Hollis

Herpetologists' League
Phylogenetics of the Genus Chironius Fitzinger, 1826 (Serpentes, Colubridae) Based on

Morphology
Author(s): Jennifer L. Hollis
Source: Herpetologica, Vol. 62, No. 4 (Dec., 2006), pp. 435-453
Published by: Allen Press on behalf of the Herpetologists' League
Stable URL: http://www.jstor.org/stable/3893601
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Herpetologica, 62(4), 2006, 435-453
? 2006 by The Herpetologists' League, Inc.
PHYLOGENETICS OF THE GENUS CHIRONIUS FITZINGER, 1826
(SERPENTES, COLUBRIDAE) BASED ON MORPHOLOGY

JENNIFER L. HOLLIS
Department of Biology and Museum of Southwestern Biology, University of New Mexico, 167 Castetter Hall,
Albuquerque, NM 87131-0001, USA
ABSTRACT: I present a phylogeny of the Latin American snake genus Chironius based on 36 external
morphological and hemipenal characters. Data from the literature for the 20 recognized species of Chironius
were combined with data I collected from 20 outgroup taxa. I assessed the monophyly and in-group
relationships of Chironius using both parsimony and methods. Two optimal trees were obtained using
parsimony criteria; the genus Chironius is supported as monophyletic with 93% bootstrap support from six
unambiguous synapomorphies, including an even number of scale rows at midbody. A Bayesian analysis of
the data supports the monophyly of Chironius with a posterior probability of 100%. Both analyses inferred
similar topologies; recognized species groups are partially supported, but this analysis suggests that revision of
the genus is necessary. I propose elevation to full species status of taxa within the carinatus, fuscus,
multiventris, and quadricarinatus species groups on the basis of morphological features unique to each taxon.
Future molecular work is necessary to resolve the remaining taxonomic issues within Chironius.
Key words: Chironius; Colubridae; Morphology; Phylogeny; Snakes; Systematics; Taxonomy
COLUBRID snakes of the genus Chironius aredorsal scales is unusual among snakes worlddiurnal, active, and terrestrial-arboreal. They wide; Chironius shares this characteristic only
are visually-oriented hunters that primarily eat
with those taxa of the genus Ptyas that were
small frogs such as Eleutherodactylus; preformerly recognized as Zaocys (e.g., Pope,
dation of lizards and small birds has also been 1934), which usually have 16 scale rows at
documented. Snakes of the genus Chironius
midbody (e.g., Gunther, 1864). Ptyas (Zaocys)
inhabit 17 countries in Central and South
is an Asian rat snake genus related to New
America, and they are distributed from the
World racer and whipsnake genera such as
Caribbean coast of Honduras and Nicaragua
Salvadora and Masticophis (Nagy et al., 2004).
south to northeastern Chile, Argentina, and theThus, among Latin American snake genera, an
Rio de la Plata of Uruguay. Chironius habitat even number of dorsal scale rows are uniquely
varies from lowland rainforests to savannas todiagnostic of Chironius.
montane cloud forests; most species occur in
Dixon et al. (1993) hypothesized that the low
low- to mid-elevation forests. There are
even-numbered dorsal scale-row count for all
currently 20 recognized taxa of Chironius
taxa of Chironius is derived from the slightly
comprising 13 species (Dixon et al., 1993).
higher odd-numbered dorsal scale-row count
A monograph by Dixon et al. (1993) is the
for all species of Dendrophidion; Dendrophionly published study to examine Chironius
dion was proposed as the extant sister taxon to
comprehensively. Species relationships within Chironius. Stuart (1932) suggested that DenChironius were inferred qualitatively on the drophidion was ancestral to most if not all other
basis of morphological characters, but subcolubrid snake genera in Latin America,
species placements were not estimated on the although the paper did not specifically address
resulting tree (Fig. 1). The data were not
the placement of Chironius. Dixon et al. (1993)
analyzed rigorously (i.e., using computer-based
suggested that Dendrophidion was ancestral to
techniques) to form a phylogenetic tree, and Chironius;
no
no data analyses were used to
outgroup taxa were included in the study. support Dixon et al.'s hypothesis. In their
Chironius is morphologically distinct from
scenario, a proto-Dendrophidion stock split
sympatric snake genera: Chironius is the only
into two lineages: one lineage led to Chironius,
Latin American snake genus with either 10 or
and the other lineage led to Drymobius,
12 scale rows at midbody. An even number of
Mastigodryas, and other whipsnakes with an
odd number of dorsal scale rows (as originally

suggested by Stuart, 1932). Chironius and
CORRESPONDENCE: e-mail, jhollis@unm.edu

435
436
HERPETOLOGICA
Chironius laevicollis
Chironius scurrulus
Chironius fuscus
Chironius grandisquamis
[Vol.
62,
No.
collected from 44 alcohol-preserved museum

specimens representing 20 outgroup taxa from
10 genera (Appendix I). Additional outgroup
taxa data were obtained from Lancini (1979)
and Ortenburger (1928) for three characters.
When possible, one to four adult male
specimens were examined from each out-
group species. Exceptions to this are: females

only of Dendrophidion dendrophis (n = 2);
inclusion of one female and one juvenile male
Chironius flavolineatus
of Dendrophidion nuchalis (n = 3); inclusion
Chironius monticola
of one female of Dendrophidion pericarinatum (n = 2); females only of Drymobius
Chironius carnnatus
rhombifer (n = 2); inclusion of one female of
Mastigodryas bifossatus (n = 2); and one
Chironius exoletus
female only for Mastigodryas heathi and
Pseustes sulphureus. In outgroup taxa for
Chironius laurenti
which at least one adult male was available,
four characters known to be sexually dimorChironius vincenti
phic in Chironius (maximum snout-vent
length, number of ventrals, number of subChironius multiventns
caudals, tail length as a percentage of total
length) were scored from males only. Those
Chironius quadricarinatus
characters were scored from males only for all
FIG. 1.-Phylogeny of 13 species of Chironius proposed
Chironius taxa.
by Dixon et al. (1993).
I used Thiele's (1993) method of gapweighting for quantitatively-scored characters
with extrema coded as 0 and 9 for the
Dendrophidion share some morphological
parsimony analysis. Extrema were coded as
features (e.g., relatively long tail length, large
0 and 5 for the Bayesian analysis. Polymorphic
eye size), suggesting that Chironius and
characters were frequency-coded (Wiens,
Dendrophidion may be part of a shared
1995). All characters were weighted so that
lineage, although the shared feature of large
the costs to reach the extrema were equal
eye size could be a product of convergent
regardless of the number of character states.
evolution due to similar ecomorphology (Marx
All characters were ordered except character
and Rabb, 1972). The unique morphological
14 (modal condition of the temporals; see
features of Chironius (i.e., low even dorsal
below), which was scored in the parsimony
scale row counts) support the idea that
analysis using a stepmatrix and was unordered
Chironius is a monophyletic genus, but this
in the Bayesian analysis. All characters were
contention has never been tested.
In this analysis, I use 36 external morpho- given equal weight.
Two approaches were initially used to
logical and hemipenal characters to estimate
assign scores to the data for the parsimony
the phylogeny of all 20 Chironius taxa,
analysis. A data matrix was created in which
evaluate the monophyly of the genus Chironius, and investigate the monophyly of the the gap-weighted characters of outgroup taxa
were scored relative to the range of Chironius
species C. carinatus, C. fuscus, C. multivalues ("Chironius-based matrix"). For examventris, and C. quadricarinatus.
ple, in character 19 (mean ratio of eye
diameter
to snout length; see below), the
MATERIALS AND METHODS
range of values within Chironius is 0.679 to
Characters
0.942. These extrema received scores of 0 and
Morphological data for Chironius from
9, respectively. When outgroup taxa were
Dixon et al. (1993) were combined with data
examined, each taxon with a character value
Chironius bicarinatus
December
20061
HERPETOLOGICA
437
or dissected
hemipenes forDrymar
some species).
less than 0.679
(e.g.,
0.530) was scored as 0 and any taxon with
Characters that could not be scored for
outgroup taxa were retained in this analysis
a character value greater than 0.942 (e.g.,
Dendrophidion pericarinatum, 1.038) refor their value to clarify ingroup relationships.
ceived a score of 9. Outgroup taxa with
One character, mean number of pairs of
intermediate values received the score that
genials, was scored but subsequently discorresponded to that gap weight in Chironius. carded as being uninformative because all
It was expected that such an approach would
but one taxon had two pairs of genials in every
provide the best assessment of ingroup
specimen examined. Fewer than 1% of
relationships as supported by bootstrap values
Chironius exoletus specimens examined (4
(Felsenstein, 1985). A second data matrix was
out of 764 specimens) had three pairs of gecreated in which the data from both in- and
nials; all remaining individuals had two pairs.
outgroup taxa were considered when assigning This character does not appear in the data
gap weighting extrema ("all-taxa matrix"). This matrices in Appendix II and Appendix III.
approach was expected to better establish the
Character Descriptions
monophyly of Chironius because character
values potentially unique to taxa within
Characters 1-34 are drawn from Dixon et
Chironius would receive similar scores, thus
al. (1993); information for characters 35 and
differentiating Chironius as a genus from the 36 was obtained from Dixon et al. (1993) but
outgroup taxa. Preliminary analyses suggested the characters as defined in this study were
that the Chironius-based matrix would not
not used in that monograph. Characters 1-25
produce a tree with significantly better
are used as described by Dixon et al. (1993);
support for ingroup phylogeny than the alloriginal citations for methodology follow
taxa matrix, and internal resolution of some
where appropriate. Counts and measurements
intra-clade relationships was in fact less well
for characters 4, 8-15, 17-20, and 24 were
resolved than on the tree generated from the made on the left side of the animal. Charall taxa matrix. The Chironius-based matrix
acters 26-34 are hemipenal features based on
was recoded for Bayesian analysis but similar the terminology and methods of Dowling and
results were also achieved in preliminary
Savage (1960); the left hemipene was scored
Bayesian analyses. Therefore the results from
when possible.
the Chironius-based matrix are not presented Indicated extrema 0-9 were used in charfor either the parsimony analysis or the
acter state coding for the parsimony analysis;
Bayesian analysis, and the all-taxa matrix wasextrema for the Bayesian analysis were coded
used for all subsequent analyses. The charac- 0-5 (see Appendix III).
ter states assigned to each taxon in the all-taxa
1. Maximum snout-vent length (SVL); 542 mm (0)
matrix are listed in Appendix II (parsimony
to 1813 mm (9). Gap-weighted.
analysis) and Appendix III (Bayesian analysis).
2. Tail length as a percent of total length; 16.2% (0)
StatView 5.0 (Caldarolo et al., 1992) was
to 42.5% (9). This character was not significantly
employed to test four quantitative characters dependent when linearly regressed on SVL (P =
0.3169), so the raw data were scored. Gapfor nonindependence with size using linear
weighted.
regression (see characters 2-4 and 20, below).
3. Mean number of ventrals; 144.5 (0) to 213.0 (9).
Characters and size measurements were

natural-log transformed prior to regression.
One character showed significant dependence
Counted following the method of Dowling (1951).

