to the width of a subcaudal.
etal on either side of head; mental slightly lon- Due to excessive fading of the holotype, sev-
ger than wide, contacting chinshields; strong eral aspects of coloration could not be assessed.
mental groove; length of suture between first Most significantly, black pigmentation had fad-
pair of chinshields equal to suture between pre- ed to brown and was indistinguishable from the
frontals; 7/7 infralabials, first four contacting ground coloration in places such as the head.
anteriormost chinshield, fourth and fifth con- The following description of a recently collect-
tacting second chinshield; chinshields elongate ed Bolivian specimen (UTA 44687) provides a
and well differentiated from gulars; chinshields more detailed depiction of the species's appear-
separated from ventrals by six gulars and one ance and reveals some noteworthy variation in
preventral; 207 ventrals; 26 subcaudals; anal and color pattern within this species: dorsal surface
subcaudals divided; in situ hemipenis (left) ex- of head tan with black reticulation on parietals;
tending 7 subcaudals; dorsals smooth, 15-15-15 edges of other dorsal head plates black; irreg-
reducing to eight at level of tenth subcaudal; no ular blotch occupying caudal third and center
apical pits; tail conical except for laterally com- of frontal; rostral and anterior half of nasals
pressed terminal scale; terminal scale short and cream except for vertical black band in center
pointed. of rostral; prominent white labial spot on parts
Measurements: SVL 206, tail length 17; head of supralabials 3 and 5 and all of 4; anterior half
length 5.55; head width 4.97; eye-nostril dis- of first supralabial cream; remaining supralabi-
tance 1.16; eye diameter 0.58; frontal length als black with tan blotches on supralabials one
1.97; suture between parietals 1.73; suture be- and two; ventral aspect of head white with black
tween prefrontals 1.04. My measurement of pattern; mental and first four infralabials edged
head width may not accurately represent the in black and with black blotches; first and sec-
species. The upper jaw of the holotype has been ond pairs of chinshields edged in black medi-
extensively dissected, and the head likely ap- ally; black interrictal band occupying third pair
pears wider than it was in life. of chinshields and one to two rows of gulars;
Color in preservative (the specimen is faded occipitals and first row of dorsals black; white
and stored in 70% alcohol): Dorsal surface of neck collar one scale row long; white collar fol-
head brown with irregular cream blotches on all lowed by an incomplete black collar one scale
large cephalic plates; cream blotches particular- row long extending dorsally from first to sev-
ly prominent on prefrontals, supraoculars, and enth dorsals; vertebral stripe beginning in mid-
parietals; cream nuchal collar extending for 2- dle of eighth dorsal within last black collar; ven-
3 dorsals and edged posteriorly by irregular ter and half of first row of dorsals cream ante-
brown collar 0.5 to 1 dorsal long; chin and gular riorly, gradually becoming light gray at 40th
region immaculate cream; cream supralabial ventral; charcoal band extending from middle
blotch occupying supralabials 3-4/3-4; dorsum of first to middle of fourth dorsal scale rows;
bearing five stripes: pair of lateral bands cover- dorsal margin of band edged in black; fifth
ing dorsals 1-4, pair of narrow paravertebral through eighth scale rows tan; narrow black ver-
stripes along junctions between dorsals 5-6, and tebral stripe; tip of tail white.
narrow vertebral stripe along dorsal 8; brown Most specimens of A. nigroterminata come
band encompassing caudal fourth of tail (6/7 from a single locality, Buena Vista, allowing a
subcaudals black); terminal scale cream with rare chance to examine intrapopulational vari-
band of melanophores dorsally and proximally. ation within a species of Apostolepis.Within the
population from Buena Vista, the chin ranges
Variation.-The upper jaw of the holotype has from immaculate (e.g., UMMZ 60773) to mod-
been heavily dissected by earlier researchers erately pigmented (e.g., UMMZ 67962) to hav-
and its teeth are no longer present. However, ing black pigment on many of the infralabials
the maxillary teeth of the other specimens from and chinshields (e.g., NK 472). A gular band is
the British Museum and from UTA were ex- present in 3 specimens (BMNH 1927.8.1.180,
amined. In each of these four specimens, the 182, and NK 472), as well as in the single spec-
first five maxillary teeth increase in size poste- imen from nearby Finca Dos Milanos. All the
riorly (1 < 2 = 3 < 4 < 5). A short diastema other specimens lack a gular band. The suprala-
separates the fifth tooth from two large, bial blotch occupies supralabials 3-6 on both
grooved fangs, each about three times as long sides of all specimens from Santa Cruz, Bolivia.

