Herpetologists' League

Revision of the Micrurus frontalis Complex (Serpentes: Elapidae)

Author(s): Nelson Jorge da Silva, Jr. and Jack W. Sites, Jr.
Source: Herpetological Monographs, Vol. 13 (1999), pp. 142-194
Published by: Herpetologists' League
Stable URL: http://www.jstor.org/stable/1467062
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HerpetologicalMonographs,13, 1999, 142-194
? 1999 by The Herpetologists' League, Inc.

REVISION OF THE MICRURUS FRONTALIS COMPLEX

(SERPENTES: ELAPIDAE)
NELSON JORGE DA SILVA, JR.,' AND JACK W. SITES, JR.2
'Centro de Estudos e Pesquisas Biologicas, Departamento de Biologia, Universidade Cat6olicade Goias,
Ave. Universitdria, 1440-Setor Universitdrio-74605-010-Goidnia, GO-BRAZIL.
E-mail: herp@nutecnet.com.br
2Department of Zoology and Monte L. Bean Museum, Brigham Young University, Provo, UT 84602 U.S.A.
E-mail: jack.sites@byu.edu

ABSTRACT: The New World coralsnake genus Micrurus has been taxonomically problematic due
to extreme variation in color pattern and highly conserved morphological variation. These problems
are exemplified in the Micrurus frontalis "group", and we present a monographic revision based on
an analysis of over 2100 specimens (including all available types). We recognize the following seven
species on the basis of unambiguous combinations of characters: M. altirostris, M. baliocoryphus,
M. brasiliensis, M. diana, M. frontalis, M. pyrrhocryptus, and M. tricolor. A complete synonymy is
given for each taxon, as well as a re-description of the holotypes. We also characterize morphological
variation for each, and provide distribution maps and a key to the species.

RESUMO:As serpentes corais do Novo Mundo, genero Micrurus, tem sido taxonomicamente
problemSticas devido a extrema variacao de padr6es de cores e variagao morfologica altamente
conservadora. Esses problemas sao exemplificados no "grupo" Micrurus frontalis, e nos apresenta-
mos uma revisao monografica baseada na analise de mais de 2100 esptecimes (incluindo todos os
tipos disponfveis). Nos reconhecemos as seguintes sete especies baseado na combina9ao nao am-
bfgua de caracteres: M. altirostris, M. baliocoryphus, M. brasiliensis, M. diana, M. frontalis, M.
pyrrhocryptus e M. tricolor. Uma sinonfmia completa e oferecida para cada taxon, bem como a
redescri5ao dos holotipos. N6s tambem caracterizamos a variacao morfolo6gica de cada um e apre-
sentamos mapas de distribuiVao e uma chave para as especies.

Key words: Serpentes; Elapidae; Micrurus; Morphological variation; Species status

CORALSNAKES (Serpentes, Elapidae) but the Micrurus frontalis complex is es-

range from the southern United States to
southern South America and are arranged
into three genera: Leptomicrurus Schmidt,
Micruroides Schmidt, and Micrurus Wag-
ler. The vast majority of the species were
originally recognized and described on the
basis of unique colors and patterns, and
this is generally true for the entire genus
(Roze, 1996).
Coralsnakes can generally be divided
into two groups based on the color pat-
tern: (1) monadal coralsnakes-with only
one black ring combined with red and
white (or yellow) rings (red-white-black-
white-red); and (2) triadal coral snakes-
with three black rings combined with red
and white (or yellow) rings (red-black-
white-black-white-black-red). The South
American triadal coralsnakes include 17
currently recognized species (Campbell
and Lamar, 1989; Roze, 1996; Savage and
Slowinski, 1992; Slowinski, 1995).
Most of the triad taxa are poorly known,
142
pecially problematic. This is an assemblage
of poorly defined taxa (Campbell and La-
mar, 1989; Roze, 1983, 1994, 1996; Scroc-
chi, 1990) ranging from the Cerrado for-
mations of central Brazil and Bahia,
through eastern Bolivia and the temperate
areas of southeastern Brazil, to the Pam-
pean regions of south Brazil, Uruguay,
Paraguay, and Argentina (Fig. 1). It has the
most southerly distribution, and is also the
most diverse (currently four subspecies are
recognized, but there have been as many
as eight) of all the South American triadal
coralsnakes.
Since the original description by Du-
meril et al. (1854), all later specimen de-
scriptions and taxonomic arrangements
have done little to clarify the problems re-
lated to morphological variation, geo-
graphic ranges, and species boundaries.
Most authors used small sample sizes, and
revisions were based on inadequate cov-
erage of the literature, failure to examine
1999] HERPETOLOGICAL MONOGRAPHS 143

E
i:i'!i:i II?!
-
iiiil : II : A
G
@*t
B

C..

tI- F

FIG. 1.-Generalized distribution of Micrurus frontalis complex, with color patterns (gray shading = red)
"diagnosing" the following recognized taxa: A. Micrurusf frontalis; B. Micrurus f. altirostris; C. Micrurus f.
baliocoryphus; D. Micrurus brasiliensis; E. Micrurus f. diana; F. Micrurus p. pyrrhocryptus; G. Micrurus p.
tricolor.
144
144 HERPETOLOGICAL MONOGRAPHS
HERPETOLOGICAL MONOGRAPHS
holotypes, and in some cases lack of
knowledge of basic coralsnake biology
(Amaral, 1944; Barrio and Miranda, 1966;
Roze, 1967, 1970, 1983, 1994, 1996;
Scrocchi, 1990, 1995; Slowinski, 1995;
Striissmann and Sazima, 1993). Others
have published checklists and bibliograph-
ic compilations (Boulenger, 1896; Golay,
1985; Roze, 1967, 1983; Schmidt, 1936),
but ignored details of geographic variation.
Among recent taxonomic changes, Scroc-
chi (1990) elevated M. frontalis pyrrho-
cryptus to a full species (with M. frontalis
tricolor as a junior synonym), interpreted
M. frontalis multicinctus as an intergrade
between M. frontalis frontalis and M. fron-
talis altirostris, and placed M. frontalis bal-
iocoryphus in synonymy with M. frontalis
mesopotamicus. In a few words, Strtiss-
mann and Sazima (1993) resurrected M.
frontalis tricolor as full species without any
comparative work, reference to museum
material, or use of appropriate literature.
Roze (1994) elevated M. frontalis diana to
a full species, placed M. frontalis mesopo-
tamicus as a junior synonym of M. frontalis
baliocoryphus, changed the status of M.
pyrrhocryptus to M. pyrrhocryptus pyr-
rhocryptus to accomodate M. frontalis tri-
color as M. pyrrhocryptus tricolor, and res-
urrected M. f multicinctus as a subspecies
of M. frontalis. None of these studies were
based on solid data analyses or, a priori de-
fense of a particular species concept as a
testable hypothesis, and consequently they
have not contributed to the clarification of
species boundaries or phylogenetic rela-
tionships of diagnosable units in this group.
The extensive museum material now avail-
able suggests that recognition of subspecies
may be based on nothing more than early
descriptions of clinal variation in color pat-
terns (clinal variation is evident in M. fron-
talis frontalis and in M. frontalis altirostris),
and no constancy appears to be evident for
characters previously thought to be diag-
nostic.
Because of their contiguous geographic
distributions, better representation in mu-
seum collections, uncertainties of species
boundaries, and nomenclatural instability,
the Micrurus frontalis complex was chosen
for detailed studies of morphological and
[No.
[No. 13
color pattern variation. Our goals in this
study are to describe variation in meristic,
cranial, and hemipenial characters as com-
pletely as possible for all recognized taxa,
test the validity of the currently recognized
species and subspecies, and present syn-
onymies, distributions and a key to the taxa
we recognize. In another paper we esti-
mate phylogenetic relationships within this
group relative to other South American tri-
ad species (Jorge da Silva and Sites, in re-
view).
MATERIAL AND METHODS
Museum material for the M. frontalis
complex examined for morphological data
is deposited in collections in Brazil, Boliv-
ia, Uruguay, Paraguay, Argentina, France,
and the United States. The following col-
lections loaned or provided access to spec-
imens (acronyms follow Leviton et al.,
1985).
AMNH-American Museum of Natural
History, New York, NY, USA.
ANSP-Academy of Natural Sciences,
Philadelphia, PA, USA.
CCG-Coleccion Carlos Grisolfa,
Buenos Aires, Argentina.
CEPB-Centro de Estudos e Pesquisas
Biol6gicas-Universidade Catolica de Goias,
Goiania, GO, Brazil.
CHINM-Coleccion Herpetologica del
Instituto Nacional de Microbiologfa Carlos
G. Malbran, Buenos Aires, Argentina. This
collection once belonged to A. Barrio and
was donated to the MACN, but it is not
yet fully incorporated to this museum.
CHC-Coleccion Herpetologica Corri-
entes, Corrientes, Argentina.
COBAP-Cole,ao Ofiologica da Bacia
do Alto Paraguai-Serpentario da Divisao
de Toxicologia e Farmacologia da Secre-
taria da Saude do Estado do Mato Grosso
do Sul, Campo Grande, MS, Brazil.
CZVR-Coleccion de Vertebrados-Rep-
tiles, Facultad de Humanidades y Cien-
cias, Universidad de la Republica, Mon-
tevideo, Uruguay.
FML-Fundacion Miguel Lillo, San
Miguel de Tucuman, Argentina.
IB-Instituto Butantan, Sao Paulo, SP,
Brazil.
MACN-Museo Argentino de Ciencias
1999] HERPETOLOGICALMONOGRAPHS
Naturales Bernardino Rivadavia, Buenos
Aires, Argentina.
MCP-Museu de Ciencias e Tecnolo-
gia-Pontiffcia Universidade Catolica do
Rio Grande do Sul, Porto Alegre, RS, Bra-
zil.
MCZ-Museum of Comparative Zool-
ogy, Harvard University, Cambridge, MA,
USA.
MHNCI-Museu de Historia Natural
Capao da Imbuiia, Curitiba, PR, Brazil.
MLP-Museo de La Plata, La Plata, Ar-
gentina.
MNHN-Museum National d'Histoire
Naturelle, Paris, France.
MNHNP-Museo Nacional de Historia
Natural del Paraguay, Asuncion, Paraguay.
MNRJ-Museu Nacional do Rio de Ja-
neiro, Rio de Janeiro, RJ, Brazil.
MPHNLP-Museo Provincial de His-
toria Natural de La Pampa, La Pampa, Ar-
gentina.
MZUSP-Museu de Zoologia-Universi-
dade de Sao Paulo, Sao Paulo, SP, Brazil.
NKR-Museo de Historia Natural Noel
Kempff Mercado, Santa Cruz de la Sierra,
Bolivia.
UMMZ-University of Michigan Mu-
seum of Zoology, Ann Arbor, MI, USA.
A total of 2110 specimens of the Micru-
rus frontalis complex was examined and
2060 specimens were used in the statistical
analyses. From this total 626 (30.5%) were
M. frontalis, with 494 males and 132 fe-
males; 1084 (52.7%) M. altirostris, with
765 males and 319 females; 98 (4.6%) M.
baliocoryphus, with 81 males and 17 fe-
males; 11 (0.5%) M. brasiliensis, with 8
males and 3 females; 7 M. diana (0.3%),
with 5 males and 2 females; 215 (10.5%)
M. pyrrhocryptus, with 196 males and 19
females; and 19 M. tricolor (0.9%), with
13 males and 6 females. The remaining
specimens (50) were not included because
they lacked more than 3 morphological
characters due to damage, incomplete
bodies, or were collected during the final
revisions of this study. The following set of
characters were recorded from all but
these 50 snakes.
Head Coloration
This includes the overall color pattern
of the snout (internasals, nasals, and ros-
'45
145
tral), head (prefrontals, supraoculars, fron-
tal, parietals, and temporals), mouth (su-
pralabials and infralabials), and gular
region (infralabials, genials and first ven-
trals). Also it includes the color and dis-
tance (number of scales) from the poste-
rior of the head to the first triad.
Meristic Characters and Triad Pattern
The ventral and subcaudal scales were
counted following Dowling (1951). The
position of a character along the body was
quantified by recording the number of the
ventral or subcaudal scales opposite to
which it was situated. Triad sizes were
quantified on the basis of single rings and
the triads as a whole, by counting the
number of dorsal scales (along the mid-
vertebral line) involved, and also measur-
ing each to the nearest 0.1 mm. We used
the median triad as a standard for all spec-
imens, and these were quantified as fol-
lows (see Fig. 1): length of middle black
ring; length of two lateral black rings;
length of two white/yellow rings; and
length of red rings. The snout-vent length-
(SVL) and tail length (TL) were measured
to the nearest 0.1 mm, and the number of
triads recorded as the total number of tri-
ads on the body plus the number on the
tail, including any incomplete triad.
Cranial Osteology
Skulls were prepared from museum
specimens, and five specimens of each
species were used for the general descrip-
tion except M. brasiliensis (n = 2), M. di-
ana (n = 1), and M. tricolor (n = 3) due
to the small representation of these taxa in
museum collections. Selected skulls were
drawn using a Zeiss Stemi SV 6 stereo-
microscope.
Hemipenial Morphology
Everted organs were examined in ten
specimens of each species for the general
description except M. brasiliensis (n = 2),
M. diana (n = 1), and M. tricolor (n = 6)
for the same reasons of skull preparations.
When necessary, specimens were prepared
following the method of Pesantes (1994),
and representative examples were drawn
using a Zeiss Stemi SV 6 stereomicro-
146 HERPETOLOGICAL MONOGRAPHS
HERPETOLOGICAL MONOGRAPHS LNo.
[No. 13