This character was not significantly dependent
when linearly regressed on SVL (P = 0.2638), so
the raw data were scored. Gap-weighted.
4. Mean number of paired subcaudals; 64.7 to 187.4.
Counted beginning with first scale posterior to the
anal plate and also including the terminal unpaired
scale (Dixon et al., 1993). This character was
on size (character 3; see below); consequently,

the residual values were used in gap-weighting that character.
Some characters could not be scored in
outgroups because of inapplicability (e.g.,
outgroups do not undergo a tail reduction
from six to four scale rows; see character 20) 5.
or lack of material (i.e., unavailability of skulls
significantly dependent when linearly regressed on

tail length (P < 0.0001), so the residuals for the
data were scored. Gap-weighted.
Anal plate condition; single (0) or divided (9).
Frequency coded.
438
HERPETOLOGICA
[Vol.
62,
No.
6. Mean number of scale rows keeled, counted at

midbody; 0 rows (0) to 19 rows (9). Gap-weighted.
7. Apical pits, entirely absent/present on neck only

(0) or present on dorsum (9). Frequency coded.
8. Mean number of supralabials; 6.50 (0) to 9.50 (9).

Gap-weighted.
9. Modal condition of supralabials entering orbit; 3 +

4 (0); 3 + 4 + 5 (1); 4 + 5 (2); 4 + 5 + 6 (3); 5 + 6 (4);
6 (5), in which the formula indicates which
supralabials contact the eye orbit. For example,
3+4 signifies that the third and fourth supralabial
(a)
(b)
(C)
(d)
contact the orbit. Gap-weighted.
10. Mean number of infralabials; 8.50 (0) to 12.50 (9).

Gap-weighted.
11. Mean number of infralabials contacting genials;

5.00 (0) to 7.50 (9). Gap-weighted.
12. Mean number of preoculars; 1.000 (0) to 3.000 (9).
Gap-weighted.
13. Mean number of postoculars; 1.87 (0) to 3.00 (9).
Gap-weighted.
FiG.
2.-Common
(a) 1 + 1 (b) 1 + 2 (c) 1 + 1 + 1 (d) 1 + 1 + 2. Condition
a leads to condition c by a diagonal splitting of the first
temporal; condition b leads to condition d by a similar
fission event. Modified from Dixon et al. (1993).
14. Modal condition of temporal scales; I + 1 (0); 1 + 2

(1); 1 + I + 1 (2); 1 + 1 + 2 (3), in which the formula
indicates the splitting pattern of the temporals (see
Figure 2). Character changes were weighted according to the following matrix:
[0] [1] [2] [3]

[0]
[1]
[2] 1 2 1
[31 2 1 1
15. Mean number of loreals; 0.996 (0) to 2.333 (9).
Gap-weighted.
16. Mean number of postcephalics; 5.93 (0) to 12.33

(9). Gap-weighted.
22. Modal number of dorsal scale rows at midbody; 10

rows (0) to 21 rows (9). Counted over the middle
ventral. Gap-weighted.
23. Modal number of dorsal scale rows anterior to
vent; 8 rows (0) to 17 rows (9). Counted over the
tenth ventral anterior to the anal plate. Gapweighted.
24. Ratio of maximum male SVL to maximum female
SVL; 0.874 (0) to 1.362 (9). Gap-weighted.
25. Ratio of male tail percent to female tail percent;
0.935 (0) to 1.049 (9). Gap-weighted. Data for
outgroup taxa Coluber constrictor constrictor,
Masticophis flagellum testaceus, and Masticophis
17. Mean number of maxillary teeth; 17.0 (0) to 44.2

taeniatus taeniatus are from Ortenburger (1928).
(9). Counted following the method of Myers
26. Mean subcaudal number to which everted hemi(1974). Gap-weighted. Data for outgroup taxa
pene extends; 4.0 (0) to 12.0 (9). Gap-weighted.
Drymarchon corais, Drymoluber dichrous, and
27. Mean subcaudal number at which hemipene
Spilotes pullatus are from Lancini (1979). Data
retractor muscle originates; 16.0 (0) to 31.0 (9).
for outgroup taxa Coluber constrictor constrictor,
Gap-weighted.
Masticophis flagellum testaceus, and Masticophis
28. Termination of sulcus spermaticus; not in naked
taeniatus taeniatus are from Ortenburger (1928).
basal area (0) or in naked basal area (9).
18. Mean number of dentary teeth; 28.2 (0) to 44.3 (9).
Frequency coded.
Gap-weighted. Data for outgroup taxa Coluber
29. Extent of naked basal pocket; small (0) or large
constrictor constrictor, Masticophis flagellum tes- (9). Frequency coded.

taceus, and Masticophis taeniatus taeniatus are
30. Mean subcaudal number to which naked base
from Ortenburger (1928).
extends; 1.0 (0) to 5.5 (9). Gap-weighted.
19. Mean ratio of eye diameter to snout length; 0.530
31. Condition of basal spinules; none (0), minute
(0) to 1.038 (9). Eye diameter is measured
spinules (5), or spinules only slightly smaller than
horizontally; snout length is defined as the
proximal spines (9). Gap-weighted.
straight-line distance from the tip of the snout to
32. Mean number of central spines on dorsal surface
the posterior end of the interprefontal suture
of hemipene; 2.0 (0) to 13.0 (9). Gap-weighted.
(Dixon et al., 1993). Gap-weighted.
33. Mean number of central spines along sulcus
20. Mean subcaudal at which tail reduction from six
spermaticus; 1.0 (0) to 11.0 (9). Gap-weighted.
to four scale rows occurs; 6.0 (0) to 28.5 (9). This
34. Mean number of distal calyces on dorsal surface of
character was not significantly dependent when
hemipene; 3.0 (0) to 25.0 (9). Gap-weighted.
linearly regressed on tail length (P = 0.2352), so
35. Odd versus even modal number of dorsal scale
the raw data were scored. Gap-weighted.
rows, counted at mid-body; odd (0) or even (9).
21. Modal number of dorsal scale rows on neck; 10
Frequency coded.
rows (0) to 23 rows (9). Counted over the tenth
ventral scale posterior to the head. Gap-weighted.
36. Raised vertebral keel; absent (0) or present (9).

Frequency coded.
December
2006]
HERPETOLOGICA
Parsimony
Analysis71 Chironnius multiventris septentnonalis

12 Chironius vincenti
I used PAUP* 4.0blO (Swofford, 2002) to

perform parsimony analyses on the data (e.g.,
Kluge and Farris, 1969). A total of 1000
heuristic random addition searches were
performed on the all-inclusive matrix to
search for the optimal tree, with one tree
held per replicate. A starting tree was
obtained via random stepwise addition; an
optimal tree for each replicate was found
using the tree-bisection-reconnection (TBR)
branch-swapping algorithm.
Clade support was estimated using the
nonparametric bootstrap method (Felsenstein, 1985). Bootstrap values were obtained
from 1000 replicates in which taxa were
randomly added to produce a starting tree.
The optimal tree for each replicate was found
using TBR.
439
79 1 3 Chironius multiventns foveatus
74 Chironius multiventris cochranae

14 Chironius multiventris multiventris
78 15 Chironius laurenti
16 < Chironius fuscus fuscus
Chironius fuscus leucometapus
6 Chironius laevicollis
9 Chironius scurrulus
- 17 Chironius flavolineatus
Chironius cannatus flavopictus

Chironius carinatus spixi
18 Chironius cannatus carinatus

Chironius exoletus
93 Chironius bicarinatus
_ 19 77 Chironius quadncarinatus maculoventris

20 Chironius quadricarinatus quadricarinatus
Chironius monticola
Drymobius marganitiferus
Dendrophidion dendrophis
Dendrophidion nuchalis
Dendrophidion pericarinatum
Drymobius rhombifer
Mastigodryas melanolomus
Mastigodryas heathi
Pseustes poecilonotus
Mastigodryas bifossatus
Mastigodryas boddaerti
Mastigodryas pleei
Pseustes sulphureus
Spilotes pullatus
Drymarchon corais
Characters were optimized in PAUP* using

Drymoluber dichrous
Salvadora grahamiae
ACCTRAN (accelerated transformation) and
63 Salvadora hexalepis
Masticophis flagellum testaceus
DELTRAN (delayed transformation) to genMasticophis taeniatus taeniatus
erate a list of apomorphies for clades in
Coluber constrictor constrictor
Chironius (Appendix IV) with the tree rooted
FIG. 3.-Strict consensus maximum parsimony phylogat Coluber constrictor constrictor.
eny of Chironius taxa and 20 outgroup taxa from 36
morphological characters. Numbers above branches in-
Bayesian Analysis
I used MrBayes 3.1.2 (Huelsenbeck and

Ronquist, 2001) to perform a Bayesian analysis on the data using the Markov chain
Monte Carlo method (e.g., Geyer, 1991;
Mau and Newton, 1997; Mau et al., 1999).
The Markov k (Mk) maximum likelihood
model for discrete morphological data
(Lewis, 2001) was applied to the data
set. Two replicate searches using four
chains for 500,000 generations each were
run, with sampling every 100 generations.
dicate support from 1000 nonparametric bootstraps.

Numbers to the right of splits indicate nodes listed in
Appendix IV (Apomorphy List).
RESULTS AND DISCUSSION
The 1000 heuristic searches in the parsimony analysis inferred two equally parsimonious trees from the same tree-island. Bootstrap values greater than 50% are listed on the
strict consensus of the most parsimonious
trees (Fig. 3) with the following properties:

tree length = 114.9, consistency index (CI) =
Coding was changed to informative and
0.31 (Kluge and Farris, 1969), and retention
the gamma distribution was used for
index (RI) = 0.67 (Farris, 1989). Nonparacharacter rate changes; default priors were
metric bootstrap values at or above 70% are
used for all other settings. Coluber constrictor
considered credible (Hillis and Bull, 1993).
constrictor was designated as the outgroup
The numbers of apomorphies supporting
taxon.
clades and lineages within Chironius are
Log likelihood stationarity was achieved
shown in Fig. 4; although for the most part
within approximately 5000 generations for only unambiguous synapomorphies are dis-
both replicates; burn-in was conservatively

cussed below in relation to clade support,
set at 150. Majority-rule consensus was used
complete information about apomorphies
to estimate a topology from the pool of post- within and supporting Chironius is listed in
burn-in trees. Clade support on the consensus Appendix IV.
tree was estimtated using Bayesian posterior
Posterior probabilities are listed on the
probability values.
majority-consensus cladogram generated in
440
HERPETOLOGICA
42 1
Chironius
[Vol.
multiventris
No.
86 Chironius multiventris septentrionalis

septentrionalis
69 : Chironius vincenti
7412 Chironius multiventbis foveatus

47 Chironius multiventris cochranae
4 / 0 12 Chironius vincenti
13 Chironius muttiventris muttiventris
3/ 2 Chionius mu/tiventns foveatus
6 1 2 13
62,
14 Chironius laurenti
8 /1 Chironius multiventris cochranae
6 Chironius fuscus fuscus
21 14 0_
14 / Chironius multivent/rs multiventns
87 Chironius fuscus leucometapus
73 Chironius laevicollis
8 Chironius scurrulus
15 3 /2 Chironius laurenti
15 Chironius monticola
Chironius carinatus cannatus
10 / 1 Chironius grand/squamis
1 86 Chironius carinatus tiavopictus

100 61 4 Chironius carinatus spixi
3 2 5 Chironius fuscus fuscus

16
Chironius exoletus
07
Chironius
fuscus
leucometapus
Chironius bicarinatus
Chironius flavolineatus
10
Chironius
laevicollis
2/1 7/2
55 Chironius quadricarinatus macubventns