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402 COPEIA, 1999, NO. 2

Size of this blotch decreases in specimens north Distribution.-Brazil. Estado Mato Grosso: (con-
of Santa Cruz, occupying supralabials 2-4 in a fluence of Rio Araguaia and Tapirape, Tapirape
specimen from Beni and 3-4 in the two Peru- Village) AMNH 87942, (Urucum, holotype of A.
vian specimens. borellinot examined herein) MUZT 962.
A second temporal (0 + 1) is present on both Bolivia (Fig. 8). Departamento Beni: (Prov-
sides of the head in one of the specimens from incia Vaca Diez, Tumi Chucua) USNM 280371;
Buena Vista (BMNH 1927.8.1.181) and on one Departamento Santa Cruz: (Provincia Chiqui-
side only of the head in the Brazilian specimen tos, Finca Dos Milanos) UTA 44687; (Provincia
(AMNH 87942). All other specimens lack tem- Ichilo, Buena Vista) BMNH 1927.8.1.180-182,
porals. One specimen (BMNH 1927.8.1.182) CM 2909, UMMZ 60773, UMMZ 67962-63;
has the fourth through the sixth supralabials (Provincia Nuflo de Chives, Las Trancas) NK
contacting the parietals on both sides, another 942; (Provincia Velasco, El Remanso) NK 472.
(CM 2909) has the fourth contacting the pari- Peru. Departamento Ayacucho: (La Mar, Siv-
etal on one side only; all other specimens from ia on Apurimac River, approximately 760 m.
Buena Vista have the fifth only or fifth and sixth above sea level) FMNH 39646; Departamento
supralabials contacting the parietals. Ucayali: (Callaria) BMNH 1946.1.9.77.
Obvious scale anomalies are present in one
specimen (NK 942). Supralabials 1-4 are fused Remarks.-The specimen from Beni may repre-
on one side of the head, and the mental is fused sent a cryptic species but, in the absence of
to the first infralabial on one side and the first more convincing diagnostic features, is here re-
chinshield is fused to the first infralabial on the ferred to A. nigroterminata.Unlike any other
other. Except for these anomalies, all specimens specimens of A. nigroterminata,the Beni speci-
have divided anals, dorsals 15 + 15 + 15, single men has rows of ventrolateral spots and a much
pre- and postoculars, six supralabials, and su- bolder color pattern overall.
pralabials 2-3 entering the eye. Most specimens
have seven infralabials with six and eight infral-
abials occurring once and on one side only. All Apostolepisphillipsi New Species
but one specimen have infralabials 1-5 contact- Figure 11
ing the chinshields; BMNH 1927.8.1.181 has 1- Holotype.-University of Texas at Arlington
6 infralabials contacting the chinshields on one (UTA) 43940, (field number MBH 2471), an
side only. The preocular and nasal are in con- adult female collected by Barbara Phillips on 17
tact in all specimens except FMNH 39646 which September 1993 from the grounds of Estancia
has a loreal on one side only. The lateral band El Refugio, Provincia Velasco, Santa Cruz, Boliv-
encompasses dorsals 1-4 in all specimens ex- ia, 14?45'S; 61?00'W.
cept AMNH 87942 where it encompasses dorsals
3-4. Definition.-A moderately long, gracile species
Noteworthy variation in other meristic and of Apostolepis,SVL 401 mm, differing from all its
morphometric characteristics follow. Means and congeners by the following combination of
standard deviations are reported in parenthe- characteristics: (1) snout pointed, extent of ros-
ses. Unless otherwise noted, ranges are based tral visible slightly greater than one-half its dis-
on specimens examined and do not include tance to the frontal; (2) six supralabials; (3) na-
published reports on this species. Four females sal separated from preocular by contact be-
(CM 2909, UMMZ 60773, 67962, and UTA tween prefrontal and second supralabial; (4)
44687) and 10 males were examined: preven- temporals absent; (5) four infralabials contact-
trals 1-3 (x = 1.5, SD = 0.76); ventrals 200-231 ing first pair of chinshields; (6) ventrals 227,
(x = 211.9, SD = 10.1) in males, 222-231 (x = subcaudals 24; (7) pair of small, cream blotches
225.8, SD = 4.1) in females; subcaudals 26-33 on prefrontals and rostral, remaining dorsal sur-
(x = 28.7, SD = 2.0) in males, 23-26 (x = 25, faces of head dark brown; (8) light supralabial
SD = 1.4) in females; white nuchal collar oc- blotch extensive, covering parts of all suprala-
cupying 1.5-3; black nuchal collar 0-2; tail bials; (9) white nuchal collar 1 dorsal long fol-
length 8-13% (x = 10.4, SD = 1.4) of SVL in lowed by irregular (0.5-2 dorsals) black nuchal
males, 7-9% (x = 8.3, SD = 1.2) in females; collar; (10) five dorsal stripes: lateral stripe wid-
head width 62-82% (x = 71, SD = 6.5, holotype est and occupying scale rows 1-4; narrow para-
and BMNH 1927.8.1.180 excluded) of head vertebral stripe on rows 5 and 6; narrow verte-
length; eye-nostril distance 44-66% (x = 53, SD bral stripe on row 8; (11) terminal scale black
= 7.2) of eye diameter;
length of suture be- with a white tip; (12) dorsum brown and tan.
tween prefrontals 41-62% (x = 54, SD = 5.3) Apostolepisphillipsi is one of the most distinc-
of frontal length. tive species of Apostolepisand is unlikely to be