scope. Hemipenial terminology follows crurus (Slowinski, 1995), and the fact that
Dowling and Savage (1960) and Savage most material is available only as museum
(1997). vouchers, we will use concordance as a
necessary criterion for species recognition.
Statistics Species are defined under this concept on
Species were first designated by color- the basis of their constituent populations
ation patterns of the head and triad, and sharing at least two unique character
these were quantified by univariate and states. Ideally concordance of characters
multivariate statistical treatments. should be defined by the genealogical re-
Standard univariate descriptive statisticslationships of the alternative states (Avise
were summarized with the PROC and Ball, 1990; Avise and Wollenberg,
MEANS option of SAS (1990) for all var- 1997), but we use simple presence/ab-
iables by species, and are summarized in sence as a surrogate measure. We return
Table 1. The same variables were also an- to these issues in the Discussion section.
alyzed by three multivariate approaches,
including a canonical discriminant analysis Species Accounts
(CDA), a canonical correlation analysis A summary of the morphological data
(CCA), and a principal component analysis related to scutelation and triad pattern and
(PCA). Prior to the multivariate analyses, number, and quantitative features of cra-
we tested for significant correlations be- nial osteology and hemipenial morphology
tween all paired combinations of charac- are provided with descriptions of each tax-
ters; all individuals were combined across on.
sex and preliminary species boundaries for
these tests (n = 2060). This allowed us to RESULTS
eliminate redundant (i.e., significant cor- Our review recognizes the following
related) characters (see Results) and be- taxa of the Micrurus frontalis complex as
cause most characters tested significant for full species: M. frontalis, M. altirostris, M.
deviations from normality, morphometric baliocoryphus, M. brasiliensis, M. diana,
variables were log-transformed, and meris- M. pyrrhocryptus, and M. tricolor.
tic characters were square-root trans- Tables 1 and 2 summarize the descrip-
formed. The CDA was then used to iden- tive statistics for all meristic and morpho-
tify particular combinations of characters metric data collected for this study. These
that maximized separation of the samples, data reveal interspecific overlap in the ex-
and the CCA was used to maximize the treme values for many characters, even
separation between groups (species) rela- those showing pronounced differences be-
tive to within-group variance, taking into tween means (i.e., compare means and
account any remaining within-group cor- ranges for ventral scale counts between M.
relation between characters. The PCA was frontalis and M. altirostris [Table 1]).
also used to reduce the number of re- Correlation tests revealed significant
maining variables to a small number of in- positive correlations for head-length (HL)
dices (called principal components) rep- vs. snout-vent length (SVL) (Pearson cor-
resenting linear combinations of the relation coefficient = 0.859), HL vs. tail-
original variables (Manly, 1991). length (TL) (c = 0.826), and SVL vs. TL
(c = 0.915; P < 0.0001 in all cases).
Species Concepts Among the meristic characters, highly sig-
A number of alternative operational nificant positive correlations were also
species concepts can be framed as hypoth- found for anterior vs. posterior red ring
eses for empirical testing (reviewed by length (c = 0.843), anterior vs. posterior
Sites and Crandall, 1997), but given the black ring length (c = 0.728), and anterior
prior importance of color pattern in spe- vs. posterior white ring length (c = 0.785;
cies recognition in this group (Roze, 1996), P < 0.0001 in all cases). We therefore
the conservative nature of some data par- eliminated correlated variables from the
titions within monophyletic groups of Mi- data matrix by retaining only SVL, and the
19991
199IEPTLGCLMNGAH HERPETOLOGICAL MONOGRAPHS
anterior ring lengths (for red, black, and
white rings) in the multivariate analysis.
We found one significant negative corre-
lation between the total number of triads
and the length of the middle black ring (c
= -0.759; P < 0.0001), but retained both
of these variables because of their unam-
biguous phenotypic expression and high
discriminant power.
To maximize separation of samples in
multivariate space, the eight remaining
transformed variables were subjected to a
canonical discriminant analysis, by species
(n = 7) by sex (7 x 2 = 14). A matrix of
pairwise Mahalanobis distances between
all species-sex combinations revealed sig-
nificant divergence among groups across
the entire matrix (Wilks' Lambda = 0.073;
F = 62.55 with 104 df; P < 0.0001), and
small between-sex D values within most
species relative to between-species D val-
ues.
Table 3 summarizes the eigenvalues for
the eight canonical vectors, the proportion
of total variation explained by each vector,
and the total-sample standardized canoni-
cal coefficients for the first two canonical
vectors. The eigenvalues of the first two
canonical vectors explain >80% and >14%
of the total variation (>94% cumulative).
The characters most influential on the first
vector are the number of triads (STRIAD)
and length of the middle black ring
(SMIDBL in Table 3), while on vector
two, the dominant variables are SMIDBL
and number of ventral scales (SVENSC).
Figure 2 illustrates the distribution of sam-
ple means (for the 14 classes) along the
two axes represented by canonical vari-
ables (CVs) 1 and 2. The first CV clearly
separates, by both sexes, M. altirostris, M.
baliocoryphus, M. pyrrhocryptus, and M.
tricolor along an axis representing an in-
verse relationship between SMIDBL and
STRIAD. At one extreme, M. altirostris
(squares in Fig. 2) is characterized by a
large number of triads, each with a short
middle black ring, while M. pyrrhocryptus
(hexagons) shows exactly the opposite pat-
tern. The second CV clearly distinguishes
this entire group from a second group (M.
brasiliensis, M. diana, and M. frontalis);
the two groups are separated by a combi-
4
147
nation of middle black ring lengths and
ventral scale numbers (Table 3). Snakes
with a low average SMIDBL have a high
SVENSC (M. brasiliensis, M. diana, and
M. frontalis), and vice-versa (all others in
Fig. 2). Even though there is considerable
overlap in the ranges of some groups (i.e.,
all 3 species of the 'frontalis" group), class
means are clearly distinct across all species
by this conservative analysis. Results were
the same with the other multivariate analy-
ses (data not shown).
As demonstrated by Savitzky (1978),
Scrocchi (1993), and Slowinski (1995) cor-
alsnakes have very few informative mor-
phological characters, and many of these
are autapomorphies found in species with
extremely specialized diet or habitat re-
quirements (i.e., M. surinamensis; see
Campbell and Lamar, 1989; Jorge da Silva
and Sites, in review; Savitsky, 1978; Slow-
inski, 1994). We present a general over-
view of the conservative nature of suites of
morphological characters in the triadal Mi-
crurus, and then complete synonymies,
descriptions, diagnoses, summaries of var-
iation, illustrations, and range maps for the
seven species recognized in this revision.
All Micrurus have 15 rows of dorsal
smooth scales without reduction, anal
plate divided (except M. hemprichii) and
subcaudals also divided. The head scute-
lation is very similar to any colubrid snake.
The snout is rounded with very small eyes
and the head is hardly distinguishable
from the neck. Behind the rostral scale are
2 internasals, 2 nasals, 2 prefrontals, 1
frontal, 2 supraoculars, and 2 parietals. Lo-
reals are absent. There are 1 preocular, 2
postoculars, 1 + 1 temporals (rarely 1 +
2); 7-8 supralabials and infralabials; 1
mental; 2 anterior genials; and 2 posterior
genials. Body sizes are extremely variable,
but within the M. frontalis complex they
range from 260 to 1641 mm (total length).
The skull of any of the species of the
study group has the following general
characteristics: the premaxilla and septo-
maxillae are compact and robust bones;
there is a considerable inter- and intra-spe-
cific variation of the nasals and vomer
without distinct fixed characters; the pre-
frontals may or may not be in contact with
148 HERPETOLOGICAL MONOGRAPHS [No. 13

TABLE 1.-Summary of means, standard deviations (second line, in parentheses), and ranges for all samples
of the Micrurus frontalis complex examined in this study. Characters are defined in the text (scale counts and
measurements), and samples are separated by sex.
Meristic Morphometric
Taxon (n) Ventrals Subcaudals Triads HL SVL TL