Chironius quadricannatus quadricannatus
Drymarchon corais
Chironius scurrulus
Pseustes sulphureus
76 Spilotes pu/latus
4 / 3 17 1 0 / 2 Chironius flavolineatus
Chiron/us cannatus flavopictus
Drymoluber dichrous
71 Mastigodryas heathi
n7 Dendrophidion dendrophis
Dendrophidion nuchalis
5 0 Chironius cnat/naus spixi
53 Mastigodryas bifossatus
Mastigodryas boddaerti
2 2 3 /2 4 Chironius cnatrnaus carnnatus
2 / 1 5 14/ Chironius exoletus
Dendrophidion pericarinatum
Drymobius margantiferus
Drymobius rhombifer
Masticophis taeniatus taeniatus
Chironius bicat/natus
_ 19 / 5 3 Chironius quadricar/natus maculovent/s

/ Chironius quadt/cat/natus quadt/cat/natus
5/8
Mastigodryas melanolomus
Mastigodryas pleei
Pseustes poecilonotus
96 Sahvadora grahamiae
95 Salvadora hexaiepis
Masticophis flagellum testaceus
Chironius monticola
Coluber constnctor constrictor
FIG. 4.-Apomorphy support within ChironiusFiG.

on the
5.-Majority-rule consensus Bayesian
stnct consensus maximum parsimony cladogram of
Chironius taxa and 20 outgroup taxa from
Chironius. Numbers above branches indicate apomorical characters. Numbers above branches indicate postephies: numbers to the left of the slash indicate
rior probability support from post burn-in pool of trees
apomorphies unambiguous under both ACCTRAN and
(harmonic log likelihood mean = -1549.43; two replicate
DELTRAN optimization; numbers to the right of the
chains, 500,000 generations each, burn-in = 150).
slash indicate apomorphies that occur under DELTRAN
optimization only. Numbers to the right of splits indicate
nodes listed in Appendix IV (Apomorphy List). The genus also support the monophyly of Chironius:
Chironius is supported by six unambiguous synapomorscale rows keeled at midbody (6: 6-3),
phies and three ambiguous synapomorphies.
dentary teeth (18: 0--4), and distal calyces

on dorsal surface of hemipene (34: 3->5).
Chironius monophyly is supported in the
the Bayesian analysis (Fig. 5; tree length =
Bayesian analysis with a posterior probability
123.3, CI = 0.29, RI = 0.61). A phylogram
of 100% (node 16, Fig. 5).
estimating branch lengths for in-group taxa
The strong support for Chironius monoand the outgroup taxon Coluber constrictor
phyly is not surprising given that four of the
constrictor is shown in Fig. 6.
unambiguously synapomorphic characters
Monophyly of the Genus Chironius
identified by the parsimony analysis (those
The monophyly of Chironius (node 19,
related to dorsal scale row formulae) have
Fig. 3) has 93% bootstrap support from six
long been recognized as uniquely identifying
unambiguous synapomorphies: dorsal scale
characters of the genus. Although there is
rows at neck (character 21: 5-* 1), dorsal scale
considerable variation between characters 21rows at midbody (22: 6->l), dorsal scale rows
23 in the outgroups, it seems likely that these
anterior to vent (23: 7--0), spinules along the characters (and also character 35) are not
sulcus spermaticus (33: 2--5), even versus
completely independent of one another (i.e., it
odd modal number of scale rows as midbody
seems unlikely that a snake could possess, e.g.,
(35: 0-*9), and presence of a vertebral keel 10 dorsal rows at midbody and 17 dorsal rows
(0-*9). Three additional characters apoanterior to the vent). In order to assess the
morphic under DELTRAN optimization only
potential effects of nonindependent correla-
December
20063
0.60
0.800
1.00
0
1.200
HERPETOLOGICA
Chironius
400
1.400 6 0
septentnonalis
Chironius is not definitively
identified
multiventris
(Fig. 5).
Chironius
C0bhironius
-0 600Chironius multiventns cochranae
441
vincenti
foveatus
Intrageneric Relationships
of Chironius
multiventris
A clade containing laevicollis, scurrulus,

and both fuscus subspecies (node 9, Fig. 3)
has 55% bootstrap support from seven unambiguous synapomorphies: tail percent (2:
7->6), subcaudals (4: 6->5), anal plate (5:
9->0), apical pits (7: 8-*9), dentary teeth (18:
5->7), dorsal scale rows at neck (21: 1--0),
and dorsal scale rows at midbody (22: 1->0).
This clade is also supported in the Bayesian
analysis with a posterior probability of 87%
(node 7, Fig. 5). The two subspecies of C.
fuscus are paired in 92% of replicates (node 7,
Fig. 3) and supported by six unambiguous
synapomorphies: maxillary teeth (17: 6-*9),
dentary teeth (18: 7--8), eye-snout ratio (19:
4->6), male-female tail percent ratio (25:
0 o Chironius multiventns multiventns
2.800 1.-60-0 Chironius laurenti

2700 Chironius grandisquamis
1.200 Chironius fuscus fuscus
0.600
0 40 Sironius fuscus leucometapus

2.100
1.600 Chironius laevicollis

2.400
1.40 D
1.800 Chironius scurrulus
1.733 2.200 Chironius monticola

0.933 Chironius carinatus cannatus
0.200
1.600 Chironius carnatus flavopictus

2.200 Chironius cannatus spixi
7.933
2.000 Chironius exoletus

.900 Chironius bicannatus
5-*3), hemipene length (26: 5-.>3), and
retractor muscle origin (27: 4-*2). The fuscus

clade has a Bayesian posterior probability of
Chironius quadncarnatus maculoventns
81% (node 6, Fig. 5). Sister to fuscus is
a paired arrangement of laevicollis and
0 0?Chironius quadncannatus quadrncannatus
scurrulus
(node 6, Fig. 3; 62% bootstrap
3.000 Coluber constrictor constnctor
support) from three unambiguous synapomorFIG. 6.-Majority-rule consensus Bayesian phylogram
phies: SVL (1: 4--3), subcaudals (4: 5->2),
of Chironius taxa and outgroup taxon Coluber constrictor
and tail reduction (20: 1->4). This pairing has
constrictor. Numbers above branches indicate branch
lengths based on estimated rate of character change. a Bayesian posterior probability of 90% (node
5, Fig. 5). This is in agreement with Dixon et
tion between these characters, I ran a parsi- al. (1993) (Fig. 1). In a departure from both
mony analysis excluding characters 21, 23, and
the phylogeny proposed by Dixon et al. (1993)
35. Results (not shown) are nearly identical to and the parsimony analysis, the Bayesian
Fig. 3, including monophyly of Chironius, but
analysis offers 73% posterior probability
with lower bootstrap levels. This result
support for the placement of monticola as
suggests that these characters are not unduly
sister taxon to the clade containing laevicollis,
influencing the placement of taxa in the
scurrulus, and the two fuscus subspecies
analysis.
(node 8, Fig. 5).
All included members of the genera DenThe four multiventris subspecies are monodrophidion and Drymobius are among the
phyletic with the inclusion of vincenti in both
taxa most closely related to Chironius in the
analyses (node 13, Fig. 3; node 12, Fig. 5).
parsimony analsysis (Fig. 3), although bootThe five-taxon clade is supported with
strap support is below 50% for those relation- a Bayesian posterior probability of 74%, and
ships. These findings are in agreement with
bootstrap support is 73% from six unambiguDixon et al.'s (1993) hypothesis that Chironius ous and two ambiguous synapomorphies.
is the sister taxon of Dendrophidion and
Unambiguous synapomorphies are: eye-snout
Drymobius. Drymobius appears paraphyletic
ratio (19: 4-*6), tail reduction (20: 1-*0),
in this analysis, with Drymobius margaritihemipene length (26: 3->2), origin of referus as sister taxon to Chironius. The
tractor muscle (27: 4-4), extent of hemipene
Bayesian cladogram is sufficiently unresolved
naked basal area (30: 2-*1), and distal calyces
at the outgroup level such that a sister taxon
(34: to
4->3). Two multiventris subspecies,
3.300 Chironius flavolineatus
442
HERPETOLOGICA
[Vol.
62,
No.
m. cochranae and m. multiventris, are paired

duced a relatively long branch length leading
in 58% of bootstraps (node 10, Fig. 3), with
to the clade (Fig. 6; branch length = 1.600).
Furthermore, such a grouping is supported in
Bayesian posterior probability of 77% (node 9,
Fig. 5) from two unambiguous and three
the parsimony analysis by five unambiguous
synapomorphies: SVL (1: 4-*5), tail percent
ambiguous synapomorphies. Unambiguous
(2: 7-*6), apical pits (7: 8->9), postcephalics
synapomorphies are ventrals (3: 3-*5) and
(16: 3- >2), and basal spinules (31: 5-*0). The
eye-snout ratio (19: 6--7). A pairing of m.
septentrionalis and vincenti is supported withmore inclusive parsimony-based clade that
86% Bayesian posterior probability (node 10,
also contains bicarinatus and exoletus (node 5,
Fig. 3) is supported by two unambiguous
Fig. 5) and in 65% of bootstraps (node 11,
synapomorphies, tail reduction (20: 2->4)
Fig. 3) by four unambiguous synapomorphies:
and basal spinules (31: 9-.5), and one
SVL (1: 7-4), tail percent (2: 7-*8), subambiguous synapomorphy. However, it also
caudals (4: 6- >8), and male-female SVL ratio
(24: 6->3). Dixon et al. (1993) presented
has less than 50% bootstrap support. The
Bayesian supports the placement of exoletus
multiventris and vincenti as sister species
as sister taxon to the carinatus grouping (node
(Fig. 1).
4, Fig. 5; 61% posterior probability), a clade
In both the parsimony and Bayesian analyses, a more inclusive clade containing grand- that has a relatively long branch length
isquamis, laurenti, m. cochranae, m. foveatus, (Fig. 6), but placement of bicarinatus is
m. multiventris, m. septentrionalis, and vinunresolved on the Bayesian cladogram.
centi (node 15, Fig. 3; node 14, Fig. 4) is
The two quadricarinatus subspecies form
supported with a Bayesian posterior probabila clade (node 1, Fig. 3; node 1, Fig. 5) with
ity of 89% and in 67% of bootstraps by five
75% bootstrap support and 55% Bayesian
unambiguous and one ambiguous synapomorposterior probability. Five unambiguous and
phies (Fig. 4). Unambiguous synapomorphies
three ambiguous synapomorphies support the
are: SVL (1: 4-*7), ventrals (3: 1-*2), malequadricarinatus pairing in the parsimony
female tail percent ratio (25: 4--2), hemipene
analysis. The unambiguous synapomorphies
basal pocket size (29: 9--0), and extent of
are: supralabials entering the orbit (9: 3--2),
hemipene naked basal area (30: 3--2). The
maxillary teeth (17: 6-45), eye-snout ratio (19:
node placing laurenti as sister taxon to multi4-*3), male-female SVL ratio (24: 5-1), and
ventris and vincenti (node 14, Fig. 3; node extent
13,
of hemipene naked basal area (30:
Fig. 5) has 64% Bayesian posterior probability
3-1). Dixon et al. (1993) hypothesized that
and 62% bootstrap support. It arises from four quadricarinatus was the basal species in
unambiguous synapomorphies: ventrals (3:
Chironius (Fig. 1); the Bayesian analysis
2->3), apical pits (7: 8-4), hemipene length
supports that placement (node 16, Fig. 5).
(26: 5--3), and distal calyces (34: 5-4); this
The parsimony analysis places monticola
agrees with Dixon et al.'s (1993) hypothesized
basally within Chironius (node 19, Fig. 3),
placement of laurenti relative to multiventris with the quadricarinatus subspecies as sister
and vincenti (Fig. 1). However, Dixon et al.
taxa, but has less than 50% bootstrap support.
(1993) placed exoletus basal to multiventris,
As mentioned previously in this Results
vincenti, and laurenti; as stated above, both of section, monticola has a susbstantially differthese analyses supports grandisquamis as
ent placement on the Bayesian cladogram.
basal to muliventris, vincenti, and laurenti
Selected Species Considerations
(node 15, Fig. 3). The branch leading to this
within Chironius
clade (Fig. 6) is relatively long, with a length
of 2.800.
The placement of vincenti within the
Although bootstrap support for a monophy- multiventris clade (node 13, Fig. 3; node 12,
letic arrangement of the three carinatus
Fig. 5) is significantly supported with a 73%
subspecies (node 3, Fig. 3) is below 50%,
bootstrap value and 74% Bayesian posterior
the Bayesian posterior probability is signifiprobability. This has intriguing implications
cantly higher (node 3, Fig. 5; 86% posterior
because vincenti is an island species, occurprobability) and the Bayesian analysis proring only on Saint Vincent's Island in the West
December
2006]
HERPETOLOGICA
443
internally paired in the parsimony analysis