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HARVEY-BOLIVIAN APOSTOLEPIS 403

'
tebral stripe lies on the sixth row (fifth and
sixth). Finally, Koslowsky (1898) described the
dorsal ground color of A. intermediaas "blanco
sucio" and questioned whether it was red in life
(ground color tan in life, in the new species).
This latter comment and the species's lack of a
white nuchal collar are reminiscent of A. dimi-
diata.

Descriptionof the holotype.-An adult female; tail


short, 7.2% of total length; head not distinct
from neck (Fig. 2B), its width 71% of its length;
pupil round; eye-nostril distance 30% of head
length; snout pointed in dorsal, lateral, and ven-
tral (Fig. 3B) views; rostral clearly visible from
above, its extent visible dorsally slightly greater
than half the length of prefrontal suture; pre-
frontal-rostral contact broadly separating nasals;
suture between prefrontals about 70% of the
length of hexagonal frontal; frontal about 70%
as wide as long; parietals about half as wide as
long; suture between parietals slightly longer
than frontal; nasal entire, separated from single
preocular by broad contact between prefrontal
and second supralabial; preocular rectangular,
about one-fourth length of nasal; naris centrally
positioned in anterior half of nasal, directed lat-
eral of anterior; 6/6 supralabials: first contact-
ing rostral and nasal; second contacting nasal
and preocular; second and third entering orbit;
third, fourth, and fifth contacting single round
fifth and sixth contacting parietal.
Fig. 11. Dorsaland ventralviewsof Apostolepis phil- postocular;
UTA Temporals absent; three occipitals; median oc-
lipsi (Holotype, 43940).
cipital positioned between caudal tips of parie-
tals and same size as adjacent dorsals; lateral oc-
confused with other members of the genus. Su- cipitals enlarged and filling space just caudal to
it resembles A. and contact of sixth supralabial with parietal on ei-
perficially, nigroterminata
snakes of the A. quinquelineata complex with ther side of head; mental slightly longer than
which it is likely sympatric. However, these latter wide (Fig. 3B), separated from chinshields by
medial contact between first infralabials; mod-
species have a rounded snout (Fig. 2C) and
have the preocular and nasal in contact. erate mental groove; length of suture between
I erred in identi- first pair of chinshields 63% of length of suture
Previously (Harvey, 1998),
between prefrontals; 7/7 infralabials, first four
fying this specimen as A. intermedia, a striped
anteriormost chinshield, fourth and
species lacking preocular-nasal contact and contacting
known from the vicinity of Miranda in adjacent fifth contacting second chinshield; chinshields
Mato Grosso, Brazil (Koslowsky, 1898). Unlike elongate and well differentiated from gulars;
A. phillipsi (characteristics in parentheses), A. in- chinshields separated from ventrals by 3 gulars
termedia lacks a white nuchal collar (distinct and one preventral; 227 ventrals; 24 subcaudals;
white collar present) and has only four infrala- anal and subcaudals divided; dorsals smooth,
bials contacting the chinshields (five): 1-3 con- 15-15-15, 6 at level of tenth subcaudal; no apical
tact the first pair (1-4), 3-4 contact the second pits; tail conical except for laterally compressed
pair (4-5). Koslowsky's (1898) description of terminal scale; terminal scale short and round-
the striping pattern is not consistent with the ed (Fig. 5A).
pattern of A. phillipsi in some details: the lateral Measurements: SVL 401, tail length 31; head
band occurs on the third and fourth dorsals length 8.0; head width 5.7; eye-nostril distance
(prominent on second, third, and fourth dor- 2.4; eye diameter 0.8; frontal length 2.4; suture
sals, diffuse on first, Fig. 4A), and the paraver- between parietals 2.5; suture between prefron-