Males
Micrurus frontalis (494) 225.03 21.90 13.62 21.54 830.77 50.92
(8.31) (1.99) (1.31) (6.21) (288.51) (16.00)
(212-238) (19-27) (10-15) (8-65) (205-1551) (13-90)
Females
Micrurus frontalis (132) 224.63 21.03 13.13 17.55 632.16 37.03
(11.51) (2.82) (1.36) (4.63) (203.07) (12.12)
(213-234) (15-26) (11-14) (8-31) (222-1164) (11-74)
Males
Micrurus altirostris (765) 211.32 20.42 15.29 17.69 628.20 39.21
(7.18) (2.25) (1.44) (4.53) (184.32) (11.01)
(199-230) (15-25) (13-18) (5-31) (223-1235) (11-75)
Females
Micrurus altirostris (319) 209.97 18.75 15.15 15.98 528.50 32.81
(8.84) (2.81) (1.30) (5.00) (159.21) (10.71)
(119-254) (14-23) (14-18) (6-75) (177-829) (10-65)
Males
Micrurus baliocoryphus (81) 220.79 22.81 13.15 19.57 769.95 46.32
(8.51) (3.07) (1.63) (4.97) (241.52) (12.39)
(210-230) (18-27) (9-16) (8-35) (253-1374) (17-75)
Females
Micrurus baliocoryphus (17) 220.24 20.12 13.00 15.60 558.12 33.65
(7.38) (2.74) (1.00) (3.08) (162.51) (10.12)
(208-233) (17-23) (11-15) (10-21) (262-818) (16-51)
Males
Micrurus brasiliensis (8) 227.38 28.25 13.00 22.50 801.00 46.25
(9.23) (18.67) (2.07) (8.05) (304.43) (17.34)
(218-234) (20-25) (11-14) (14-39) (352-1431) (28-82)
Females
Micrurus brasiliensis (3) 232.33 19.67 10.50 17.00 575.33 27.00
(4.16) (3.51) (0.58) (5.57) (239.14) (8.19)
(229-237) (16-23) (12-13) (11-22) (302-746) (18-34)
Males
Micrurus diana (5) 222.50 23.00 10.50 21.00 795.50 47.50
(9.98) (2.00) (0.58) (3.56) (134.67) (5.51)
(215-229) (22-24) (10-11) (17-24) (627-954) (42-54)
Females
Micrurus diana (2) 227.00 23.00 10.50 18.50 646.50 47.50
(0) (0) (0.71) (3.54) (229.81) (7.07)
(227-227) (19-19) (10-11) (16-21) (484-809) (32-42)
Males
Micrurus pyrrhocryptus (196) 228.20 24.31 8.11 17.33 683.24 42.47
(6.21) (2.73) (1.16) (4.65) (203.87) (11.82)
(218-239) (20-28) (6-11) (7-31) (214-1175) (10-66)
Females
Micrurus pyrrhocryptus (19) 234.79 22.16 9.42 17.63 673.95 38.00
(7.81) (1.89) (4.87) (4.49) (167.90) (9.30)
(221-251) (24-29) (8-10) (10-28) (403-950) (15-51)
1999] HERPETOLOGICAL MONOGRAPHS 149

TABLE1.-Continued.

Meristic Morphometric
Taxon (n) Ventrals Subcaudals Triads HL SVL TL

Males
Micrurus tricolor (13) 224.00 25.92 9.92 21.85 910.62 60.31
(6.99) (1.50) (0.95) (3.69) (147.11) (10.92)
(220-234) (24-29) (8-10) (17-30) (680-1100) (45-81)
Females
Micrurus tricolor (6) 223.00 22.17 10.12 15.83 621.00 39.33
(4.60) (2.48) (0.98) (4.79) (210.33) (12.82)
(217-231) (19-26) (9-11) (11-22) (410-889) (26-57)

each other through a dorsomedial process;
the frontals are triangular in shape with a
pointed posterior end; the braincase (pa-
rietals) is narrow and long with pointed or
rounded posterior end; the dorsal skull ta-
ble is triangular in shape with the anterior
oblique lateral ridges more evident in
some species that attain large body and
head sizes; the occipital region is short; the
transverse ridge of the supraoccipitals is
moderately developed; the tabular is short
and variable in shape sometimes touching
the parietals; the quadrate is short and tri-
angular with a variable size and shape of
the surface of contact with the tabulars.
The maxilla is a short and compact bone
that bears the short and recurved venom
fang. The contact maxilla-ectopterygoid is
broad and at mid-frontals level. There are
8-10 palatine and 3-5 pterygoid teeth.
The compound bone is long without a
pseudocoronoid process. There are 9-11
dentary teeth. Very few inter-specific char-
acters are diagnosable (see also Scrocchi,
1993).
The hemipenis is bilobed, semicapitat-
ed, with the sulcus spermaticus running
medially (centripetally) along the arms to
their tips. The ornamentation is limited to
spines with the absence of fringes, flounc-
es, or calyces, and within the M. frontalis
complex, inter-specific characters are
poorly defined.
SPECIES ACCOUNTS
Micrurus frontalis (Dumeril, Bibron,
and Dumeril)
Elaps frontalis Dumeril, Bibron, and Du-
meril 1854:1223.
Holotype: Two syntypes MNHN 854
(M) and MNHN 578 (M) are still at the
MNHN herp collection. The specimen
MNHN 578 (M) is not a M. frontalis (it is
M. isozonus, see below). We suggest that
the specimen MNHN 854 (M) be assigned
as the lectotype.
Type-locality: Corrientes and Misiones.
We suggest changing the type-locality to
Lagoa Santa, Minas Gerais, Brazil.
Elaps baliocoryphus Cope 1859:346
Elaps frontalis Boulenger 1896:427
Micrurus frontalis Amaral 1925:19
Micrurus frontalis Amaral 1929:230
Micrurus frontalis frontalis Schmidt 1936:
199
Micrurus lemniscatus frontalis Amaral
1944:92
Micrurus frontalis frontalis Roze 1967:24
Micrurus frontalis frontalis Hoge and Ro-
mano 1979:395
Micrurus frontalis frontalis Roze 1983:323
Micrurus frontalis frontalis Golay 1985:34
Micrurus frontalis frontalis Campbell and
Lamar 1989:115
Micurus frontalis frontalis Roze 1994:179
Micrurus frontalis frontalis Roze 1996:172
The original type series apparently in-
cluded eight specimens ("Le Museum pos-
sede quatre bocaux qui renferment chacun
deux individus."; Dumeril et al., 1854) but
the authors based their description on
three specimens making all of them syn-
types. One was collected by Castelnau in
Corrientes and Misiones, Argentina. A sec-
ond specimen was collected by Balper-
thuis in Cote ferme, Venezuela, and the
third was collected by Claussen in Brazil,
150 150 ---- MONOGRAPHS
HERPETOLOGICAL
HERPETOLOGICAL - - - .- - [No.
[No. 13

TABLE 2.-Summary of means, standard deviations, and ranges for the size (in number of dorsal scales) of
different rings in the middle triad, of all Micrurus frontalis complex specimens examined in this study. Data
are arranged as in Table 1, and are identified as follows: ANTRD and POSTRD = anterior and posterior red
rings; ANTBL, MIDBL, and POSTBL = anterior, middle, and posterior black rings; and ANTWH and
POSTWH = anterior and posterior white rings (see Figure 1).

Taxon ANTRD ANTBL ANTWH MIDBL POSTWH POSTBL POSTRD

Males
Micrurus frontalis (494) 7.00 2.40 1.76 2.94 1.68 2.43 6.82
(2.17) (0.65) (0.62) (0.77) (0.60) (0.58) (2.38)
(1-22) (1-10) (0.5-4.5) (1-9) (0.5-5.0) (1-6) (2-32)
Females
Micrurus frontalis (132) 6.25 2.84 2.20 3.00 2.16 2.84 6.27
(1.91) (0.61) (0.75) (1.21) (0.74) (0.65) (1.80)
(2-13) (2-4) (0.5-4) (1.5-13) (1-4) (1-4.5) (2-11.5)
Males
Micrurus altirostris (765) 4.76 2.48 1.31 2.96 1.28 2.43 4.75
(1.46) (0.60) (0.45) (0.71) (0.43) (0.57) (1.45)
(1-11) (1-9) (0.5-4) (1-7) (0.5-3) (1-9) (1-11)
Females
Micrurus altirostris (319) 4.50 2.60 1.40 3.04 1.32 2.60 4.58
(1.34) (0.54) (0.42) (0.80) (0.40) (0.58) (1.33)
(1-9) (1.5-5.7) (0.5-2.5) (1.5-9) (0.5-2.5) (1.5-7.3) (1-8)
Males
Micrurus baliocoryphus (81) 5.20 2.52 1.71 4.64 1.59 2.51 5.51
(2.14) (0.61) (0.50) (1.21) (0.52) (0.54) (2.28)
(1-10) (1-4) (1-3) (2-7) (0.5-3) (1-3) (4-12)
Females
Micrurus baliocoryphus (17) 5.18 2.71 1.68 5.06 1.74 2.56 5.41
(1.63) (0.59) (0.47) (1.21) (0.40) (0.79) (2.80)
(1-8) (2-4) (1-2.5) (2-7) (1-2.5) (2-5) (1-42)
Males
Micrurus brasiliensis (8) 5.06 2.88 2.56 3.56 2.69 2.69 5.19
(1.29) (0.58) (0.86) (0.98) (0.84) (0.56) (1.28)
(4-7.5) (2-4) (1-3.5) (2.5-5) (1-3.5) (2-3) (4-7)
Females
Micrurus brasiliensis (3) 5.83 2.83 3.0 2.67 3.0 2.83 6.50
(1.76) (0.29) (0.50) (0.29) (0.50) (0.29) (0.87)
(4-7.5) (2.5-3) (2.5-3.5) (2.5-3) (2.5-3.5) (2.5-3) (6-7.5)
Males
Micrurus diana (5) 7.50 3.38 3.25 3.88 3.13 3.63 7.38
(1.47) (0.48) (0.87) (2.10) (0.85) (0.95) (1.11)
(6-9.5) (3-4) (2-4) (2.5-7) (2-4) (3-5) (6.5-9)
Females
Micrurus diana (2) 7.75 3.50 4.0 2.5 4.0 3.25 7.25
(0.35) (0) (0) (0.71) (0) (0.35) (0.35)
(7.5-8) (3.5-3.5) (4-4) (2-3) (4-4) (3-3.5) (7-7.5)
Males
Micrurus pyrrhocryptus (196) 9.80 3.95 2.12 8.81 2.11 3.99 9.35
(3.80) (0.82) (0.67) (2.18) (0.61) (1.14) (3.54)
(1-20) (2-6) (1-6.5) (2-15) (1-6.5) (2-5) (1-19)
Females
Micrurus pyrrhocryptus (19) 10.11 4.08 2.02 8.79 2.05 4.05 9.97
(4.24) (0.87) (0.56) (2.31) (0.60) (0.72) (4.26)
(2-17) (2.5-5) (1-3) (5.5-14) (1-3) (2.5-5) (2-20)
1999] HERPETOLOGICAL MONOGRAPHS 151

TABLE2.-Continued.