Indies (Dixon et al., 1993; Henderson et al,,
(node 2, Fig. 3), carinatus flavopictus and
1988), whereas the multiventris complex is
carinatus spixi, are both found in northeastern
a widespread mainland species distributed in
eight South American countries: Brazil (multi- Colombia but occupy different habitats: carventris cochranae, multiventris foveatus, and inatusflavopictus prefers evergreen rainforest
multiventris multiventris), Colombia (multi- while carinatus spixi inhabits dry forests and
savannas (Dixon et al., 1993). Rather than
ventris multiventris), Ecuador (multiventris
being paired with carinatus spixi in the
multiventris), Guyana (multiventris cochranae), Peru (multiventris multiventris), Suri- Bayesian analysis, carinatus carinatus is sister
nam (multiventris cochranae), Trinidad &
to carinatus flavopictus (node 2, Fig. 5).
Tobago (multiventris septentrionalis), and
There is some evidence in the form of
Venezuela (multiventris multiventris and mul-morphologically intermediate specimens that
tiventris septentrionalis). The finding in thissuggests that carinatus spixi may intergrade
analysis that multiventris septentrionalis is
with carinatus carinatus in Venezuela (Dixon
sister to vincenti is geographically consistent
et al., 1993). Although carinatus carinatus
with the distributions of the two species:
occupies evergreen rainforest and would seem
multiventris septentrionalis is the northernunlikely to occur sympatrically with the llanosmost of the multiventris subspecies, known
inhabiting carinatus spixi, the localities from
from six states in northern Venezuela as well
which the possible intergrade specimens were
as the island of Trinidad (Dixon et al., 1993).
collected (in Estados Amazonas and Sucre)
Interestingly, multiventris foveatus, the taxon are from areas of very diverse habitat in which
that is sister to vincenti and multiventris
sympatry is possible (Dixon et al., 1993). In
septentrionalis (node 12, Fig. 3; node 11,
light of the lack of strong support for the
Fig. 5), is known only from four states along
carinatus clade, the disagreement between
the eastern coast of Brazil (Dixon et al., 1993).
the two analyses regarding intra-clade organiEach taxon in the clade is supported by at
zation, and the existence of possible interleast five autapomorphies (Fig. 4): multiventris
grades between two members of the species
cochranae is supported by eight unambiguous
complex, the issues of speciation within the
autapomorphies and one ambiguous autapocarinatus clade present some interesting
morphy, multiventris foveatus is supported by
questions. In particular, carinatus spixi may
three unambiguous and two ambiguous autabe undergoing strong selection pressure to
pomorphies, multiventris multiventris is sup- adapt to its ecological status as savanna
ported by five unambiguous autapomorphies,
inhabitant; carinatus spixi is the only taxon
multiventris septentrionalis is supported by five in Chironius that does not undergo an
unambiguous autapomorphies, and vincenti is
ontogenetic change in eye size, perhaps
supported by ten unambiguous and three
because it is also the only Chironius species
ambiguous autapomorphies.
that does not spend the majority of its time in
According to Dixon et al. (1993), the threeclosed-canopy forests (Dixon et al., 1993). The
carinatus subspecies are distinct from all
parsimony analysis identified at least six
other Chironius species on the basis of a suite
apomorphies for each taxon in the carinatus
of combined characteristics (divided anal
clade (Fig. 4): carinatus carinatus is supplate, modal dorsal scale row formula 12ported by one unambiguous autapomorphy
12-8, and a green-olive dorsum). However,
and five ambiguous autapomorphies, carinathe parsimony analysis did not infer strong
tus flavopictus is supported by nine unambigsupport for the grouping of carinatus. The
uous autapomorphies, and carinatus spixi is
three subspecies do form a clade (node 3,
supported by five unambiguous autapomorFig. 3) but with only 41% bootstrap support.
phies and two ambiguous autapomorphies.
The carinatus clade is much more strongly
A pairing of fuscus fuscus and fuscus
supported in the Bayesian analysis with 86%
leucometapus (node 7, Fig. 3; node 6, Fig. 5)
posterior probability (node 3, Fig. 5) and
is well supported in both analyses but has
a branch length on the phylogram of 1.600
a relatively short branch length of 0.600 on
(Fig. 6). The two carinatus species that are
the Bayesian phylogram (Fig. 6). Wiest (1978)
444
HERPETOLOGICA
[Vol.
62,
No.
recognized fuscus leucometapus as a new

Apomorphy support for each taxon as identitaxon on the basis of several morphological
fied by the parsimony analysis (Fig. 4) supfeatures, namely the presence of a white snoutports the recognition of each taxon as a unique
lineage: quadricarinatus maculoventris is supand forehead and the lack of enlarged
proximal hemipene spines (present in fuscus
ported by five unambiguous and three ambigfuscus). Other distinctive features that distin- uous autapomorphies, and quadricarinatus
guish fuscus leucometapus from fuscus fuscusquadricarinatus is supported by five unaminclude higher mean number of ventrals and
biguous and two ambiguous autapomnorphies.
lower mean number of subcaudals (Dixon et
The only taxon to show significant differal., 1993). Although the average elevation for
ence in placement between the two analyses is
fuscus fuscus is lower than that of fuscus
monticola: the parsimony analysis placed it
leucometapus (Dixon et al., 1993), the two
basally to all other Chironius (Fig. 3) whereas
fuscus subspecies apparently occur in sympatthe Bayesian analysis placed it basal to an
ry in the three central Peruvian departments
internal clade containing laevicollis, scurrulus,
from which fuscus leucometapus is documenand the two fuscus subspecies (Fig. 5).
ted (Huanuco, Junin, and San Martin),
Furthermore, neither cladogram agreed with
lending credence to the recognition of sepaDixon et al.'s (1993) internal placement of
rate species. Evidence from the parsimony
monticola as sister taxon to carinatus (Fig. 1).
analysis shows that both taxa are supported by
At this time it is unclear why the placement of
autapomorphies (Fig. 4): fuscus fuscus is
monticola varies between analyses, but it is
supported by five unambiguous autapomorworth noting that monticola was supported as
phies andfuscus leucometapus is supported by a distinct lineage by thirteen autapomorphies
six unambiguous autapomorphies.
(Fig. 4), five of which were unambiguous and
Sister to fuscus is the pairing of laevicollis
eight ambiguous, more than any taxon except
and scurrulus; a unique ontogenetic color
exoletus (fourteen unambiguous autapomorchange shared by laevicollis and scurrulus
phies). Selectional pressure on monticola may
supports their status as sister to one another
differ from that of other Chironius, as
(Marques and Sazima, 2003), which is consismonticola is the only Chironius to routinely
tent with both these analyses (node 6, Fig. 3;
occur at high elevations (Dixon et al., 1993).
node 5, Fig. 5) and that of Dixon et al. (1993)
Proposed Taxonomic Revisions to
(Fig. 1). The pairing of laevicollis and scurruChironius Taxa
lus has a relatively long branch length of 2.400
on the Bayesian phylogram (Fig. 6). These
I propose taxonomic revision of taxa within
two species are geographically disjunct from
the carinatus, fuscus, multiventris, and quadone another (laevicollis is reliably known only ricarinatus species groups. Both the parsimofrom the southeastern coast of Brazil; scurru- ny and Bayesian analyses indicate distinct
lus is more widely distributed across equatolineages for taxa even within recognized
rial South America).
species groups. This evidence, in combination
The pairing of the two quadricarinatus
with diagnostically unique morphological
subspecies is well supported in both analyses
characters for each taxon (see Selected
(node 1, Fig. 3; node 1, Fig. 5). Wiest (1978)
Species Considerations within Chironius,
recognized quadricarinatus maculoventris as
above, and the list of apomorphies in Appena taxon separate from the nominate form on
dix IV) as well as other considerations (e.g.,
the basis of two morphological features:
the well-supported placement of vincenti
quadricarinatus maculoventris has ventral
within the multiventris group), suggests that
speckling that is absent in quadricarinatus
elevation to full species status is warranted for
quadricarinatus as well as a distinct postocular
the eleven Chironius taxa historically recog-
stripe, also absent in quadricarinatus quadnized as subspecies.
ricarinatus. There is also variation in features

such as mean number of ventrals (greater in
quadricarinatus maculoventris) and subcaudals (fewer in quadricarinatus maculoventris).
(1) Chironius carinatus carinatus. This was the

nominate form of this species group so I am
resurrecting Chironius carinatus for this taxon
(Fitzinger, 1826; Linnaeus, 1758).
December
2006]
HERPETOLOGICA
445
(2) Chironius carinatus flavopictus. I am resurrecting

the name Chironius flavopictus for this taxon
et al. (1993) contended that most appearances

of Chironius cinnamomeus in the literature
(Peters, 1960; Werner, 1909).
referred to specimens of Chironius scurrulus,
(3) Chironius carinatus spixi. I am resurrecting the
adults of which are cinnamon-colored. In the
name Chironius spixi for this taxon (DonosoBarros, 1969; Hallowell, 1845).
studies using more than one Chironius
(4) Chironius fuscus fuscus. This was the nominate
species, clades were not sufficiently resolved
form of this species group so I am resurrecting
to
suggest or reject a monophyletic grouping
Chironius fuscus for this taxon (Amaral, 1929;
of Chironius species, including C. cinnamoLinnaeus, 1758).
(5) Chironius fuscus leucometapus. Based on the
meus (Buckley et al., 2000; Dowling et al.,
original description of C. f. leucometapus
by Highton et al., 2002).
1996;
Dixon et al. (1993), I suggest Chironius leucomeThe present analysis demonstrates the utility
tapus for this taxon.
of quantitatively analyzing the existing mor(6) Chironius multiventris cochranae. I am resurrecting the name Chironius cochranae for this taxon
phological data for Chironius in combination
(Hoge and Romano, 1969).
with data for outgroup taxa; my findings
(7) Chironius multiventris foveatus. I am resurrecting
support the monophyly of Chironius but
the name Chironius foveatus for this taxon
indicate that revision of the genus is warranted
(Bailey, 1955).
(8) Chironius multiventris multiventris. This was the
at the species level. There is some support for
nominate form of this species group so I am
monophyly of species groups as recognized by
resurrecting Chironius multiventris for this taxon
Dixon et al. (1993), but elevation to full species
(Schmidt and Walker, 1943).
status
is proposed for all historically recognized
(9) Chironius multiventris septentrionalis. Based on
subspecies on the basis of multiple autapomorthe original description of C. m. septentionalis by
Dixon et al. (1993), I suggest Chironius septenphies per taxon. Some intrageneric relationtrionalis for this taxon.
ships are still not strongly supported by these
(10) Chironius quadricarinatus maculoventris. Based
analyses,
and at least one taxon (C. monticola)
on the original description of C. q. maculoventris
could not be unequivocally placed within
by Dixon et al. (1993), I suggest Chironius
maculoventris for this taxon.
Chironius. A more comprehensive treatment
( 11) Chironius quadricarinatus quadricarinatus. This