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404 COPEIA, 1999, NO. 2

tals 1.9; suture between first pair of chin shields Apostolepistenuis Ruthven
1.2.
Color in preservative: (Seventy percent eth- Apostolepis tenuis Ruthven, 1927:1. Holotype
anol after unbuffered, 10% formalin. Color has UMMZ 64436; type locality Buena Vista, Santa
not faded from its condition in life). Dorsal sur- Cruz, Bolivia.
face of head dark brown except for pair of small Apostolepisambinigravittata: Amaral, 1930b:51.
cream blotches overlapping suture between ros- Apostolepisambinigra:Amaral, 1930c:224; 1935:
tral and prefrontals; supralabial blotch exten- 147; Peters and Orejas-Miranda, 1970:22.
sive, covering part of supralabials 1-6/1-6; ros- Apostolepistenuis: Hartweg, 1932:103; Peters and
tral cream except for medial brown band; first Orejas-Miranda, 1972:588.
two rows of dorsals posterior to parietals dark Apostolepisambiniger.Fugler, 1986:46.
brown; white nuchal collar 1-2 dorsals long, fol-
lowed by irregular dark brown nuchal collar re- Hartweg (1932) and Peters and Orejas-Miran-
da (1972) did not accept Amaral's synonymiza-
sulting from fusion of stripes, 0.5-2 dorsals
tion of A. tenuis with A. ambiniger.When Fugler
long; dorsum brown and tan; five dorsal stripes
(Fig. 4A); pair of lateral stripes widest, extend- (1986) reported the latter species from Bolivia,
he may have been unaware that A. tenuis had
ing from middle of dorsal 1 to middle of 4, dif-
fuse on dorsal 1, darkest on dorsals 3 and 4; been revalidated. Apostolepis ambiniger occurs
near the border of Bolivia in Brazil and Para-
paravertebral stripes overlapping half of dorsals
5-6; vertebral stripe narrow, occupying middle guay and may eventually be discovered in Boliv-
of dorsal 8; anterior third of venter and chin ia.
pale yellow grading to grayish cream on caudal
two-thirds; ventral surface of tail cream with ir- Definition.-A medium-sized species of Apostole-
regular black blotches on medial halves of most pis, differing from all congeners by the follow-
subcaudals; black caudal band extending for 5/ ing combination of characteristics: (1) snout
7 subcaudals; tip of tail white (Fig. 5A); chin round in dorsal view and in profile, length of
nearly immaculate: anterolateral edges of men- rostral visible dorsally less than one-half its dis-
tal and first and second infralabials edged in tance from frontal; (2) six supralabials; (3) na-
black; dark brown gular band present. First five sal separated from preocular by prefrontal; (4)
maxillary teeth increasing in size posteriorly; temporals absent; (5) three infralabials contact-
short diastema separating fifth tooth from two ing first pair of chinshields; (6) ventrals 245-
large, grooved fangs, each about two and a half 265, subcaudals 37-46; (7) light band across
times as long as fifth tooth. dorsal surface of snout; (8) light supralabial
blotch moderate; (9) white nuchal collar nar-
Etymology.-The specific epithet is a matronym row, 1.0-1.5 dorsals long, black nuchal collar ab-
for Barbara Phillips who discovered A. phillipsi. sent; (10) dorsum with three wide stripes; (11)
Although long interested in natural history and dorsum brown and tan in preservative; (12) ter-
active in conservation efforts in Bolivia, Phillips minal scale mostly white; (13) chin and gular
has recently become interested in herpetology region immaculate or nearly so.
and has greatly assisted in my research on Bo- Apostolepistenuis has been confused with A.
livian reptiles and amphibians. ambinigerand A. vittata by earlier authors. The
latter species both have spade-like snouts,
Distribution.-Apostolepis phillipsi is known only whereas the snout of A. tenuis is round (Fig.
from the type locality (Fig. 8). The holotype of 1A). In addition, A. tenuis has six supralabials
A. phillipsi was collected while digging a drain- (vs five in A. vittata) and is striped (the dorsum
age ditch near Estancia El Refugio on high of A. ambinigeris immaculate except for black
ground previously covered in tropical dry forest. nuchal and caudal bands).
Estancia El Refugio (base camp at 14?45'S,
61?00'W) was previously part of a privately Description.-Peters and Orejas-Miranda (1972)
owned biological reserve but has recently been noted some minor differences between the ho-
incorporated into the Parque Nacional "Noel lotype and a specimen of A. tenuis in the Smith-
Kempff Mercado." El Refugio lies in a flat plain sonian, USNM 123973. I would add only that
with only slight (0.5-10.0 m) elevational relief. the lateral bands break the white nuchal collar
The estancia sits on the north bank of the Rio and extend to the rictus in the USNM speci-
Paragua, and much of the surrounding savanna men, but not in the holotype.
and forest is seasonally inundated. A more de- Apostolepistenuis is an elongate, gracile spe-
tailed description of this site appears in Harvey cies; its head is rounded in dorsal and lateral
(1998). aspects (Fig. 1A) and is about 1-2% of SVL.