Taxon ANTRD ANTBL ANTWH MIDBL POSTWH POSTBL POSTRD

Males
Micrurus tricolor (13) 7.31 3.19 1.73 6.12 1.89 3.08 7.42
(3.07) (0.60) (0.81) (1.45) (0.80) (0.64) (2.58)
(4-15) (2-4) (1-4) (4.5-10) (1-4) (2-4) (4-12)
Females
Micrurus tricolor (6) 6.75 3.08 1.75 5.83 1.50 2.92 5.75
(1.72) (0.49) (0.42) (0.68) (0.55) (0.20) (2.09)
(5-8) (2.5-4) (1-2) ( 5-6.5) (1-2) (2.5-3) (3-8)

in 1844. From the type series six speci-
mens are missing including the specimen
from from Corrientes and Misiones, which
apparently was represented by two indi-
viduals. Only two syntypes are left at the
MNHN herpetological collection, MNHN
578 and MNHN 854 (Roux-Esteve, 1983).
The description of the specimen(s) from
Corrientes and Misiones (Dumeril et al.,
1854) may have made Schmidt (1936) des-
ignate this region as the type-locality of M.
frontalis. Amaral (1944) reported two
specimens as cotypes (= syntypes) from
Corrientes and Misiones. We do not know
if these authors (Amaral, 1944; Schmidt,
1936) ever examined the specimens or
based their comments solely on the origi-
nal description where it clearly mentions
more than one specimen from this region
("Dans les exemplaires de Corrientes .. ";
Dumeril et al., 1854). The other possibility
is that at that time (1930-1940) the syn-
types were already missing, leaving the
only two reported in our paper and Amaral
(1944) interpreted it as if they were the
specimens from the type-locality designat-
ed by Schmidt (1936).
Oddly, Dumeril et al. (1854) also men-
tions Claussen's and Balperthuis' speci-
mens as if they were more than one which
was also noted by Schreve (1953) who
tried to assign Claussen's specimens (two)
as the only ones that really represented M.
frontalis. Shreve (1953) also noted the dis-
crepancy between specimens and type-lo-
calities and suggested Brazil as the type-
locality for M. frontalis which was ignored
by most authors including the latest revi-
sions (Roze, 1994, 1996). At that time P.
E. Vanzolini (1953) pointed out that
Claussen collected mostly around the town
of Lagoa Santa (Minas Gerais) when in
Brazil and this is possibly the real type-
locality for this taxon (see Shreve, 1953).
Claussen was a well known collector who
used to sell his collections of rocks, fossils,
and animals to European museums (see
Boulenger, 1896; Glinther, 1858; Lyddek-
ker, 1890; Stangerup, 1984).
In the original paper (Dumeril et al.,

TABLE 3.-Eigenvalues, proportion of total variation explained by each of the eight canonical vectors, and
character "loading" (= total-sample standardized canonical coefficients) on the first two canonical vectors.
Abbreviations are: LSVL = log-transformed snout-vent length; SVENSC = square-root transformed ventral
scale count (S = the same transformation in the remaining six variables); SCASC = subcaudal scale count;
STRIAD = total number of triads; SANTRD = anterior red ring; SANTBL = anterior black ring; SMIDBL
= middle black ring; and SANTWH = anterior white ring.

Vectors Eigenvalue Proportion Cumulative Character Can 1 Can 2

1 4.585 0.802 0.802 LSVL -0.0683 0.1490

2 0.804 0.141 0.943 SVENSC 0.4517 0.8088
3 0.194 0.034 0.977 SCASC 0.2053 0.0850
4 0.067 0.012 0.989 STRIAD -1.2696 -0.3193
5 0.048 0.008 0.996 SANTRD -0.0224 0.2834
6 0.014 0.003 0.998 SANTBL -0.0382 -0.2203
7 0.006 0.001 0.999 SMIDBL 0.9330 -1.1161
8 0.002 0.000 1.000 SANTWH 0.0602 0.2927
152 152 HERPETOLOGICAL MONOGRAPHS 13
[No.~~~~~~~~~~~
[No.13--

3 -

2 -

CM

1 -
.(

0_
0 -
o
0.
CO

-1 -

-2 -
I I I I I I

-2 0 2 4 6 8
Canonicalvariable1
FIG. 2.-Distribution of sample means along the two axes represented by canonical variables 1 (STRIAD
and SMIDBL) and 2 (SMIDBL and SVENSC). Abbreviations are defined as follows: circles = M. frontalis;
squares = M. altirostris; triangles = M. baliocoryphus; inverted triangles = M. brasiliensis; diamonds = M.
diana; hexagons = M. pyrrhocryptus; and circles + = M. tricolor. Solid = male, open = female.

1854) the description of the three speci-
mens is vague and the specimen(s) from
Corrientes and Misiones could be either
M. altirostris or M. baliocoryphus, as these
localities are in the Argentine Mesopota-
mia and there are no records of M. fron-
talis in this region. Since these specimens
are missing it is difficult to draw any ac-
curate conclusions based solely on the
original description.
The confusion of the description of
these very different specimens can be fur-
ther highlighted by other comparisons by
the authors that suggested some similari-
ties of Elaps frontalis with Elaps marcgra-
vii (Dumeril et al., 1854). The authors ap-
parently based their comments on a color
plate by Wied-Neuwied (1820) but in fact,
Elaps marcgravii is the type-specimen for
Micrurus ibiboboca (see also Merrem,
1820), and they might have made some
comparisons with the MNHN 578 (M)
syntype (= M. isozonus), which also has
long triad rings and a mostly red head.
Only recently Roze (1996) acknowledged
the syntype situation at MNHN but did
not mention or comment on the missing
specimens or the status of the MNHN 578
specimen. He only suggested the assign-
ment of the MNHN 854 specimen as the
lectotype, which was already obvious
(Roux-Esteve, 1983) and mentioned by
Jorge da Silva (1995).
Since the specimen(s) from Corrientes
and Misiones is/are missing and the de-
scription is clearly not of a Micrurus fron-
talis, we suggest changing the type-locality
to Lagoa Santa, Minas Gerais (Brazil) and
using the specimen MNHN 854 (M) as
the lectotype (as proposed by Roze, 1996).
To accept the current type-locality with a
missing specimen can lead to erroneous
1999]
1999] HERPETOLOGICAL MONOGRAPHS 153

A A B

'\-'

C D

ms

FIG. 3.-Morphological characteristics of the Mi-

crurusfrontalis lectotype (MNHN 854 M). A. Detail
of the head pattern; B. Detail of the first /3 of the FIG. 4.-Variation of head pattern in Micrurus
body (gray shading - red); C. Photograph of the frontalis. A. IB 20542 (M)-Brasflia, DF; B. IB
specimen. 48951 (M)-Niquelandia, GO; C. CEPB 2363
(M)-Pontalina, GO; D. IB 53331 (M)-Itutinga,
MG.