of the genus including molecular data is
was the nominate form of this species group, so I
necessary
to produce a robust phylogeny
am resurrecting Chironius quadricarinatus for this
further resolve questions about Chironius
taxon (Boie, 1827; Bailey, 1955).
Future Directions
and
species diversity and relationships.
Acknowledgments.-I thank S. Poe for critically review-
At most three taxa of Chironius identified

ing early versions of this manuscript and for his valuable
advice throughout the duration of this project. Thanks to
to species have been included in molecularA. Harrison, E. Hulebak, H. Maclnnes, and E. W. Schaad
based approaches to the study of snake
for reading an early version of this manuscript. Helpful
evolution (Buckley et al., 2000; Dowling etsuggestions by F. T. Burbrink and two anonymous
al., 1996; Heise et al., 1995; Highton et al., reviewers greatly improved a later version of this
2002; Lopez and Maxson, 1995), so at this

manuscript. I thank J. R. Dixon and K. Vaughan (Texas
A&M University) for permission to examine specimens in
time there are not enough existing molecular
the Texas Collective Wildlife Collection. Last, I wish to

data to be useful in Chironius phylogenetic
acknowledge the extensive work of J. A. Wiest, Jr. and J.
analyses. Three of the studies including
R. Dixon in collecting the Chironius data from which this
Chironius taxa (Buckley et al., 2000; Dowling
study was inspired.
et al., 1996; Highton et al., 2002) were based
at least in part on the same allozyme data set
LITERATURE CITED
and included a specimen of Chironius cinna-
momeus, a taxon with a history of problematic

diagnoses. Based on the original description of
the type specimen of Natrix cinnamomea

Wagler 1824, Hoogmoed and Griiber (1983)
believed that it should subsequently have
been placed in the genus Pseustes, making
Chironius cinnanomeus a nomen dubium; da
Cunha and do Nascimento (1983) and Dixon
AMARAL, A. 1929. Estudos sobre ophidios ne6tropicos.

XVII. Valor sistematico de varias formas de ophidios
neotropicos. Mem6rias do Instituto Butantan (Sao
Paulo) 4:3-68.
BAILEY, J. R. 1955. The snakes of the genus Chironius in

southeastern South America. Occasional Papers of the
Museums of Zoology, University of Michigan 571:1-21.
BUCKLEY, L., M. KEARNEY, AND K. DE QUEIROZ. 2000.
Slowly-evolving protein loci and higher-level snake

phylogeny: a reanalysis. Herpetologica 56:324-332.
446
HERPETOLOGICA
[Vol.
62,
No.
BowE, F. 1827. Bemerkungen uber Merrem's Versuch

eines Systems der Amphibien, Marburg, 1820. Erste
Lieferung: Ophidien. Isis von Oken 20:508-566.
HOGE, A. R., AND S. A. ROMANO. 1969. A new species of

Chironius (Serpentes-Colubridae). Mem6rias do Instituto Butantan (Sao Paulo) 34:93-95.
CALDAROLO, J., A. DILMAGHANI, J. GAGNON, K. HAYCOCK, J.
HUELSENBECK, J. P., AND F. RONQUIST. 2001. MRBAYES:
ROTH, C. SOPER, AND E. WASSERMAN. 1992. StatView.
Bayesian inference of phylogeny. Bioinformatics 17:
Version 5.0. SAS Institute Inc., Cary, New York, U.S.A.
754-755.
DA CUNHA, 0. R., AND F. P. Do NASCIMENTO. 1983. Ofidios
KLUGE, A. G., AND J. S. FARRIS. 1969. Quantitative
phyletics and the evolution of the anurans. Systematic

Zoology 18:1-32.
LANCINI, V. A. R. 1979. Serpientes de Venezuela. Talleres
46:139-172.
de Graficas Armitano, C.A., Caracas, Venezuela.
DIXON, J. R., J. A. WIEST, JR., AND J. M. CEI. LEWIS,
1993. P. 0. 2001. A likelihood approach to estimating
Revision of the Neotropical Snake Genus Chironius
phylogeny from discrete morphological character data.
Fitzinger (Serpentes, Colubridae). Monografie XIII.
Systematic Biology 50:913-925.
da Amaz6nia 15: as especies de Chironius da Amazonia

oriental (Para, Amapa e Maranhao). (Ophidia: Colubridae). Mem6rias do Instituto Butantan (Sao Paulo)
Museo Regionale di Scienze Naturali, Torino, Italy.

DONoso-BARROS, R. 1969. Contribuci6n al conocimiento
del genero Chironius (Serpentes: Colubridae). Boletin
de la Sociedad de Biologia de Concepci6n 41:189-197.
LINNAEUS, C. 1758. Systema naturae per regna tria
naturae, secundum classes, ordines, genera, species,

cum characteribus, differentiis, synonymis, locis. Tomus 1. Editio Decima, reformata. Holmiae: Laurentii
DOWNLING, H. G. 1951. A proposed standard system of Salvii.

counting ventrals in snakes. British Journal of Herpe- LOPEZ, T. J., AND L. R. MAXSON. 1995. Mitochondrial DNA
sequence variation and genetic differentiation among
DONVLING, H. G., C. A. HASS, S. B. HEDGES, AND R.
colubrine snakes (Reptilia: Colubridae: Colubrinae).
Biochemical Systematics and Ecology 23:487-505.
HIGIITON. 1996. Snake relationships revealed by slowevolving proteins: a preliminary survey. Journal of
MARQUES, 0. A. V., AND I. SAZIMA. 2003. Ontogenetic color
Zoology, London 240:1-28.
changes may strengthen suggestion about systematic
DOWLING, H. G., AND J. M. SAVAGE. 1960. A guide to the
affinities between two species of Chironius (Serpentes,
snake hemipenis: a survey of basic structure and
Colubridae). Phyllomedusa 2:65-67.
systematic characteristics. Zoologica 45:17-28.
MARX, H., AND G. B. RABB. 1972. Phyletic analysis of fifty
FARRIS, J. S. 1989. The retention index and the rescaled
characters of advanced snakes. Fieldiana (Zoology)
63:1-321.
consistency index. Cladistics 5:417-419.
MAU, B., AND M. NEWTON. 1997. Phylogenetic inference
FELSENSTEIN, J. 1985. Confidence limits on phylogenies:
for binary data on dendrograms using Markov chain
an approach using the bootstrap. Evolution 39:783-791.
Monte Carlo. Journal of Computational and Graphical
FITZINGER, L. I. 1826. Neue Classification der Reptilien
Statistics 6:122-131.
nach ihren naturlichen Verwandschaften nebst einer
Verwandschafts - tafel und einem Verzeichnisse der
MAU, B., M. NEWTON, AND B. LARGET. 1999. Bayesian
phylogenetic inference via Markov chain Monte Carlo
Reptilien Sammlung des K. K. Zoologischen Museums
methods. Biometrics 55:1-12.
zu Wien. J. G. Heubner, Wien, Germany.
GEYER, C. J. 1991. Markov chain Monte Carlo maximum
MYERS, C. W. 1974. The systematics of Rhadinaea
likelihood. Pp. 156-163. In E. M. Keramidas (Ed.),
(Colubridae), a genus of New World snakes. Bulletin
of the American Museum of Natural Historv 153:1-262.
Computing Science and Statistics: Proceedings of the
23rd Symposium on the Interface. Interface FoundaNAGY, Z. T., R. LAWSON, U. JOGER, AND M. WINK. 2004.
tion, Fairfax Station, Virginia, USA.
Molecular systematics of racers, whipsnakes, and
GUNTHER, A. C. L. G. 1864. The Reptiles of British India.
relatives (Reptilia: Colubridae) using mitochondrial
Robert Hardwick, London, England.
and nuclear markers. Journal of Zoological Systematics
HALLONW7ELL, E. 1845. Descriptions of reptiles from South and Evolutionary Research 42:223-233.
America supposed to be new. Proceedings of the
ORTENBURGER, A. I. 1928. The whip snakes and racers:
Academy of Natural Sciences of Philadelphia 2:
genera Masticophis and Coluber. Memoirs of the
241-250.
University of Michigan Museums 1:1-247.
HEISE, P. J., L. R. MAXSON, H. G. DOWLING, AND S. B.
PETERS, J. A. 1960. The snakes of Ecuador. A checklist and
HEDGES. 1995. Higher-level snake phylogeny inferred
key. Bulletin of the Museum of Comparative Zoology
from mitochondrial DNA sequences of 12S rRNA and
122:491-541.
16S rRNA genes. Molecular Biology and Evolution
POPE, C. H. 1934. List of Chinese turtles, crocodilians,
12:259-265.
and snakes, with keys. American Museum Novitates
tology 1:97-99.
HENDERSON, R. W., R. A. SAJDAK, AND R. M. HENDERSON.
1988. The rediscovery of the West Indian colubrid

snake Chironius vincenti. Amphibia-Reptilia 9:
733:1-29.
SCHNIIDT, K. P., AND W. F. WALKER. 1943. Peruvian snakes
from the University of Arequipa. Zoological Series of

Field Museum of Natural History 24:279-296.
HIGHTON, R., S. B. HEDGES, C. A. HASS, AND H. G.
STUART, L. C. 1932. Studies on Neotropical Colubrinae. I.
DOWLING. 2002. Snake relationships revealed by slowlyThe taxonomic status of the genus Drymobius Fitzinevolving proteins: further analysis and a reply. Herpeger. Occasional Papers of the Museums of Zoology,
tologica 58:270-275.
University of Michigan 236:1-16.
415-416.
HILLIS, D. M., AND J. J. BULL. 1993. An empirical test of
bootstrapping as a method for assessing confidence in

phylogenetic analysis. Systematic Biology 42:182-192.
SWN'OFFORD, D. L. 2002. PAUP*. Phylogenetic Analysis

Using Parsimony (*and Other Methods). Version 4.0.
Sinauer Associates, Sunderland, Massachusetts, U.S.A.
December
2006]
HERPETOLOGICA
447
THIELE, K. 1993. The holy grail of the perfect character:
Maqueyes, Rio Pilon: TCWC 52507; 31 mi N Gonzales:
the cladistic treatment of morphometric data. Cladistics

9:275-268.
WERNER, F. 1909. Uber neue oder seltens Reptilien des
Naturhistorischen Museums in Hamburg. Mitteilungen
aus dem Zoologischen Museum in Hamburg 26:
205-247.
WIENS, J. J. 1995. Polymorphic characters in phylogenetic
systematics. Systematic Biology 44:482-500.
WIEST, J. A. 1978. Revision of the neotropical snake genus
TCWC 57346.
Chironius Fitzinger (Serpentes, Colubridae). Ph.D.