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HARVEY-BOLIVIAN APOSTOLEPIS 405

Apostolepistenuis has six supralabials (2-3 or 2- Apostolepis ambinigra var. vittata: Boulenger,
4 enter the orbit), five to six infralabials (1-3 1896:237.
contacting first chinshield, 3-4 contacting sec- Apostolepisambinigra:Amaral, 1930c:224; 1935:
ond), one preocular, one postocular, and no 147.
temporals. The prefrontal separates the nasal Apostolepis vittata: Peters and Orejas-Miranda,
and preocular. The naris is directed anterolat- 1972:588; Lema, 1978:39.
erally and slightly ventral of center in the mid-
dle of the nasal. The tail of A. tenuis is short, Definition.-A medium-sized species of Apostole-
ends in a pointed and laterally compressed ter- pis, SVL to 460 mm, differing from all conge-
minal scale, and is about 10% of the total body ners by the following combination of character-
length. istics: (1) snout strongly projecting, pointed in
The dorsum of A. tenuis is brown and tan in dorsal view and in profile, length of rostral vis-
preservative; however, the two specimens ex- ible dorsally twice its distance from frontal; (2)
amined have faded considerably and live speci- five supralabials; (3) nasal contacting preocular;
mens may have a red ground coloration. In the (4) temporals absent; (5) three infralabials con-
holotype, a triangular, cream band covers the tacting first pair of chinshields; (6) ventrals 228
snout and encloses the first supralabials, nasals, in females, 238-243 in males; subcaudals 23 in
prefrontals, and anterior one-fourth of the fron- females, 27-28 in males; (7) dorsal head plates
tal. This band is smaller in USNM 123973, being
immaculate; (8) light supralabial blotch on su-
restricted to the prefrontal and frontal and cov-
pralabials extensive; (9) white nuchal collar ab-
ering two-thirds of the former and one-fourth sent; black nuchal collar 2-3 dorsals long; (10)
of the latter. Diffuse light mottling covers the dorsum with five stripes; (11) dorsum brown
remaining dorsal head scales. The supralabial and tan in preservative; (12) terminal scale
blotch is small in both specimens and covers
white; (13) chin and gular region immaculate
portions of supralabials 3 and 4. The chin, gular to moderately pigmented.
region, and venter are immaculate in the ho-
Only one species is likely to be confused with
lotype. Some melanistic patches occur on in- A. vittata. Apostolepisgoiasensis is striped dorsally
fralabials 3 and 4 of USNM 123973. A narrow
and has five supralabials. Unlike A. vittata, A.
black stripe runs along the sutures connecting
the subcaudals in both specimens. The white goiasensis has four infralabials contacting the
first pair of chinshields and the nasal separated
nuchal collar is narrow (1.0-1.5 dorsals long); a
black collar is absent. The lateral band increases from the preocular (Prado, 1943; Ferrarezzi,
in width caudally. Just behind the white collar, 1993a). Apostolepisgoiasensis is known only from
it occupies dorsals 3 to the lower half of 6; at the holotype (IB 10260) from Rio Verde, Goias,
Brazil.
midbody it occupies the upper third of 1 to the
lower half of 5. The vertebral band is wide (Fig.
4C), covering all of dorsal 8 and the upper third Description.-Cope's (1887) original description
of 7. The black band on the tail extends for 5 of A. vittata mainly pointed out differences be-
to 7 scale rows dorsally; none of the subcaudals tween it and A. ambiniger.Peters and Orejas-Mi-
are black. The terminal scale is mostly white. randa (1972) provided additional data on the
The following counts and measurements for holotype.
UMMZ 64436 are followed by those of USNM Apostolepisvittata reaches 460 mm; its head is
123973 in parentheses: ventrals 265 (245), sub- pointed in dorsal and lateral profile (Fig. IB)
caudals 37 (46); SVL 308 (168); tail length 36 and is 2.2-3.0% of SVL. Its tail is short, ends in
(14); head length 4.55; head width 3.30; eye- a bluntly rounded to slightly pointed terminal
nostril distance 1.25; eye diameter 0.80; frontal scale and is 7-9% of total body length. The ros-
length 1.48; suture between parietals 1.59; su- tral is pointed and concave ventrally; dorsally,
ture between prefrontals 1.14; suture between its length is twice its distance to the frontal.
first pair of chinshields 0.80. Length of the prefrontal suture is reduced rel-
ative to other species of the genus and is about
Distribution.-Bolivia (Fig. 8): Departamento one-third the length of the frontal. Apostolepis
Beni: (Provincia Vaca Diez: Guayaramerin) vittata has five supralabials (2-3 enter the orbit,
USNM 123973; Departamento Santa Cruz: 4-5 contact the parietal), five to six infralabials
(Provincia Ichilo: Buena Vista) UMMZ 64436. (1-3 contact the first chinshield, 3-4 contact
the second), one preocular, one postocular, and
Apostolepisvittata (Cope) no temporals. The nasal and preocular contact
Rhynchonyxambinigervittatus Cope, 1887:56. Ho- one another. The naris is directed laterally and
lotype ANSP 11293; type locality Chupada, located in the anteroventral corner of the nasal.
Mato Grosso, Brazil. Presumably, the dorsum of A. vittata is brown