interpretations regarding the taxonomic

position of M. altirostris or M. baliocory- this unambiguously distinguishes the M.
phus (see below). frontalis pattern from the patterns of other
taxa. Also, the middle black ring in some
Diagnosis other taxa (M. baliocoryphus, M. pyrrho-
Micrurus frontalis is a triadal coralsnake cryptus, and M. tricolor) is much longer
with black snout with white bordered compared to M. frontalis. The white and
scales, including the rostral, internasal, the red rings are usually tipped with black
prefrontals, nasals, preoculars and anterior but this is not a fixed character for this
3-4 supralabials (Figs. 3-6). Laterally, the taxon. Triads range from 10 to 15 (Table
black color usually reaches the postoculars. 1).
The top of the head is black including the
frontal, supraoculars and parietals, never Redescription of the Holotype
reaching the temporals. Inferiorly, the Of the two remaining syntypes, speci-
mental, first 3 infralabials and sometimes men MNHN 578 (M) from Venezuela is
the anterior genials are white. These scales in fact a Micrurus isozonus, identified by
may present irregular black markings. The Hoge in 1978 (Roux-Esteve, 1983) and ex-
remaining gular region is red (mostly im- amined by one of us (NJS). It has 208 ven-
maculate). The first triad is usually sepa- tral and 28 subcaudal scales and 11 (10
rated from the parietals by 2 to 7 scales. body + 1 tail) black triads. The overall
The triad pattern can either be of equally length is 782.1 mm, with a SVL of 726.7
symmetrical black and white rings or have mm and a TL of 55.4 mm. The TL/SVL
a slightly longer middle black ring (Table ratio is 0.076. In the original paper, Du-
2). Whenever the middle black ring is lon- meril et al. (1854) even mentioned some
ger than the external ones the white rings peculiarities of this specimen related to
are longer than the external black rings; head coloration that are typical of M. iso-
154 HERPETOLOGICAL MONOGRAPHS
FIG. 5.-Detail of the hemipenis of Micrurusfrontalis (IB 53331-left
view. Total length of the hemipenis = 23.1 mm.
FIG. 6.-Detail of the skull of Micrurus frontalis
(CEPB 0614). A. Dorsal view; B. Lateral view. Total
length of the skull = 28.3 mm.
[No. 13
organ). A. Sulcate view; B. Asulcate
zonus, but at that time they were probably
interpreted as morphological variation
within M. frontalis.
The M. frontalis specimen from Brazil
(MNHN 854M) was described by Dumeril
et al. (1854) as having completely black
frontal and parietal scales, and the first oc-
cipital scales also black and fused to the
first triad. The description (MNHN 854
M) is as follows: rostral wider than high;
prefrontals slightly longer than the inter-
nasals. The frontal is as long as its distance
from the snout and the parietals are short-
er than their distance from the snout.
There are 1 + 1 temporals; 7 + 7 suprala-
bials; 7 + 7 infralabials; 0 + 2 oculars. It
has 235 ventral and 25 subcaudal scales.
The snout is pale with dark markings on
the intemasals and prefrontals. The frontal
is almost completely black with a small
white margin anteriorly. The supraoculars
1999]
1999] HERPETOLOGICALMONOGRAPHS
are mostly pale red with an almost iden-
tical black marking on both scales at their
distal margin. The parietals are mostly
black with irregular margins anteriorly and
a wide semitriangular pale red spot on the
posterior part of the scales. The external
ring of the first triad is almost completely
fused with the black color of the parietal
scales. Inferiorly, the chin is immaculate
(probably red) with the exception of the
mental and first infralabials that might be
white in the live specimen. There are 12
(11 + 1) black triads with the middle black
ring longer than the external ones. All of
the middle black rings are irregular in
shape. The white rings are shorter than the
external black rings at the vertebral line.
All red and white rings are black-tipped,
with the black being stronger on the white
rings (Fig. 3). The overall length is 528.7
mm with a head length of 21.4 mm, a SVL
of 498.4 mm and a TL of 30.3 mm. The
tail length/SVL ratio is 0.061.
The specimen has some reduction of
black color on the head (parietal scales)
and irregular middle black rings on the tri-
ads. However, there are no black markings
on the chin, which is diagnostic of M. fron-
talis; all the other taxa of the complex pos-
sess mild to heavy darkening of the chin.
In the sample that we examined this is not
the most frequent character (apparently
not fixed) but even when present it is very
reduced (Fig. 4).
Variation
Males showed a range of 212 to 238
ventral scales, 19 to 27 subcaudal scales,
and 10 to 15 triads (n = 494; Table 1).
Females showed a range of 213 to 234
ventral scales, 15 to 25 subcaudal scales,
and 11 to 14 triads (n = 132; Table 1).
Thirteen specimens (2.1%) examined from
throughout the range revealed some mor-
phological variation, especially regarding
the head color pattern. Figure 4 shows
some of the head pattern variation and
Fig. 1 the most common triad pattern in
M. frontalis. In 3% of our sample we
found a condition in which the anterior
ring of the first triad was fused with the
posterior end of the parietals. The snout
pattern varies clinally from south to north
155
in the states of Goias and Minas Gerais
(with the specimens in the northern range
having a lighter snout), but elsewhere in
the range, this variation appears to be ran-
dom.
Hemipenial Morphology
The hemipenis is weakly bilobed, with
short lobes covered by uniform size spines
(Fig. 5). The everted organ is seven sub-
caudals long with lobes formed at the
sixth/seventh subcaudals level. The hemi-
penis of the IB 53331 specimen is 23.1
mm long, representing 1.7% of the SVL
(SVL = 1335 mm) and 27.8% of the TL
(TL = 83.1 mm). The sulcus spermaticus
bifurcates at about the fifth/sixth subcau-
dals level, extending centripetally to the
distal tip of the lobes. The intrasulcar sur-
face is smooth. The semicapitate condition
is evident in the asulcate surface by a line
at the second (sometimes third) subcaudal
level. Below this line to the base of the
hemipenis the surface is smooth. Above it
there are larger spines of equal size ex-
tending distally to the tip of the lobes. The
sulcate surface is also covered by large
spines above the level of the second or
third subcaudal scales. The basal region to
this level the hemipenis is mostly naked
with 20-30 large spines of equal size (Fig.
5).
Cranial Osteology
In M. frontalis the dorsomedial process-
es of the prefrontal bones are strongly de-
veloped and may or may not contact at
midline. At midline the dorsomedial pro-
cesses are narrow. An anteromedial pro-
cess of the frontals may be present and
separates the dorsomedial processes of the
prefrontals. The braincase (parietals) is
pointed at the posterior end. The suture
prefrontals + frontals is anterior to the
maxilla + pterygoid suture. There are 8 to
10 palatine, 3 to 4 pterygoid, and 10 to 12
dentary teeth (Fig. 6).
Geographic Distribution
This taxon has the most widespread dis-
tribution of all members of the M. frontalis
complex. It ranges from the Cerrado for-
mations of Central Brazil (including Sao
156 HERPETOLOGICAL MONOGRAPHS
FIG. 7.-Distribution of Micrurus frontalis (black
dots), M. brasiliensis (black diamonds), and M. diana
(open squares). Stars represent the type-localities of:
1. M. frontalis (Lagoa Santa, Minas Gerais-Brazil);
2. M. brasiliensis (Barreiras, Bahia-Brazil); 3. M. di-
ana (Serranfa de Santiago, Santa Cruz-Bolivia).
Scale represents 200 km.
Paulo and Minas Gerais) west to the bor-
der of Bolivia and the Brazilian states of
Mato Grosso and Mato Grosso do Sul. To
the east it apparently reaches the Atlantic
coast only in the state of Espirito Santo
due to the Serra do Mar mountain range
to the south. There is no evidence at this
point to the presence of this taxon in Par-
ana. To the S-SW it reaches Paraguay
where it is restricted possibly by the Rio
Parana and Rio Paraguay (Fig. 7). In dif-
ferent parts of its range it is sympatric with
M. tricolor, M. lemniscatus, M. altirostris,
and M. baliocoryphus.
Micrurus altirostris (Cope)
Elaps altirostris Cope 1859:345
Holotype: ANSP 6857 (M)
Type-locality: South America.
Elaps heterochilus Mocquard 1887:39
(Type-locality: Brazil)
[No.
[No. 13
Elaps frontalis Boulenger 1896:427
Micrurus frontalis altirostris Schmidt
1936:199
Micrurus lemniscatus multicinctus (par-
tim) Amaral 1944:91 (Type-locality:
Teixeira Soares-IB 8877 F).
Micrurus frontalis altirostris Barrio and
Miranda 1966:875
Micrurus frontalis altirostris Roze 1967:25
Micrurus lemniscatus multicinctus (par-
tim) Roze 1967:25
Micrurus frontalis altirostris Abalos and
Mischis 1975:74
Micrurus frontalis altirostris Achaval, Mel-
garejo, and Meneguel 1976:4
Micrurus frontalis altirostris Achaval 1976:
28
Micrurus frontalis altirostris Hoge and
Romano 1979:395
Micrurus frontalis altirostris Roze 1983:
323
Micrurus frontalis multicinctus (partim)
Roze 1983:325
Micrurus frontalis altirostris Esteso 1985:
41
Micrurus frontalis altirostris Golay 1985:
34
Micrurus frontalis multicinctus (partim)
Golay 1985:34
Micrurus frontalis altirostris Scrocchi
1990:350
Micrurus frontalis multicinctus (partim)
Scrocchi 1990:367
Micrurus frontalis altirostris Roze 1994:
179
Micrurus frontalis multicinctus Roze 1994:
179
Micrurus frontalis altirostris Roze 1996:
173
Micrurus frontalis multicinctus Roze 1996:
175
Micrurus frontalis altirostris Achaval 1997:
515
Micrurus frontalis multicinctus Achaval
1997:518
This is probably the most problematic
species in terms of previous literature and
taxonomic treatments. One of us (NJS) in-
vested a great deal of time examining the
holotype and the available specimens of
major collections in Brazil, Argentina, Par-
aguay, and Uruguay. Since Cope's descrip-
1999] HERPETOLOGICAL MONOGRAPHS 157

tion (1859), the holotype appears not to A

have been examined by most later authors,
who might have avoided taxonomic mis-
takes in describing new forms or suggest-
ing taxonomic arrangements (Amaral,
1944; Mocquard, 1887; Roze, 1967, 1970,
1983, 1994, 1996; Schmidt, 1936; Scroc-
chi, 1990).
The literature problems first appeared
with Schmidt (1936) and Amaral (1944).
The first author offered a preliminary ac-
count of South American coralsnakes
based on specimens available in US and C
European museums at that time, and sug
gested a restricted range for M. altirostri
in Uruguay, with M. frontalis ranging fro
Sao Paulo to Rio Grande do Sul (Brazil)
Amaral (1944) compared Schmidt's pape
with his data (obtained from a much large
sample size at IB) and contested the pro
posed geographical distribution. Th
whole discussion was about the difference
between M. frontalis and M. altirostris
Both authors were working with the entire FIG. 8.-Morphological characte-isti"o Mi-
cnintls altirostris holotype (ANSP 6857 M). A. D)etail
range of scale counts (including extreme of the head pattern; B. D)etail of the first 1/3 of the
outliers), and if this is considered as the bodv (gray shading = red); C. Photograph of the
only criterion, then all of the forms includ- specimen.
ed in the M. frontalis complex overlap (Ta-
ble 1). With regards to the geographical
range, Amaral (1944) had a much better sample of 46 specimens, and also suggest-
representation than Schmidt (1936) and ed a distribution of the latter from Uru-
his observations were closer to the real dis- guay to Sao Paulo (Brazil).
tribution. However, Vanzolini (1953) The position of M. lemniscatus multi-
pointed out that M. altirostris was never cinctus was interpreted by Roze (1967,
found in Sao Paulo; it is confined to south- 1970) and Scrocchi (1990) as an intergrade
ernmost Brazil. between M. f frontalis and M. f altirostris
Amaral (1944) also rearranged M. fron- and later (Achaval, 1997; Roze, 1983,
talis and M. ibiboboca into M. lemniscatus, 1994, 1996) as a subspecies of M. frontalis.
and raised some interesting questions
about the latter two taxa that we will ad- Diagnosis
dress under M. baliocoryphus. In this Micrurus altirostris is a triadal coral-
same paper, he described M. lemniscatus snake with a black snout and most scales
multicinctus based on two specimens from bordered by white. The frontal is usually
Teixeira Soares (IB 8877 F-holotype) and completely black, but the overall color of
Iratl (IB 7874 F-cotype), Parana, Brazil, the head is red, which covers most of the
with an extremely poor characterization of parietals, temporals, and supralabials pos-
the form. The only diagnostic feature was terior from the eyes. The posterior border
the higher number of triads in both spec- of the parietals is black (Figs. 8-11). There
imens. is a tendency toward increased melanism
Shreve (1953) suggested that "M. lem- of the red color of the head and triad
niscatus multicinctus" might be simply a rings. Inferiorly, the gular region is red
product of hybridization between M. f. with irregular but intense black markings.
frontalis and M. f. altirostris, based on a All supralabials and infralabials possess ir-
158 HERPETOLOGICAL. MONOGRAPHS
HERPETOLOGICALT [No 13
[No.