Dissertation, Texas A&M University, College Station,
Texas, USA.
Accepted: 3 June 2006

Associate Editor: Frank Burbrink
APPENDIX I
Specimenis Examined
Drymobius rhombifer (n = 2).-Peru: Loreto: Centro

Uni6n: TCWC 44595; Moropon: TCWC 44594.
Drymoluber dichrous (n = 2).-Perl: Loreto: Moropon: TCWC 42190. TCWC 42193.
Masticophisflagellum testaceus (n 2).-USA: Netw

Mexico: Guadalupe County, 1 mi S Santa Rosa: UNM
36456; Guadalupe County, 21 mi W Santa Rosa: UNM
31638.
Masticophis taeniatus taeniatus (n = 2).-USA: New

Mexico: Grant County, 10 mi S Silver City: UNM 15205;
Grant County, 13 mi S Silver City: UNM 15294.
Mastigodryas bifossatus (n = 2).-Venezuela: Gudrico: 6 km W San Fernando de Apure: TCWC 47929; 8 km
N El Manteco: TCWC 60154.
Mastigodryas boddaerti (n = 2).-Venezuela: Aragua: 2 km SE Rancho Grande Biological Station: TCWC
59015; Miranda: 34.4 km N Altagracia: TCWC 59011.
Mastigodryas heathi (n = 1).-Ecuador: El Oro: 9 mi
S Zaruma: TCWC 24132.
Mastigodryas melanolomus (n = 2).-Mexico: San

Luis Potosi: 5 mi NE Xilitla: TCWC 29490; Veracruz:
Cooperative Wildlife Collection (Texas A&M University);
Volcan
UNM = Museum of Southwestern Biology (University
of San Martin, San Andreas Tuxtla: TCWC 26430.
Mastigodryas pleei (n = 1).-Venezuela: Gudrico:
New Mexico).
Coluber constrictor constrictor (n = 2).-USA: New 8.3 km N Altagracia: TCWC 59208.
Pseustes poecilonotus (n = 2). Peri: Loreto: Cento
Jersey: Burlington County: UNM 30111, UNM 30115.
Uni6n: TCWC 44708; Venezuela: Miranda: 1 mi N Agua
Dendrophidion dendrophis (n = 2).-Peru: Loreto:
Blanca: TCWC 59079.
Mishana, on Rio Nanay: TCWC 41417, TCWC 42196.
Pseustes sulphureus (n = 1).-Perui: Loreto: Centro
Dendrophidion nuchalis (n = 3).-Venezuela: GudrUni6n: TCWC 44709.
ico: 1 km S Agua Blanca: TCWC 59019; 26 km N
Altagracia: TCWC 59024, TCWC 59026.
Salvadora hexalepis (n = 3).-USA: Arizona: Coconino County, 10 mi W Cameron: UNM 34341; Mohave
Dendrophidion pericarinatum (n = 2).-Costa Rica:
Museum acronyms are as follows: TCWC = Texas
San Jose: Parque Nacional Carara: TCWC 84083; Reserva

Biological Carara: TCWC 83370.
County, 10 mi S Wickiup: UNM 36595; Pima County,
TCWC 42197.
61825.
Drymobius margaritiferus (n = 4). Mexico: Chiapas: 1.9 mi N Puerto Arista: TCWC 57101; 800 m N
Iquitos: TCWC 48671; Venezuela: Miranda: 8 km E Santa
Cabeza Prieta Gamle Refuge: UNM 31259.

Drymarchon corais (n = 3). Mexico: Tamaulipas:
Salvadora grahamiae (n = 4).-USA: New Mexico:
0.3 mi SW Rancho Carricitos: TCWC 49906; vicinity of
Eddy County: UNM 61830, UNM 66947; Eddy County,
Rancho Carricitos: TCWC 58108; Peru: Loreto: Iquitos: Dark Canyon: UNM 48569; Last Chance Canyon: UNM
Spilotes pullatus (n =2). Peru: Loreto: 5 mi NNE
Puerto Arista: TCWC 57421; Tamauilipas: 0.7 mi NE Teresa: TCWC 58990.
448
HERPETOLOGICA
[Vol.
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No.
APPENDIX II
Data matrix of morphological character states used for parsimony analysis. Character numbers correspond to those
listed in the Character Descriptions (Materials and Methods). All characters are ordered except for 14, which was
transformed by a stepmatrix. Weighting was set so that steps had equal weight among characters regardless of number of
states in each character.
Character
Chironius
number
bicarinatus
10
11
12
Chironius carinatus carinatus 5 6 1 3 9 1 9 6 2 5 4

Chironius carinatus flavolineatus 6 6 1 4 9 1 9 8 4 5 4
Chironius carinatus spixi 5 6 1 4 8 1 9 5 4 4 4
Chironius exoletus 3 7 0 0 8 1 5 8 3 5 5
0
0
0
0
Chironius flavolineatus 0 8 1 8 9 0 8 8 3 4 4 0
Chironius fuscus fuscus 4 6 0 5 0 1 9 8 3 4 4

Chironius fuscus leucometapus 3 6 1 5 0 1 9 7 3 3 4
Chironius grandisquamis 9 7 2 3 9 3 8 8 3 4 4
Chironius laevicollis 8 6 1 1 0 0 9 7 3 4 3
Chironius laurenti 7 6 3 6 9 3 1 8 3 5 4
Chironius monticola 3 6 0 5 0 3 9 8 4 4 4
Chironius multiventris cochranae 7 8 6 8 9 1 0 8 3 4 4
Chironius multiventris foveatus 7 7 3 6 9 1 1 8 3 5 4
Chironius multiventris multiventris 8 7 5 7 9 1 3 8 3 4 4
Chironius multiventris septentrionalis 4 8 3 8 9 1 1 8 3 4 3
0
0
0
0
0
0
0
0
0
0
Chironius quadricarinatus maculoventris 1 6 1 6 9 3 9 5 2 5 5 0

Chironius quadricarinatus quadricarinatus 1 7 0 7 9 4 9 6 2 3 4 0
Chironius scurrulus 7 6 1 2 0 0 9 8 3
Chironius vincenti 1 8 3 9 9 4 3 8 3
6
5
4
6
0
0
Coluber constrictor constrictor 2 3 4 3 9 0 0 3 0 1 0 7

Dendrophidion dendrophis 0 ? 1 ? 0 7 9 8 3 0 4 0
Dendrophidion nuchalis 0 6 1 ? 6 7 6 8 3 3 2
Dendrophidion pericarinatum 0 9 1 9 9 6 0 9 3 3 4
Drymarchon corais 6 0 7 0 0 0 9 5 2 0 0
Drymobius margaritiferus 1 6 0 6 5 ? 9 8 3 5 4
Drymobius rhombifer 1 3 1 5 9 8 0 8 3 5 4
Drymoluberdichrous 1 4 3 4 0 0 0 5 2 0 4
0
0
0
0
0
0
Masticophis flagellum testaceus 2 3 8 7 9 0 0 8 4 1 0 7

Masticophis taeniatus taeniatus 2 5 8 7 9 0 5 5 2 1 0 2
Mastigodryas bifossatus 3 4 5 4 9 0 0 5 2 3 2 0
Mastigodryas boddaerti 2 ? 5 ? 9 0 0 6 2 3 6 0
Mastigodryas heathi 0 4 6 6 0 0 0 8 3 6 4 0
Mastigodryas melanolomus 1 4 4 6 9
Mastigodryas pleei 0 3 2 4 9 0
Pseustes poecilonotus 2 4 6 6 5 2
Pseustes sulphureus 4 3 9 8 0 9
Salvadora grahamiae 0 3 5 8 7 0
0 0 9 3 3 4
9 5 ? 3 8
0 5 3 9 9
0 5 2 6 8
0 5 2 4 0
Salvadora hexalepis 0 2 7 6 9 0 6 8 5 4 4
Spilotes pullatus 5 3 9 5 0 6 0 0 0 0 0
0
0
0
0
9
6
0
Character number 13 14 15 16 17 18 19 20 21 22 23 24
Chironius bicarinatus 1 0 0 3 6 6 4 4 1 1 2
Chironius carinatus carinatus 1 1 0 2 6 5 4 5 1 1 0
Chironius carinatus flavolineatus 1 1 0 2 6 ? 5 2 1 1 0
Chironius carinatus spixi 1 1 0 2 6 6 4 4 1 1 0
Chironius exoletus 1 1 0 3 4 3 5 6 1 1 0
7
6
2
4
2

5
5
9
7
Chironius
laurenti
December
20061
HERPETOLOGICA
APPENDIX
449
II
Continued.
Character
Chironius
number
13
monticola
14
15
16
17
18
19
20
21
22
23
Chironius multiventris cochranae 6 1 0 3 7 6 8 0 1 1

Chironius multiventrisfoveatus 1 1 0 4 6 ? 6 0 1 1
Chironius multiventris multiventris 1 1 0 2 7 6 7 0 1 1
Chironius multiventris septentrionalis 2 1 0 4 7 ? 7 0 1 1
24
0
0
0
0
8
8
7
2
Chironius quadricarinatus maculoventris 1 1 0 3 5 4 3 6 ? 1 0 1

Chironius scurrulus 1 0 0 1 7 7 4 4 0 0 0 4
Chironius vincenti 5 1 0 4 8 8 4 ? 1 1 0 3
Coluber constrictor constrictor 1 ? 0 5 0 0 2 ? 5 6 7 ?
Dendrophidion dendrophis 1 ? 0 4 9 ? 9 ? 5 6 7 ?
Dendrophidion nuchalis 1 ? 2 5 3 ? 9 ? 5 6 7 ?
Dendrophidion pericarinatum 1 ? 0 6 ? ? 8 ? 3 6 7 ?
Drymarchon corais 1 ? 0 2 0 ? 0 ? 6 6 6 ?
Drymobius margaritiferus 1 ? 0 4 ? ? 4 ? 5 6 7 ?
Drymobius rhombifer 1 ? 0 3 5 ? 6
Drymoluberdichrous 1 ? 0 5 2 ? 3
Masticophis flagellum testaceus 1 ? 3 7 1 1
Masticophis taeniatus taeniatus 1 ? 0 4 0 0
Mastigodryas bifossatus 1 ? 0 6 ? ? 1
? 5 6 7
? ? 4 7
6 ? ? 6 ?
4 ? 3 4 3
? 5 4 7
?
?
?
?
?
Mastigodryas boddaerti 1 ? 0 4 ? ? 4 ? 6 8 9 ?
Mastigodryas heathi 1 ? 0 6 ? ? 2 ? 6 6 7 ?
Mastigodryas melanolomus 1 ? 0 5 ? ? 6 ? ? 6 7 ?
Mastigodryas pleei 1 ? 0 6 ? ? 4 ? 5
Pseustes poecilonotus 1 ? 0 5 ? ? 5 ? 8
Pseustes sulphureus 9 ? 0 1 ? ? 2 ? 9
Salvadora grahamiae 1 ? 3 7 ? ? 4 ? 5
Salvadora hexalepis 1 ? 9 9 ? ? 3 ? 5
Spilotes pullatus 1 ? 0 2 1 ? 3 ? 3
6
?
9
6
6
?
7 ?
? ?
5 ?
5 ?
5 ?
? ?
Chironius bicarinatus 6 5 7 0 9 3 1 3 5 5 9 9
Chironius carinatus carinatus 8 5 6 4 9 3 0 4 6 5 9 9
Chironius carinatus flavolineatus 6 9 9 4 9 3 0 5 7 9 9 9

Chironius carinatus spixi 8 6 6 0 9 5 0 4 7 6 9 9
Chironius exoletus 7 5 5 4 9 4 1 4 8 5 9 9
Chironius flavolineatus 5 5 4 0 9 4 2 5 8 4 9 ?
Chironius fuscus fuscus 0 3 2 0 9 3 2 4 6 5