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406 COPEIA, 1999, NO. 2

and tan in life; however, all specimens have fad- (Provincia Nuflo de Chaves, Rio SanJulian) CM
ed considerably. A few years after its capture 2824; (Provincia German Busch: Puerto Suarez)
and presumably before it had faded substantial- BMNH 1907.10.31.62.
ly, Cope (1887) described the holotype as hav-
ing "the space between the sixth rows of op- Remarks.-Two Bolivian specimens differ from
posite sides pale brown; external to the sixth the holotype in several noteworthy characteris-
row and below, dirty white." The dorsal head tics and the status of the Bolivian population
surfaces are immaculate brown. A small cream should be reconsidered when larger samples be-
blotch is just visible dorsally and overlaps the come available. The most obvious difference is
lateral side of the rostral and front of the nasal. the vertebral stripe which is wide (Fig. 4B) and
A prominent, subrectangular to somewhat elon- extends to the tip of the tail in both Bolivian
gate supralabial blotch covers portions of su- specimens. In the holotype, the vertebral stripe
pralabials 2-4 or 1-4. The chin is moderately is a thin line like that of A. phillipsi (Fig. 4A),
pigmented to immaculate; a complete gular becomes broken at the level of the 100th ven-
band is present or not. The black nuchal collar tral, and is absent on the tail. Vertebral stripes
encompasses 2-3 rows of dorsals; a white nuchal are very common among species of Apostolepis,
collar is absent. The vertebral stripe is wide cov- species with a solid dorsal pattern such as A.
ering all of dorsal 8 and the upper edge of 7 ambiniger,A. dimidiata, and A. multicinctausually
(Fig. 4B) or a narrow pinstripe down the center have "remnants" of a vertebral stripe extending
of dorsal 8. The lateral band is slightly wider for one or two dorsals just posterior to the black
and may cover the upper one-half of dorsal 3, nuchal band (Fig. 7). A thin vertebral stripe ap-
all of 4, and the lower one-third of 5. A narrow pears in some specimens of A. flavotorquata,
paravertebral stripe covers dorsal 6. All five whereas most have a solid dorsal pattern. Poly-
stripes are prominent anteriorly; they may ex- morphism in width of the vertebral stripe was
tend unbroken throughout the body and tail not observed in any other Apostolepis,however,
(in the Bolivian specimens) or become broken variation in width of other stripes varies widely
posterior to the middle of the body. The black within some species.
band on the tail extends for about 10 subcau- Several additional differences between the
dals dorsally; 5-7 subcaudals are black. The low- Bolivian snakes and the holotype are polymor-
er and posterior one-half of the terminal scale phic in several species of Apostolepis.Unlike the
is white. A narrow black stripe may run along holotype (characteristics in parentheses), both
the sutures between the subcaudals for the Bolivian specimens have six infralabials (five);
length of the tail (CM 2824) or be absent eight scales surrounding the terminal scale
(ANSP 11293; BMNH 1907.10.31.62). (six); a gular band and moderate pigmentation
The following counts and measurements for of the chin (chin and gulars immaculate); the
CM 2824 are followed by those of BMNH lateral stripe occupying one-half of dorsal 3, all
1907.10.31.62, then ANSP 11293 (when avail- of 4, and one-third of 5 (middle one-half of 4,
able) in parentheses: ventrals 228 (243, 238), lower edge of 5; flecks are present on some
preventrals 2 (3, 2), subcaudals 23 (28, 27), SVL scales in dorsal rows 2 and 3); and the suprala-
460 (214, 280), tail length 33 (19, 25); head bial blotch covers part of supralabials 2-4 (1-
length 10.13 (6.49); head width 6.03 (3.76); eye- 4).
nostril distance 2.39 (1.37); eye diameter 1.14
(0.74); frontal length 3.30 (2.16); suture be- ACKNOWLEDGMENTS
tween parietals 2.39 (1.71); suture between pre-
frontals 1.02 (0.46); suture between first pair of During the course of this study, I was provid-
chinshields 1.59 (0.80). ed access to specimens of Apostolepisand work-
ing space at the National Museum of Natural
Distribution.-Apostolepis vittata has not been pre- History, Washington, DC, by R. McDiarmid and
viously reported from Bolivia. Cope (1887:45) R. Reynold; at the American Museum of Natural
lists the type locality for this species as near "the History, New York by D. Frost; and at the Acad-
village of Chupada, thirty miles north-east of emy of Natural Sciences, Philadelphia by T.
Cuyaba, and near the headwaters of the Xin- Daeschler. I thank E. N. Arnold, C. McCarthy,
gu." He was referring to Chapada dos Guimar- E. Censky,J. E. Cadle, W. E. Duellman, D. Frost,
tes near the city of Cuiaba. N. Franzese, I. Ineich, A. G. Kluge, R. Reynolds,
Brazil. Estado Mato Grosso: (near the "vil- D. A. Rossman, M. Suarez R., and H. K. Voris
lage of Chupada, thirty miles north-east of Cuy- for loan of specimens under their care. J. Wil-
aba," Cope, 1887:45) ANSP 11293. liams kindly verified that the holotype of A. in-
Bolivia (Fig. 8). Departamento Santa Cruz: termediahad been lost from the Museo de la