A A B

._waBil
_l^^^UEj^E

C D

_fli

FIG. 9.-Morphological characteristics of the Mi-

crurusfrontalis multicinctus holotype (IB 8877 F). A.
Detail of the head pattern; B. Detail of the first ? of
the body (gray shading = red); C. Photograph of the
specimen.
A B
FIG. 11.-Variation of head pattern in Micrurus
altirostris. A. IB 11122 (M)-Cruz Alta, Rio Grande
do Sul; B. CHINM 3633 (M)-Federaci6n, Corrien-
tes; C. CHINM 1605 (M)-Posadas, Misiones; D.
CHINM 1802 (M)-Puerto Paranaf, Misiones.

m_ regular black markings. The triads on this

form are characterized by subequal black
rings (3-4 dorsal scales) with shorter white
rings (1-2 dorsal scales) (see Table 1). The
middle black ring sometimes can be a little
longer than the external ones (4-5 dorsal
scales). The first triad begins always at 1
or 2 dorsal scales from the parietals. All
red and white rings are black-tipped. The
triad total length is the shortest among all
C D
taxa of this complex. Triads range from 13
to 18 (Table 2).
^if
Redescription of the Holotype
The holotype (ANSP 6857 M) is as fol-
lows: rostral wider than high; prefrontals
slightly longer than the internasals. The
frontal is as long as its distance from the
snout and the parietals are a little shorter
than their distance from the snout. There
are 1 + 1 temporals; 7 + 7 supralabials; 7
FIG. 10.-Variation of head pattern in Micrurus + 7 infralabials; 0 + 2 oculars. The spec-
altirostris. I. A MHNCI 4208 (F)-Irati, Parana; B.
MHNCI 4848 (F)-Campo imen has 194 ventrals and 15 subcaudals
Largo, Parana; C.
MHNCI 4884 (M)-Cascavel, Parana; D. MHNCI (the tail is damaged).
3146 (M)-Ararangua, Santa Catarina. The snout is black with few small white
1999] HERPETOLOGICAL MONOGRAPHS
HERPETOLOGICAL MONOGRAPHS 159
159

spots on the internasals and anterior part tion in black in confined to most or all of
of the frontal and right supraocular (Fig. the parietal scales only, while the pattern
8). The parietals are almost completely for M. altirostris is characterized by a re-
pale red with some black marking on the duction on the external borders of those
anterior and lateral borders with two black scales and sometimes the frontal and su-
spots posteriorly, close to the suture be- praoculars. Also, M. altirostris has a char-
tween these scales. There is a distance of acteristic feature of black marking on the
one red scale behind the parietals separat- chin. In M. frontalis there is very little or
ing them from the first triad. Inferiorly, the no black marking on the chin. The differ-
head is black with small white markings on ences between M. altirostris and the other
the borders of the infralabials and chin taxa can be further highlighted in Tables 1
shields. There are 14 (13 + 1) black body and 2. Extreme body lengths (SVL + TL)
triads with the middle black ring a little were 187 and 1310 mm.
longer than the external ones. The white The position of M. f. multicinctus is not
rings (black tipped) are shorter than the as an intergrade between M. f. frontalis
external black rings and red (black tipped) and M. f. altirostris, as proposed by Scroc-
are a little longer than the middle black chi (1990) in what he called "synonymi-
ring. The red rings are immaculate (Fig. zation of extra-Argentinian forms", or sub-
8). The overall length of the holotype is species of M. frontalis (Roze, 1994, 1996).
674 mm with a head length of 21.3 mm, a The range of M. frontalis apparently does
SVL of 639.5 mm, and a TL of 34.5 mm. not reach the northern region of Parana.
The TL/SVL ratio is 0.054. In fact, there is no specimen available for
The holotype of M. f. multicinctus (IB the entire state of Parana with any com-
8877 F) has 214 ventral and 25 subcaudal bination of the characters found in M.
scales with 19 (18 + 1) triads. The snout frontalis, and from all of the examined ma-
is pale with the rostral black, and the in- terial, this population (PR) is the most
ternasals and prefrontals are posteriorly similar to Cope's holotype. The character-
bordered with black. The frontal and su- istic of high triad number is typical of M.
praoculars are almost completely black, altirostris and M. f. multicinctus just rep-
and the parietals mostly red with anterior resents one of the upper outliers; for ex-
and porterior tips black. Inferiorly, the ample, one specimen from Sao Jose dos
mental and first and second infralabials are Pinhais (PR) has 21 (20 + 1) triads, but
completely black with the posterior part of other specimens from this same location
the fourth infralabial also black. There are show a range from 14 to 16 triads, with
black markings in the posterior part of the similar results for the additional specimens
chin shields. The black triad rings are sub- from Teixeira Soares and Irati. All of the
equal with shorter white rings (Fig. 9). other features of head, triad pattern, and
The similarities of this holotype with the scale counts are well within the range of
M. altirostris holotype (Fig. 8) are very values reported for M. altirostris. As pre-
clear. viously mentioned, taxonomic errors occur
when descriptions of variation are made
Variation without comparing a specimen of interest
Males showed a range from 199 to 230 to other specimens from the same locality
ventral scales, 15 to 25 subcaudal scales, and total range. Based on these findings
and 13 to 18 triads (n = 765; Table 1). we place M. f multicinctus as a junior syn-
Females showed a range from 199 to 220 onym of M. altirostris.
ventral scales, 14 to 23 subcaudal scales, The morphological features described
and' 14 to 18 triads (n = 319; Table 1). It above for this species remain the same for
is possible to separate M. altirostris from the state of Santa Catarina and most of Rio
M. frontalis by the higher number of ven- Grande do Sul. However, we found that at
tral scales and lower triad numbers in both least two combinations of head and triad
sexes in M. frontalis. Also, the head pat- patterns occur in Rio Grande do Sul. One
tern of M. frontalis is unique; the reduc- is the typical head pattern described for
160 HERPETOLOGICAL MONOGRAPHS
M. altirostris with subequal black triad
rings. In the other the head is mostly black
as in M. frontalis, with very little reduction
on the posterior border of the parietal
scales, with a black middle triad ring twice
as long as the external ones, and with re-
duced black coloration of the chin (see
Figs. 10, 11). Both forms are sympatric in
Alegrete, Cacequf, Carazinho, Catufpe,
Cruz Alta, Ijuf, Montenegro and Rosario
do Sul (Rio Grande do Sul). Roze (in litt.,
1992) also suggested a third form with
head pattern as in M. frontalis with black
chin as well. With regard to this third
form, we found that most of the specimens
from Uruguay and in the eastern basin of
the Rio Uruguay are melanistic to some
degree, and this is not a rare characteristic
in other species of Micrurus (Jorge da Sil-
va, Jr., 1993). The presence of these non-
typical M. altirostris patterns in Rio Gran-
de do Sul suggests that this taxon may
include other forms, but at this time we
retain the present taxonomic arrangement
due to small sample size. All the speci-
mens from Uruguay show intense mela-
nism of the red and white color of the
head and triads. However, the character of
the black reduction of the posterior pari-
etals is fixed.
Scrocchi (1990) presented data that
show three records of M. altirostris in En-
tre Rios (Argentina), and in analyzing the
same specimens we found that they do
represent M. altirostris even though the
characters are not easily distinguished. At
this point we accept them as M. altirostris
based on the lack of the transverse white
band on the head, typical of M. baliocory-
phus, together with a relatively shorter
middle black ring on the triads (typical of
M. altirostris). Additional specimens from
Concordia (CHINM 3379 M, CHINM
3571 M) and Santa Elena (MACN 1117
M) are M. altirostris, and this support
Scrocchi's (1990) interpretation of the dis-
tribution of this taxon as confined to the
Argentine Mesopotamia.
The only records for M. altirostris in the
Province of Corrientes are CHINM 2005
(M) from Santo Tome, CHINM 3537 (M)
from Torrent, and a MACN specimen
without a number, from Ituzaingo. In
[No 12
[No. 13
A B
FIG. 12.-Detail of the hemipenis of Micrurus al-
tirostris (MCP 5035-right organ). A. Sulcate view;
B. Asulcate view. Total length of the hemipenis =
12.5 mm.
July-September of 1994 the reservoir of
the Yacireta hydroelectric power plant was
filled on the Rio Parana between Argen-
tina and Paraguay, close to Ituzaingo.
From this site we received 3 live speci-
mens of M. altirostris that confirmed this
distribution (IB 55595, IB 55596, and IB
55597). The other records that Scrocchi
(1990) presents as intermediate forms in
Corrientes (e.g., General Paz) we recog-
nize as M. baliocoryphus. The records of
M. altirostris from Corrientes form a semi-
continuous line on the eastern side of the
Province and might be a natural connec-
tion with the population of Entre Rios.
Hemipenial Morphology
The hemipenis is weakly bilobed, with
short lobes covered by uniform size spines
(Fig. 12). The everted organ is five to six
subcaudals long with lobes formed at the
fourth or fifth subcaudal level. The hemi-
penis of the MCP 5035 specimen is 12.5
mm long, representing 1.7% of the SVL
(SVL = 735 mm) and 29.1% of the TL
(TL = 43.0 mm). The sulcus spermaticus
bifurcates at almost the same level of the
lobes, extending centripetally to the distal
tip of the very discrete lobes. The intra-
sulcar surface is smooth. The semicapitate
199
19991 EREOOGCLMOORPH
HERPETOLOGICAL MONOGRAPHS
FIG. 13.-Detail of the skull of Micrurus altirostris
(CEPBc 5911). A. Dorsal view; B. Lateral view. Total
length of the skull = 17.4 mm.
line is evident on the asulcate surface at
the level of the second subcaudal. From
this line above the spines are large and un-
iform. Below the line the spines are fewer
and decrease in size towards the base of
the hemipenis. The sulcate surface is com-
pletely covered by large spines of equal
size. From the base of the hemipenis up
to the level of the second subcaudal there
are smaller spines of extremely variable
number. The base of the hemipenis is na-
ked (Fig. 12).
Cranial Osteology
In M. altirostris the dorsomedial pro-
cesses of the prefrontal bones are strongly
developed and do not contact at midline.
They are united by connective tissue. At
midline the dorsomedial processes are nar-
row. An anteromedial process of the fron-
tals may be present separating the dorso-
medial processes. The braincase (parietals)
is rounded at the posterior end. The su-
ture prefrontals + frontals is posterior to
the maxilla + pterygoid suture. There are
8 to 10 palatine, 3 to 4 pterygoid, and 10
to 11 dentary teeth (Fig. 13).
Geographic Distribution
This taxon ranges from the Brazilian
state of Parana south to Uruguay, and the
161
6
FIG. 14.-Distribution of Micrurus altirostris
(black dots) and M. baliocoryphus (open triangles).
Stars represents the type-localities of: 1. M. lemni-
scatus multicinctus (Teixeira Soares, Parana-Brazil);
2. M. baliocoryphus (Villa Federal, Entre Rios-Ar-
gentina). Known records of M. lemniscatus for South-
ern Brazil, Argentina and Paraguay are represented
by black stars. Scale represents 100 km.
Provinces of Misiones, Corrientes, and
northern Entre Rios, in Argentina and
eastern Paraguay (Fig. 14). This taxon is
sympatric with M. baliocoryphus in Mis-
siones and Entre Rios (Argentina), M. lem-
niscatus in Brazil and Paraguay, possibly
with M. decoratus in Parana (Brazil), and
with M. frontalis in Paraguay. There are no
records available of the distribution of this
taxon north into the Brazilian state of Sao
Paulo which agrees with Vanzolini's obser-
vations (1953).
The distribution of M. altirostris in Cor-
rientes and Entre Rios suggests that these
rivers form natural barries to dispersal
(Fig. 14). In this case, M. altirostris seems
to have invaded the Argentine Mesopota-
mia and Paraguay from Brazil through the
northern border of Misiones. In this re-
gion the Rio Parana is the western barrier
and the Rio Uruguay appears not to re-
162 HERPETOLOGICAL MONOGRAPHS [No. 13
[No.