Chironiusfuscus leucometapus 3 3 2 0 9 3 2 3 6 4
Chironius grandisquamis 2 5 4 0 0 2 2 7 9 5
Chironius laevicollis 4 4 4 4 9 3 2 4 7 5
9
9
9
9
9
9
9
?
Chironius laurenti 2 3 4 0 0 2 2 4 6 4 9 ?
Chironius monticola 9 6 8 0 9 9 2 2 5 7 9 9
Chironius multiventris cochranae 1 1 1 0 0 1 2 4 6 3 9 9
Chironius multiventrisfoveatus 4 1 0 4 0 1 2 4 7 3 9 9
Chironius multiventris multiventris 2 2 1 0 0 1 2 3 6 2 9 9
Chironius multiventris septentrionalis 3 2 0 4 0 1 2 3 5 3 9 9
Chironius quadricarinatus maculoventris 8 6 7 9 9 1 2 2 6 7 9 ?
Chironius quadricarinatus quadricarinatus 8 6 5 0 9 1 2 2 4 7 9 ?
Chironius vincenti 3 1 0 9 0 0 2 4 6 3 9 9
Coluber constrictor constrictor 9 2 ? ? ? 4 0 2 3 2 0 0
Dendrophidion dendrophis ? ? ? ? ? ? ? ? ? ? 0
Dendrophidion nuchalis ? 2 ? ? ? 2 0 2 2 0 0
Dendrophidion pericarinatum ? 2 ? ? ? 4 2 2 2 4 0
Drymarchon corais ? 5 ? ? ? 3 0 ? ? 6 0
0
0
0
0
Drymobius margaritiferus ? 5 ? ? ? 4 2 2 2 ? 0 0
Drymobius rhombifer ? ? ? ? ? ? ? ? ? ? 0 0
Drymoluber dichrous ? 0 ? ? ? 0 0 3 3 3 0 0
450
HERPETOLOGICA
[Vol.
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APPENDIX
No.
II
Continued.
Masticophis flagelluim testaceus 6 5 ? ? ? 6 0 2 1 1 0 0

Masticophis taeniatus taeniatus 8 3 ? ? ? 4 2 0 1 3 0
Mastigodryas bifossatus ? 6 ? ? ? 8 0 0 3 7 0
Mastigodryas boddaerti ? 9 ? ? ? 6 0 2 4 7 0
Mastigodryas heathi ? ? ? ? ? ? ? ? ? ? 0
Mastigodryas melanolomus ? ? ? ? ? ? ? ? ? ? 0
Mastigodnjas
Pseustes
pleei
poecilonotus
3
?
?
8
?
?
?
?
8
?
3
0
0
0
0
0
0
Pseustes sulphuireuis ? ? ? ? ? ? ? ? ? ? 0 0
Salvadora grahaniae ? 7 ? ? ? 2 0 0 0 9 0 0
Salvadora hexalepis ? 7 ? ? ? 2 0 0 0 3 0 0
Spilotes puillatus ? 3 ? ? ? 4 0 9 9 3 0 0
APPENDIX III
Data matrix of morphological character states used for Bayesian analysis. Character numbers correspond to those
listed in the Character Descriptions (Materials and Methods). All characters are ordered except for 14, which was
unordered. Weighting was set so that steps had equal weight among characters regardless of number of states in each
character.
Character
number
10
11
12
Chironius bicarinatus 2 4 1 3 5 1 4 3 2 2 2 0
Chironius carinatutsflavolineat.us 3 3 1 2 5 1 5 4 2 3 2 0


Chironius grandisquam,is 5 4 1 2 5 2 4 4 2 2 2
Chironius multiventris foveatus 4 4 2 3 5 1 1 4 2 3 2
0
0
0
0
0
0
0
0
Chironius muiltiventris septentrionalis 2 4 2 4 5 1 1 4 2 2 2 0
Chironius quadricarinatus maculoventris 1 3 1 3 5 2 5 3 1 3 3 0
Chironius quadricarinattis quiadricarinatus 1 4 0 4 5 2 5 3 1 2 2 0

Chironius scurrulus 4 3 1 1 0 0 5 4 2 3 2
Chironius vincenti 1 4 2 5 5 2 2 4 2 3 3
Coluber constrictor constrictor 1 2 2 2 5 0 0 2 0 1 0
Dendrophidion dendrophis 0 ? 1 ? 0 4 5 4 2 0 2
Dendrophidion nuchalis 0 3 1 ? 3 4 3 4 2 2 1
Dendrophidion pericarinatum 0 5 1 5 5 3 0 5 2 2 2
Drymarchon corais 3 0 4 0 0 0 5 3 1 0 0
0
0
4
0
0
0
0
Drynobius mnargaritiferuls 1 3 0 3 3 ? 5 4 2 3 2 0
Drymobius rhombifer 1 2 1 3 5 4 0 4 2 3 2 0
Drymoluiber dichrous 1 2 2 2 0 0 0 3 1 0 2 0
Masticophisflagellum testaceus 1 2 4 4 5 0 0 4 2 1 0 4
Masticophis taeniatus taeniatus 1 3 4 4 5 0 3 3 1 1 0 1

Mastigodryas bifossatus 2 2 3 2 5 0 0 3 1 2 1
Mastigodryas boddaerti 1 ? 3 ? 5 0 0 3 1 2 3
Mastigodryas heathi 0 2 3 3 0 0 0 4 2 3 2
Mastigodryas melanolornus 1 2 2 3 5 0 0 5 2 2 2
Mastigodryas pleei 0 2 1 2 5 0 5 3 ? 2 4
Pseustes poecilonotus 1 2 3 3 3 1 0 3 2 5 5
0
0
0
0
0
0
Salvadora
Pseustes sulphuretus 2 2 5 4 0 5 0 3 1 3 4 0
grahamiae
December
20061
HERPETOLOGICA
451
APPENDIX III
Continued.
Character
number
10
11
12
Salvadora hexalepis 0 1 4 3 5 0 3 4 3 2 2 3
Spilotes pullatus 3 2 5 3 0 3 0 0 0 0 0 0
Chironius
bicarinatus
Chironius carinatus flavolineatus 1 1 0 1 3 ? 3 1 1 1 0 1

Chironinis flavolineatus 1 0 0 1 3 3 3 0 1 1 0 0
Chironius fuscus fuscus 1 0 0 0 5 5 4 1 0 0 0 3

Chironiusfuscus leucometapus 0 0 0 1 5 4 3 1 0 0 0 3
Chironius grandisquamis 1 3 0 1 4 4 2 1 0 0 0 5
Chironinis laevicollis 0 0 0 0 3 4 2 4 0 0 0 4
Chironius laurenti 1 1 0 2 3 3 1 5 1 1 0 3
Chironius multiventris foveatus 1 1 0 2 3 ? 3 0 1 1 0
Chironius multiventris multiventris 1 1 0 1 4 3 4 0 1 1 0
3
4
4
4
Chironius multiventris septentrionalis 1 1 0 2 4 ? 4 0 1 1 0 1

Chironius quadricarinatus maculoventris 1 1 0 2 3 2 2 3 ? 1 0 1
Chironinis vincenti 3 1 0 2 4 4 2 ? 1 1 0 2
Colmber constrictor constrictor 1 ? 0 3 0 0 1 ? 3 3 4 ?
Dendrophidion dendrophis 1 ? 0 2 5 ? 5 ? 3 3 4 ?
Dendrophidion nuchalis 1 ? 1 3 2 ? 5 ? 3 3 4 ?
Dendrophidion pericarinatum 1 ? 0 3 ? ? 4 ? 2 3 4 ?
Drymarchon corais 1 ? 0 1 0 ? 0 ? 3 3 3 ?
Drymobius margaritiferus 1 ? 0 2 ? ? 2 ? 3 3 4 ?
Drymobius rhombifer 1 ? 0 2 3 ? 3 ? 3 3 4 ?
Drymoluber
dichrous
Masticophis flagellum testaceus 1 ? 2 4 1 1 3 ? ? 3 ? ?
Masticophis taeniatus taeniatus 1 ? 0 2 0 0 2 ? 2 2 2 ?

Mastigodryas bifossatus 1 ? 0 3 ? ? 1 ? 3 2 4 ?
Mastigodryas boddaerti 1 ? 0 2 ? ? 2 ? 3 4 5 ?
Mastigodryas heathi 1 ? 0 3 ? ? 1 ? 3 3 4 ?
Mastigodryas melanolomus 1 ? 0 3 ? ? 3 ? ? 3 4 ?
Mastigodryas pleei 1 ? 0 3 ? ? 2 ? 3 3
Pseustes poecilonotus 1 ? 0 3 ? ? 3 ? 4 ?
Pseustes sulphureus 5 ? 0 1 ? ? 1 ? 5 5
Salvadora grahamiae 1 ? 2 4 ? ? 2 ? 3 3
Salvadora hexalepis 1 ? 5 5 ? ? 2 ? 3 3
Spilotes pullatus 1 ? 0 1 1 ? 2 ? 2 ?
4
?
3
3
3
?
?
?
?
?
?
?
Chironiuis bicarinatus 3 3 4 0 5 2 1 2 3 3 5 5
Chironius carinatus flavolineatus 3 5 5 2 5 2 0 3 4 5 5 5

Chironius flavolineatus 3 3 2 0 5 2 1 3 4 2 5 ?

Chironiusfitscus leucometapus 2 2 1 0 5 2 1 2 3 2 5
5
5
5
?
?
Chironius nonticola 5 3 4 0 5 5 1 1 3 4 5 5
452
HERPETOLOGICA
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APPENDIX III
Continued.
Chironius multiventris cochranae 1 1 1 0 0 1 1 2 3 2 5 5

Chironius multiventris foveatus 2 1 0 2 0 1 1 2 4 2 5 5
Chironius multiventris septentrionalis 2 1 0 2 0 1 1 2 3 2 5 5
Chironius quadricarinatus maculoventris 4 3 4 5 5 1 1 1 3 4 5 ?
Chironius quadricarinatus quadricarinatus 4 3 3 0 5 1 1 1 2 4 5 ?
Chironius vincenti 2 1 0 5 0 0 1 2 3 2 5 5
Coluber constrictor constrictor 5 1 ? ? ? 2 0 1 2 1 0 0
Dendrophidion dendrophis ? ? ? ? ? ? ? ? ? ? 0 0
Dendrophidion nuchalis ? 1 ? ? ? 1 0 1 1 0 0 0
Dendrophidion pericarinatum ? 1 ? ? ? 2 1 1 1 2 0 0
Drymarchon corais ? 3 ? ? ? 2 0 ? ? 3 0 0
Drymobius margaritiferus ? 3 ? ? ? 2 1 1 1 ? 0 0
Drymobius rhombifer ? ? ? ? ? ? ? ? ? ? 0 0
Drymoluber
dichrous
Masticophis flagellum testaceus 3 3 ? ?