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HARVEY-BOLIVIAN APOSTOLEPIS 407

Plata. I thank T. Schulenberg and A. Forsyth for e Squamata).I. Publ. Avul. Mus. ParaensesEmilio
inviting me to participate in Conservation In- Goeldi 40:9-92.
ternational expeditions to the type locality of A. DA SILVAJR., N. J.. 1993. The snakes from Samuel
Hydroelectric Power Plant and vicinity, Rond6nia,
phillipsi in eastern Bolivia. Finally, I express my Brazil. Herpetol. Nat. Hist. 1:37-86.
sincerest thanks to my good friends B. Phillips
DIXON,J. R., ANDF. S. HENDRICKS. 1979. The worm-
and I. Phillips for their encouragement and lo- snakes (family Typhlopidae) of the Neotropics, ex-
gistical support. I sincerely thank L. Gonzales clusive of the Antilles. Zool. Verh. Rijks. Nat. Hist.
and I. Fernandez and local collectors in Pam- Leiden 173:1-39.
pagrande and A. Langer for collecting the type DOWLING,H. G. 1951. A proposed standard system
series and referred specimens of A. multicincta. of counting ventrals in snakes. Brit. J. Herpetol. 1:
97-99.
, ANDJ. M. SAVAGE.1960. A guide to the snake
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408 COPEIA, 1999, NO. 2