strict dispersal. If the population of Entre certainly did not see Cope's holotype, as it
Rios originated in Uruguay and crossed was never mentioned in their paper, so
the Rio Uruguay, we would expect a much these authors missed all of the similarities
larger representation of this taxon in this between these two forms. Hoge and Ro-
region. Also, the populations of M. altiros- mano (1979) did not deal with the prob-
tris from Uruguay show intense melanism, lem and just recognized it as M. f balio-
and this character is rare in the specimens coryphus without any further comments.
from Argentina. All of the specimens from Roze (1983, 1994, 1996) synonymized M.
Uruguay are otherwise typical of M. alti- f. mesopotamicus with M. f baliocoryphus,
rostris. The localities of museum vouchers and also mentioned that the form de-
in Uruguay have been augmented with the scribed by Barrio and Miranda (1966) was
data from Achaval (1997). the same as Elaps baliocoryphus (Cope,
Micrurus baliocoryphus (Cope) 1862). This author (Roze, 1983) was also
aware of a possible type-locality error in
Elaps baliocoryphus Cope 1862:346
Cope's description, suggesting Villa Fed-
Holotype: ANSP 6842 (M) eral, Entre Rios, Argentina as the type-lo-
Type-locality: Buenos Aires, Argentina. cality. At the time (1983) Roze not only
Corrected and restricted to Villa Federal, had seen Cope's holotype but also the ma-
Entre Rios, Argentina by Roze (1983:324) terial from Barrio and Miranda (1966).
Micrurus frontalis mesopotamicus Barrio The paratype that these authors (Barrio
and Miranda 1966:872 (Type-locality: and Miranda, 1966) used to illustrate their
Villa Federal, Entre Rios, Argentina- paper (CHINM 1627 F- Charaji, Entre
CHINM 1823 M). Rios, Argentina) is currently part of the
Micrurus frontalis mesopotamicus Abalos AMNH herpetological collection (=
and Mischis 1975:74 AMNH 99985) also mentioned by Roze
Micrurus frontalis baliocoryphus Hoge (1996). The correction of the type-locality
and Romano 1979:396 of M. f baliocoryphus by Roze (1983) was
Micrurus frontalis baliocoryphus Roze based on the previously accepted restrict-
1983:323 ed geographic distribution of this form in
Micrurus frontalis mesopotamicus Esteso the mesopotamic region of Argentina (see
1985:41 below).
Micrurus frontalis baliocoryphus Golay Scrocchi (1990) synonymized M. f. bal-
1985:34 iocoryphus with M. f. mesopotamicus
Micrurus frontalis mesopotamicus Scroc- based solely on Roze's (1983) and Barrio
chi 1990:353 and Miranda's (1966) papers, without an
Micrurus frontalis baliocoryphus Roze analysis of Cope's holotype, or any refer-
1994:179 ence to the original description. In the
Micrurus frontalis baliocoryphus Roze original description, Cope (1862) clearly
1996:174 mentions the triad pattern (middle ring
two times longer than the external ones)
Since its original description this taxon and the head coloration.
has been the subject of extensive discus-
sion, but much of the taxonomic confusion Diagnosis
could have been avoided if some of the Micrurus baliocoryphus is a triadal cor-
authors had analyzed the type specimen. alsnake with a white snout with most of
The most important literature is the orig- the scales bordered by black, including the
inal description of this taxon by Cope first 3-4 supralabials. Sometimes the pre-
(1862), and the revisions of some authors frontals are completely white with a few
(Barrio and Miranda, 1966; Roze, 1983, little black markings. The head is black
1994, 1996; Scrocchi, 1990). with (the vast majority of our sample) or
Barrio and Miranda's (1966) description without a white band between the parie-
of M. f mesopotamicus agrees well with tals and frontal and supraoculars. The re-
Cope's Elaps baliocoryphus and they most mainder of the head is red, including the
 _
1999] HERPETOLOGICAL MONOGRAPHS
A A
As
.
IB
i3XE
_ _
_ _
3Efi
C
zS-_Z
3_
FIG. 15.-Morphological characteristics of the Mi-
crurus baliocoryphus holotype (ANSP 6842 M). A.
Detail of the head pattern; B. Detail of the first % of
the body (gray shading = red); C. Photograph of the
specimen.
posterior 3-4 infralabials with all scales
black-tipped (Figs. 15-17). Inferiorly, the
chin is white, ranging from the mental and
first 2 infralabials to the anterior genials
and first 3 infralabials. All the scales have
irregular black markings with a strong ten-
dency to melanism. The posterior part is
red rarely with black markings. The triads
have a fixed character of the middle black
ring at least twice as long as the external
ones and immaculate white rings. The red
rings are always black-tipped. Triads range
from 9 to 16.
Redescription of the Holotype
The description of the holotype (ANSP
6842 M) is as follows: rostral wider than
high; prefrontals twice as long as the in-
ternasals. The frontal is as long as its dis-
tance from the snout, and the parietals are
a little longer than their distance from the
snout. There are 1 + 1 temporals; 7 + 7
supralabials; 7 + 7 infralabials; 0 + 2 oc-
ulars. It has 226 ventrals and 26 subcau-
163
_
_
.
c
- FIG. 16.-Morphological characteristics of the Mi-
crurus frontalis mesopotamicus holotype (CHINM
1823 M). A. Detail of the head pattern; B. Detail of
the first V3of the body (gray shading = red); C. Pho-
tograph of the alotype CHINM 1627 (F) = AMNH
99985.
dals. The scales of the snout are pale with
dark spots on the internasals and on the
third and fourth supralabials. The supra-
oculars are black. The frontal and parietals
are mostly black with a white band ante-
rior to the parietals and posterior to the
frontal and supraoculars. Inferiorly, the
head is pale with dark spots on the first
three supralabials and mentals. There are
15 (14 + 1) black body triads, and the
middle black ring two times longer than
the external ones. The white rings (im-
maculate) are equal and the same length
as the external black rings. The red rings
are about the same length of the triad and
all of the red scales are black tipped (Fig.
15). The overall length of the holotype is
785.4 mm with a head length of 20.6 mm,
a SVL of 739.7 mm, and a TL of 45.7 mm.
The TL/SVL ratio is 0.062.
Barrio and Miranda's (1966) holotype
(CHINM 1823 M) has the same head col-
oration, including the characteristic white
band on the head (between the parietals
and frontals and supraoculars scales), with
164 HERPETOLOGICAL MONOGRAPHS [No. 13

A B on triad number and pattern and scale

counts (see Figs. 1 and 2). Extreme body
49 lengths (SVL + TL) were 270 and 1449
mm.
In Ituzaing6 (Corrientes, Argentina) M.
baliocoryphus (CHC 126 M) is sympatric
with M. altirostris. In General Paz, the
specimens (CHINM 2870 M and CHINM
2871 M) mentioned by Scrocchi (1990) as
intermediary forms are in fact M. balio-
coryphus, and this has been corroborated
by an additional specimen (FML 1986 M).
C D The specimens from Saladas (Corrientes)
(CHINM 737 M and CHINM 738 M)
have the characteristic low triad number
(10 in both) and the triad pattern diagnos-
tic of M. baliocoryphus. In Argentina only
two specimens (CHINM 3053 M-Saladas,
Corrientes; and CHINM 1518 M-Posadas,
Misiones) lacked the typical head pattern
but still have the triad pattern and number
for M. baliocoryphus (Fig. 17). All speci-
FIG. 17.-Head pattern in Micr-urus baliocory- mens from Paraguay lacked the white
phus. A. CHINM 3053 (M)-Saladaw s, Corrientes; B. band on the head. The specimens from
CHINM 1518 (M)-Posadas, Misiories; C. CHINM
3382 (M)-Micrurus lemniscatus-
Corrientes and Misiones together with
siones; D. COBAP 058 (M)-Micrur us tricolor An- those from Paraguay have the scales of the
astacio, Mato Grosso do Sul. snout without the typical white bordering.
All the specimens from Paraguay are in-
terpreted as M. baliocoryphus in this pa-
220 ventrals and 24 subcauda ds. The spec- per, but we acknowledge that they might
imen has 14 (13 + 1) black body triads. represent an intermediate form.
The middle black ring is als;o two times Scrocchi (1990) also mentioned two in-
longer than the external ones ;, and is bor- termediate forms (CHINM 1473 M;
dered by immaculate white rings of equal CHINM 3382 M) from Posadas, Misiones.
length (Fig. 16). Undoubted Ily the same In fact these specimens are M. lemniscatus
taxon as Cope's. The problEem with the (Fig. 17) which were misidentified, as was
type-locality and the opinions of other au- specimen CHINM 2403 (M) (identified as
thors on the subject are not sufficient to M. f mesopotamicus), and a fourth speci-
support the changes proposec I by Scrocchi men (CHINM 3043 M) from Puerto Aza-
(1990). In this case, Cope's name (balio- ra. These are the first records of M. lem-
coryphus) has priority over BEarrio and Mi- niscatus for Argentina (Jorge da Silva and
randa's (mesopotamicus). ]Roze (1994, Silva, 1996), and in general pattern they
1996) recognized that but retained balio- resemble M. baliocoryphus, but the head
coryphus as a subspecies of A4. frontalis. coloration is unambiguously characteristic
of M. lemniscatus.
Variation
Males showed a range of 21 0 to 230 ven- Hemipenial Morphology
tral scales, 18 to 27 subcaudal scales, and 9 The hemipenis is weakly bilobed, with
to 16 triads (n = 81; Table 1). Females had short lobes covered by uniform size spines.
a range of 208 to 233 ventral scales, of 17 The everted organ is six subcaudals long
to 23 subcaudal scales, and 1]1 to 15 triads with lobes formed at the fifth/sixth subcau-
(n = 17; Table 1). M. baliocor yphus can be dals level. The hemipenis of the MZUSP
differentiated from the other species based 10809 specimen is 18.1 mm long, repre-
19991
-9991 HERPETOLOGICAL
HERPETOLOGICAL MONOGRAPHS
MONOGRAPHS
senting 1.7% of the SVL (SVL = 1078
mm) and 27.4% of the TL (TL = 66.0
mm). The sulcus spermaticus bifurcates at
the fifth subcaudal level extending centrip-
etally to the distal tip of the lobes. The
bifurcation of the sulcus spermaticus pre-
cedes by one subcaudal the bifurcation of
the lobes. The intrasulcar surface is
smooth. The semicapitate line is very evi-
dent on the asulcar surface at the level of
the second/third subcaudals. Above this
line the spines are large and uniform ex-
tending to the tip of the lobes. Below the
line to the base of the hemipenis the sur-
face is completely naked. At the sulcate
surface from the level of the third subcau-
dal to the tip of the lobes the spines are
large and uniform. From this level down
to the base of the hemipenis the spines are
smaller and irregularly distributed. The
base of the hemipenis is naked (Fig. 18).
Cranial Osteology
In M. baliocoryphus the dorsomedial
processes of the prefrontal bones are
strongly developed and do not contact at
midline. They are united by connective tis-
sue. At midline the dorsomedial processes
are wide. The anteromedial processes of
the frontals are absent. The braincase (pa-
rietals) is rounded at the posterior end.
The skull table is elongated and covers the
anterior 2/3 of the braincase. The suture
prefrontals + frontals is anterior to the
maxilla + pterygoid suture. There are 7 to
9 palatine, 3 to 4 pterygoid, and 8 to 10
dentary teeth (Fig. 19).
Geographic Distribution
This taxon is found in the Argentine
Mesopotamia, through the provinces of
Entre Rios, Corrientes, and Misiones be-
tween Rio Paranai and Rio Uruguay (Fig.
14). It is also present in SW Paraguay
along the Rio Paraguay from the provinces
of Neembucu through Central and Para-
guari to Presidente Hayes, and the adja-
cent Argentine province of Formosa (Fig.
13). This distribution corroborates an ex-
tra-mesopotamian range as suggested by
Campbell and Lamar (1989). There are no
records of this taxon in Brazil or Uruguay.
There are no records of M. lemniscatus in
165
165
the adjacent Brazilian states of Rio Grande
do Sul and Santa Catarina but they do oc-
cur in NW-W Paranai, southern Mato
Grosso do Sul and Paraguay (Fig. 14; data
from AMNH, IB, and MACN herpetolog-
ical collections). The distribution of this
taxon lends support to the hypothesis that
the Rio Uruguay and Rio Paranai act as a
partial natural barriers to M. baliocory-
phus.
Micrurus brasiliensis Roze
Micrurus frontalis brasiliensis Roze 1967:
25
Holotype: UMMZ 108880 (M)
Type-locality: Barreiras, Bahia, Brazil.
Paratypes: UMMZ 108881 (F)-Barreir-
as, Bahia; UMMZ 108878 (M)-Januario (=
Januaria), Minas Gerais; AMNH 90361
(M)-Santa Isabel (= Santa Isabel do Mor-
ro), Ilha do Bananal, Goias (now state of
Tocantins); MNRJ 2494 (M)-Barreira,
Bahia; MNRJ 2497 (F)-Manga, Minas Ge-
rais; MCZ 3298 (F)-Santa Cruz, Bahia (=
Santa Cruz de Cabralia).
Micrurus frontalis brasiliensis Hoge and
Romano 1979:396
Micrurus frontalis brasiliensis Golay 1985:
34
Micrurus frontalis brasiliensis Campbell
and Lamar 1989:115
Micrurus frontalis brasiliensis Roze 1994:
179
Micrurus frontalis brasiliensis Roze 1996:
174
Roze (1967) described this form based
on the pale snout with black bordered
scales (internasals and prefrontals) and
fewer subcaudal scales in females (16 to
18) when compared to M. frontalis. Since
then very little has been done to clarify the
status of this taxon, and the latest revisions
are not based on any better sample size
than the type series (Roze, 1994, 1996).
Diagnosis
Micrurus brasiliensis is a triadal coral-
snake with a white snout including the
anterior border of the frontal and supra-
ocular scales reaching 3-4 anterior su-
pralabials. All scales show some degree
166 HERPETOLOGICAL MONOGRAPHS [No.
[No. 13