Masticophis taeniatus taeniatus 4 2 ? ?
Mastigodryas bifossatus ? 3 ? ? ?
Mastigodryas boddaerti ? 5 ? ? ?
?
?
4
3
3
2
0
0
0 1 1 1 0
1 0 1 2 0
0 2 4 0
1 2 4 0
0
0
0
0
Mastigodryas heathi ? ? ? ? ? ? ? ? ? ? 0 0
Mastigodryas melanolomus ? ? ? ? ? ? ? ? ? ? 0 0
Mastigodryas
pleei
Pseustes poecilonotus ? 4 ? ? ?
Pseustes sulphureus ? ? ? ? ?
4
4
?
0
1
?
1 2 0 0
? ? ? 0
0
0
0
0
Salvadora grahamiae ? 4 ? ? ? 1 0 0 0 5 0
Salvadora hexalepis ? 4 ? ? ? 1 0 0 0 2 0
Spilotes pullatus ? 2 ? ? ? 2 0 5 5 2 0 0
December
2006]
HERPETOLOGICA
APPENDIX JV
Apomorphy List
453
(7X=6) 4 (645) 5 (9#>O) 7 (8=>9) 18 (5=X7) 21 (1>0) 22

(140) 8-4 17 (649) 18 (748) 19 (446) 25 (5>3) 26
(543) 27 (442) 7-4fuscusfuscus 3 (140) 16 (1X0) 18
Characters in the parsimony analysis were optimized

(849) 19 (647) 25 (340) 7-->fuscus leucometapus 1
(443) 8 (847) 10 (4=3) 13 (140) 32 (4=43) 34 (5=>4)
using ACCTRAN (accelerated transformation) and DELTRAN (delayed transformation) with the tree rooted at 8--6 1 (447) 4 (542) 6 (1-*0) 20 (144) 33 (6-*7)
6-1aevicollis 1 (748) 4 (241) 8 (8=47) 11 (443) 13
Coluber constrictor constrictor. Only those apomorphies
(140) 16 (1>0) 19 (443) 20 (447) 24 (547) 25 (5-4)
relevant to the monophyly of the genus Chironius or
species therein are listed. Changes that occur under both 26 (5->4) 28 (044) 6-*scurrulus 10 (4>6) 17 (6-*7) 24
(5=4 4) 25 (5=>7) 27 (4=>5) 30 (344) 32 (4>5) 34 (549)
ACCTRAN and DELTRAN optimization are shown with
36 (9--0) 16-15 1 (4=47) 3 (142) 24 (5--6) 25 (5=42)
a double arrow (=>). Changes that occur under DEL29 (9=>0) 30 (3=42) 15-*grandisquamis 1 (749) 4
TRAN optimization only are shown with a single arrow
(643) 14 (1=>3) 16 (3-*2) 17 (6=>8) 18 (548) 21 (140)
(). Apomorphy descriptions consist of: branch change
22 (1=>0) 24 (6>9) 32 (447) 33 (6>9) 15-14 3 (243)
(node numbers or taxon name; see Fig. 3) in bold and the
character numbers diagnostic of that branch change, with 7 (841) 26 (5=43) 34 (544) 14-4aurenti 2 (7=46) 10
(4-*>5) 13 (1-*2) 19 (4>2) 20 (1>9) 14->13 4 6 (3-l)
transformations in parentheses.
18 (5--6) 19 (4=>6) 20 (1=0) 26 (3=42) 27 (441) 30
20-19 6 (6--3) 18 (O-*4) 21 (5#.1) 22 (6X1) 23
(7=?0) 33 (2=>5) 34 (3-*5) 35 (0=?9) 36 (0=X~9) 19-*18 5
(2=>1) 34 (4>3) 13-10 3 (3=>5) 4 (6-7) 17 (6-*7) 19
(6=47) 24 (6-*7) 1O->multiventris cochranae 2 (7=48) 3
(5X=9) 14 (0-*1) 25 (9-*8) 30 (4X3) 18-*17 1 (1?.4) 2
(6-*7) 7 (9?>8) 16 (4-*3) 18 (4-*5) 25 (8=?7) 32 (2X~3)
(5=>6) 4 (7=>8) 7 (1=>0) 13 (1>6) 19 (7=>8) 24 (748) 25
17--5 6 (3-*1) 20 (2=>4) 31 (9?>5) 5-*bicarinatus 7
(241) 26 (2-1) 1O--multiventris multiventris 1 (748)
(8-*7) 8 (6a?5) 14 (1=0) 18 (5-*6) 23 (0X:2) 24 (5X?7)
7 (143) 16 (3=>2) 32 (443) 34 (342) 13-12 16 (344)
27 (6-7) 5--4 4 (6>4) 10 (4X5) 28 (0>4) 32 (3-4) 33
25 (2=43) 27 (140) 28 (044) 12-*multiventrisfoveatus
(5-*6) 4-*3 1 (4X5) 2 (7=?6) 7 (8?9) 16 (3?2) 1 (5=0)
10 (4-*5) 24 (6=>8) 25 (344) 26 (2-4) 33 (647) 12-41
3--carinatus carinatus 4 (4--3) 8 (8-*6) 9 (3X2) 20
1 (7=44) 2 (7=>8) 4 (648) 13 (1-*2) 17 (6-*7) 24 (643)
(4-*5) 24 (5-6) 25 (7->8) 3--2 9 (3X4) 24 (5--4) 26
11-multiventris septentrtonalis 11 (4=43) 19 (6>7) 24
(34 12) 32 (4=43) 33 (645) 11-vincenti 1 (4=>1) 4

(5=?6) 33 (6-*7) 34 (5=>6) 2- carinatus flavopictus
(5X6) 19 (4X5) 20 (4=?2) 24 (4X2) 25 (7X6) 26 (6>9)
27 (6X9) 32 (4?5) 34 (6X9) 2--carinatus spixi 5

(9?8) 8 (8X5) 10 (5=?4) 18 (5-*6) 25 (7-*8) 28 (4?0)
30 (3=>5) 4--exoletus 1 (4=>3) 3 (10=>) 4 (40=>) 5 (9X8)
7 (8=>5) 11 (4=>5) 17 (6X4) 18 (5=>3) 19 (4?5) 20
(4=>6) 24 (5X2) 27 (6X5) 30 (3X4) 33 (6X8) 17-16 20
(2X1) 25 (7X5) 27 (6=?4) 32 (3-4) 33 (5-*6) 16-*9 6
(8=49) 6 (1=>4) 7 (143) 10 (4-*5) 11 (446) 13 (245) 17

(7>8) 19 (644) 26 (2-l) 28 (449) 30 (140) 18-4 8
(8-*6) 9 (3=>2) 17 (645) 19 (443) 24 (5=>1) 26 (5-*6)
30 (3>1) 34 (5-*7) 1-*quadricarinatus maculoventris
8 (6=>5) 10 (4=>5) 11 (445) 16 (4->3) 20 (2>6) 27
(6-7) 28 (049) 33 (5-*6) 1-quadricarinatus quad-
ricarinatus 2 (6--7) 3 (1-*0) 4 (6>7) 6 (344) 10 (443)

(3-*1) 14 (1=>0) 16 (3X1) 9-*flavolineatus 1 (4X0)
2
27 (645)
33 (5=>4) 19--monticola 1 (1-*3) 3 (1-*0) 4
(7X8) 4 (6>8) 6 (1-*0) 13 (1X2) 19 (4-*5) 20 (1?0) 24
(6=>5) 5 (54=>0) 9 (344) 14 (0-) 16 (445) 18 (4--5) 24
(5X0) 30 (3X4) 32 (4X5) 33 (6>8) 34 (5X4) 9-*8 2
(5-*6) 26 (5-*6) 27 (6--8) 30 (4>9) 34 (5->7)
Herpetologica, 62(4), 2006, 453-465
?) 2006 by The Herpetologists' League, Inc.
A NEW SPECIES OF THE BOKERMANNOHYLA PSEUDOPSEUDIS
GROUP FROM CENTRAL BAHIA, BRAZIL (AMPHIBIA, HYLIDAE)

LUCIANA LUGLI AND CtLIO F. B. HADDADI
Departamento de Zoologia, IB, Universidade Estadual Paulista, C. P. 199, 13506-900 Rio Claro, Sio Paulo, Brasil
ABSTRACT: A new species of treefrog is described from Chapada Diamantina in the central State of
Bahia, Brazil. The new species is a member of the Bokermannohyla pseudopseudis group, characterized by
its small to medium size, head as long as wide, snout short, rounded in dorsal view and nearly truncate in lateral
view, eyes and tympanum large, and forearm and prepollex well developed. The new species can be
distinguished from the other species of the B. pseudopseudis group by its voice and the shape of its head and
snout. It is smaller than B. saxicola and B. pseudopseudis, and has a less developed forearm and prepolex than
B. ibitiguara males. The tadpole and vocalizations are described and information on natural history is provided.
Key words: Bokeenrannohyla; Chapada Diamantina - Bahia; New species
CORRESPONDENCE: e-mail, haddad@rc.unesp.br

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Herpetologists' League

Phylogenetics of the Genus Chironius Fitzinger, 1826 (Serpentes, Colubridae) Based on

Herpetologica, 62(4), 2006, 435-453

? 2006 by The Herpetologists' League, Inc.

PHYLOGENETICS OF THE GENUS CHIRONIUS FITZINGER, 1826

(SERPENTES, COLUBRIDAE) BASED ON MORPHOLOGY

Key words: Chironius; Colubridae; Morphology; Phylogeny; Snakes; Systematics; Taxonomy

odd number of dorsal scale rows (as originally

CORRESPONDENCE: e-mail, jhollis@unm.edu

collected from 44 alcohol-preserved museum

group species. Exceptions to this are: females

Characters and size measurements were

Counted following the method of Dowling (1951).

on size (character 3; see below); consequently,

significantly dependent when linearly regressed on

6. Mean number of scale rows keeled, counted at

7. Apical pits, entirely absent/present on neck only

8. Mean number of supralabials; 6.50 (0) to 9.50 (9).

9. Modal condition of supralabials entering orbit; 3 +

contact the orbit. Gap-weighted.

10. Mean number of infralabials; 8.50 (0) to 12.50 (9).

11. Mean number of infralabials contacting genials;

14. Modal condition of temporal scales; I + 1 (0); 1 + 2

[0] [1] [2] [3]

16. Mean number of postcephalics; 5.93 (0) to 12.33

22. Modal number of dorsal scale rows at midbody; 10

Masticophis flagellum testaceus, and Masticophis

17. Mean number of maxillary teeth; 17.0 (0) to 44.2

constrictor constrictor, Masticophis flagellum tes- (9). Frequency coded.

36. Raised vertebral keel; absent (0) or present (9).

Analysis71 Chironnius multiventris septentnonalis

I used PAUP* 4.0blO (Swofford, 2002) to

79 1 3 Chironius multiventns foveatus

74 Chironius multiventris cochranae

16 < Chironius fuscus fuscus

Chironius fuscus leucometapus

Chironius cannatus flavopictus

18 Chironius cannatus carinatus

_ 19 77 Chironius quadncarinatus maculoventris

Characters were optimized in PAUP* using

I used MrBayes 3.1.2 (Huelsenbeck and

dicate support from 1000 nonparametric bootstraps.

RESULTS AND DISCUSSION

trees (Fig. 3) with the following properties:

both replicates; burn-in was conservatively

86 Chironius multiventris septentrionalis

7412 Chironius multiventbis foveatus

13 Chironius muttiventris muttiventris

3/ 2 Chionius mu/tiventns foveatus

8 /1 Chironius multiventris cochranae

6 Chironius fuscus fuscus

14 / Chironius multivent/rs multiventns

87 Chironius fuscus leucometapus

1 86 Chironius carinatus tiavopictus

3 2 5 Chironius fuscus fuscus

55 Chironius quadricarinatus macubventns

5 0 Chironius cnat/naus spixi

2 2 3 /2 4 Chironius cnatrnaus carnnatus

2 / 1 5 14/ Chironius exoletus

Masticophis taeniatus taeniatus

_ 19 / 5 3 Chironius quadricar/natus maculovent/s

Coluber constnctor constrictor

FIG. 4.-Apomorphy support within ChironiusFiG.

dentary teeth (18: 0--4), and distal calyces