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PARKER,H. W. 1928. Notes on reptiles and batrachi-
E-mail: mbh0417@utarlg.uta.edu. Submitted:
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PERACCA,M. G. 1897. Viaggio del dott. Alfredo Borelli
editor: A. H. Price.
nel Chaco boliviano e nella Republica Agentina.
Boll. Mus. Zool. Anat. Comp., Univ. Torino 12:1-
19.
1904. Viaggio del Dr. A. Borelli nel Matto
APPENDIX
Grosso brasiliano e nel Paraguay, 1899. ix. Rettili
ed amfibii. Boll. Mus. Torino 19 (460):1-15. SPECIMENSEXAMINED
PETERS, J. A., AND B. OREJAS-MIRANDA. 1970. Cata- Museum numbers follow names of species ex-
logue of the Neotropical Squamata. Part I. Snakes. amined and locality information in parentheses.
Smithsonian Institution Press, Washington, DC.
AND . 1972. The taxonomic validity of Many specimens obtained from Alberto and
Apostolepistenuis Ruthven and Apostolepisvittata Adolpho Breyer from La Rioja actually come
(Cope) (Serpentes: Colubridae). Copeia 1972:588-
from adjacent countries (Dixon and Hendricks,
590. 1979; Scrocchi, 1990). No other specimens of
PRADO,A. 1943. Notas ofiol6gicas 14. Comentfrios Apostolepisambinigerare known from Argentina
acerca de algumas serpentes opist6glifas do genero (G. Scrocchi, pers. comm.) and the locality in-
Apostolepis,com a descricao de uma nova especie. formation for USNM 73458 is probably incor-
Mem. Inst. Butantan 16:7-12 (dated 1942). rect.
ROZE,J. A. 1996. Coral snakes of the Americas: bi-
Apostolepisambiniger(4): (Argentina: La Rioja,
ology, identification, and venoms. Krieger Publ.
Co., Malabar, FL. probably in error) USNM 73458; (Paraguay)
RUTHVEN,A. G. 1927. Description of an apparently
MCZ 47002, MVZ 110991, UMMZ 108809.
new species of Apostolepisfrom Bolivia. Occ. Pap. Apostolepis assimilis (13): (Brazil: Bahia)
Mus. Zool. Univ. Mich. 188:1-2. UMMZ 108810-11; (Brazil: Goias) AMNH
SAVITZKY,A. H. 1979. The origin of the New World 87941; (Brazil: Distrito Federal) USNM 148790;
proteroglyphous snakes and its bearing on the (Brazil: Minas Gerais) MCZ 17898; (Brazil: Sao
study of venom delivery sytems in snakes. Unpubl. Paulo) AMNH 102251, MCZ 51493, 20763,
Ph.D. diss., Univ. of Kansas, Lawrence.
20765, UMMZ 204112, USNM 207743 (Brazil:
SCHLEGEL,H. 1837. Essai sur la physionomie des ser-
Unknown) LSUMZ 46269, MVZ 176333.
pens. Partie descriptive. Leide, Arnz, and Compag-
nie, Paris. Apostolepiscearensis(1): (Brazil: Ceara) USNM
56401.
SCROCCHI,G.J. 1990. El genero Micrurus(Serpentes:
Elapidae) en la Repfblica Argentina. Bol. Mus. Apostolepisdimidiata (9): (Brazil: Distrito Fed-
Reg. Sci. Nat. Torino 8:343-368. eral) USNM 148789-90; (Brazil: Mato Grosso)
SERIE,P. 1915. Suplemento a la fauna herpetol6gica AMNH 62192; (Brazil: Sao Paulo) AMNH 7245,
argentina.An. Mus. Hist. Nat. Buenos Aires 27:93- 102252, FMNH 69934, MCZ 27661, USNM
109. 76369-70.
STRAUCH, A. 1884 (printed 1888) Bemerkungen fiber
Apostolepis dorbignyi (2): (Bolivia: Tarija)
die Schlangengattung Elapomorphusaus der Familie MZUT 963; ("Chile") MNHN 3664 (Holotype).
der Calamariden.Mel. Biol. Acad. Imper. Sci. St.
Apostolepisflavotorquata (4): (Brazil: Bahia)
Petersbourg12:141-211. UMMZ 108808; (Brazil: Mato Grosso) AMNH
THOMAS,R. A. 1976. A revision of the South Ameri-
can colubrid snake genus PhilodryasWagler,1830. 93559-61.
Unpubl. Ph.D. diss., TexasA&MUniv.,College Sta- Apostolepislineata (1): (Brazil: Mato Grosso)
tion. ANSP 11211 (Syntype).
,M. B. HARVEY, AND L. DIRKSEN. 1999. A re- Apostolepis multicincta (2): (Bolivia: Santa
considerationof the systematicstatus of Phylodryas Cruz) NK 729 (Holotype), 878 (Paratype).
aestivuslevisquamusThomas, 1999 (Serpentes, Co- Apostolepisnigroterminata(14): (Bolivia: Beni)

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HARVEY-BOLIVIAN APOSTOLEPIS 409

USNM 280371; (Bolivia: Santa Cruz) BMNH Apostolepistenuis (2): (Bolivia: Beni) USNM
1927.8.1.180-1082, CM 2909, NK 472, 942, 123973; (Bolivia: Santa Cruz) UMMZ 64436
UMMZ 60773, 67962-63, UTA 44687; (Brazil: (Holotype).
Mato Grosso) AMNH 87942; (Peru: Ayacucho) Apostolepisvittata (3): (Brazil: Mato Grosso)
FMNH 39646; (Peru: Ucayali) BMNH ANSP 11293 (Holotype), (Bolivia: Santa Cruz)
1946.1.9.77 (Holotype). BMNH 1907.10.31.62, CM 2824.
Apostolepisphillipsi (1): (Bolivia: Santa Cruz) Elapomorphus quinquelineatus (4): USNM
UTA 43940 (Holotype). 6180, 76371-72, 207744.
Apostolepispymi/A. quinquelineata (8): (Brazil: Phalotris bilineatus (2): (Uruguay: Canelones)
Amazonas) AMNH 101954-55, 140772; (Brazil: UTA 7603; (Uruguay: Montevideo) UTA 7604.
Para) KU 127256, 128094, 140153; (Guyana) Micrurus frontifasciatus (2): (Bolivia: Santa
FMNH 26665; (Surinam) FMNH 121828. Cruz) UTA 34561-62.

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