A B
FIG. 18.-Detail of the hemipenis of Micrurus baliocoryphus (MZUSP 10809-left organ). A. Sulcate view;
B. Asulcate view. Total length of the hemipenis = 18.1 mm.

of black markings. The head is mostly 2 infralabials) sometimes with irregular

red with the posterior part of the frontal
and supraoculars black (Figs. 20-21).
The anterior part and along the medial
suture of the parietals is black with vary-
ing reduction of the black posteriorly.
This is a fixed character of this taxon. All
red scales are mostly immaculate. Infe-
riorly, the chin is white (mental and first
black markings. The remainder is red
with or without black markings. The tri-
ads are also unique. The white rings are
of the same length or slightly longer than
any of the black rings (Table 2). The red
rings are usually immaculate and the
white rings are only mildly black-tipped.
Triads range from 11 to 14 (Table 1).
1999] HERPETOLOGICAL MONOGRAPHS
FIG. 19. Detail of the skull of Micrurus baliocor-
yphus (MZUSP 10820). A. Dorsal view; B. Lateral
view. Total length of the skull = 20.4 mm.
A
B
C 7 infralabials; 0 + 2 oculars. It has 228
FIG. 20.-Morphological characteristics of the Mi-
crurus brasiliensis holotype (UMMZ 108880 M). A.
Detail of the head pattern; B. Detail of the first ? of
the body (gray shading = red); C. Photograph of the
specimen.
167
A B
C D
FIG. 21.-Head pattern in Micrurus brasiliensis. A.
CEPB 2301 (M)-Barreiras, Bahia; B. IB 51310
(M)-Barreiras, Bahia; C. IB 54921 (M)-Brumado,
Bahia; D. IB 9152 (M)-Cana Brava, Goias.
Redescription of the Holotype
The description of the holotype
(UMMZ 108880 F) is as follows: rostral
wider than high; prefrontals twice as long
as the intemasals. The frontal is shorter
than its distance from the snout and the
parietals are slightly shorter than their dis-
tance from the snout (Fig. 20). There are
1 + i temporals; 7 + 7 supralabials; 7 +
ventral and 22 subcaudal scales. The snout
is pale with black bordered scales (inter-
nasals and prefrontals). The frontal and su-
praoculars are almost completely black
with the exception of an irregular margin
lateral and posteriorly and between the
scales. There are 12 (10 + 2) black body
triads as reported by Roze (1967). The
white triad rings (black tipped) are longer
than the black in the holotype and all the
paratypes and additional specimens of this
form. The red rings are mostly immaculate
and longer than the white ones. The first
triad reaches to within 1-3 scales of the
parietals, which are mostly red with the
168 HERPETOLOGICAL MONOGRAPHS
anterior part black, together with the fron-
tal and supraoculars; a transverse black
band crosses the head through the eyes
and is longest at the midline of the head
(Fig. 20). The overall length of the holo-
type is 865.4 mm with a head length of
22.9 mm, a SVL of 817.3 mm, and a TL
of 48.1 mm. The TJL/SVLratio is 0.059.
Variation
The males showed a range of 218 to 234
ventral scales, 20 to 25 subcaudal scales,
and 10 to 14 triads (n = 8; Table 1). Fe-
males showed a range of 229 to 237 ven-
tral scales, 16 to 23 subcaudal scales, and
12 to 13 triads (n = 3; Table 1). Extreme
body lengths were 320 to 1513 mm. It is
possible to separate this species from M.
frontalis based on the fixed characters of
white snout and longer white triad rings in
M. brasiliensis. This species does not have
any other geographic distribution contact
with the remaining taxa of this complex.
Additional specimens from the type-lo-
cality (IB 51310 M; CEPB 2301 M) and
vicinity (Sao Desiderio, Bahia-IB 54848 F)
present a similar pattern of head colora-
tion described for the holotype (Fig. 21).
Roze (in litt., 1992) suggested an intergra-
dation zone between M. f frontalis and M.
f brasiliensis north of Brasilia with the
specimen IB 40205 (M) from Padre Ber-
nardo (Goias) as an intermediary form.
The head coloration pattern and scale
counts on this specimen are widespread
within the populations of M. frontalis of
central-eastern Brazil. Specimens from
Brasilia (IB 20541 M; IB 20542 M; IB
20543 F; IB 33895 M) show a variable de-
gree of darkening on the snout. In Nique-
landia (about 180 km N of Brasflia) spec-
imens have darker snout (IB 48813 F; IB
48951 M; CEPB 0336 F) (Fig. 21), but the
sample from Serra da Mesa (n = 70) (80
km N of Niquelandia) already show some
mixed characters between M. frontalis and
M. brasiliensis (i.e., longer white triad
rings) but none are fixed. A specimen (IB
9152 M) from Cana Brava (90 km N-NE
from Niquelandia) matches the descrip-
tion of M. brasiliensis with respect to head
coloration and triad pattern (Fig. 21) and
we tentatively assign this specimen to this
[No. 13
taxon. In Guarani de Goias (186 km E of
Serra da Mesa) there is a recently collect-
ed specimen of M. brasiliensis (IB 55883
M) which is also only 280 km SW of the
type-locality (Barreiras, BA). The M. brasi-
liensis X M. frontalis transition zone is
somewhere between 15? and 14? S E-NE
of Serra de Mesa.
Hemipenial Morphology
The hemipenis is weakly bilobed, with
very short lobes covered by uniform size
spines. The everted organ is six subcaudals
long with lobes formed at the fifth/sixth
subcaudal level. The hemipenis of the
CEPB 2301 specimen is 15.7 mm long,
representing 1.9% of the SVL (SVL = 820
mm) and 32.7% of the TL (TL = 48.0
mm). The sulcus spermaticus bifurcates at
almost the same level of the lobes extend-
ing centripetally to the distal tip of the
lobes. The intrasulcar surface is smooth.
The semicapitate line is at the second sub-
caudal level. In the sulcate and asulcate
surfaces the spines are large and uniform
to the base of the hemipenis which is na-
ked. There is no evident change in spine
size below the semicapitate line (Fig. 22).
Cranial Osteology
In M. brasiliensis the dorsomedial pro-
cesses of the prefrontal bones are strongly
developed and may or may not contact at
midline. At midline the dorsomedial pro-
cesses are wide. An anteromedial process
may be present and separates the dorso-
medial processes. The braincase (parietals)
is rounded at the posterior end. The su-
ture prefrontals + frontals is anterior to
the maxilla + pterygoid suture. There are
8 to 9 palatine, 3 to 4 pterygoid, and 9 to
10 dentary teeth (Fig. 23).
Remarks
At approximately 580 km N-NW from
Brasilia is Ilha do Bananal (Tocantins), the
locality for one of the paratypes of M.
brasiliensis (AMNH 90361 M-Santa Isabel
do Morro). This specimen has some re-
duced black color on the snout but not as
much as in the holotype or the specimen
from Cana Brava (IB 9152 M). East of Ilha
do Bananal the next record is in Barreiras
1999]
1999] HERPETOLOGICAL
HERPETOLOGICAL MONOGRAPHS
MONOGRAPHS 169
169

A B
FIG. 22.-Detail of the hemipenis of Micrurus brasiliensis (CEPB 2301-left organ). A. Sulcate view; B.
Asulcate view. Total length of the hemipenis = 15.7 mm.

(Type-locality), a distributional gap of ap-
proximately 620 km. Also, there is a gap of
375 km from Barreiras south to Januaria,
Minas Gerais (UMMZ 108878 M-para-
type). In assuming the validity of this tax-
on, the distribution is triangular and in-
cludes the specimens from Januaria
(UMMZ 108878 M), Ilha do Bananal
(AMNH 90361 M), Guarani de Goias (IB
55883 M), and Barreiras (UMMZ 108880
M; UMMZ 108881; MNRJ 2494 M; IB
51310 M; CEPB 2301 M). In this case, the
records for Cana Brava (IB 9152 M) and
Sao Desiderio (IB 54848 F; IB 55385 M)
would fall within this western range. East
of the type-locality there are only three ad-
ditional records for this taxon in Brumado
(IB 50142 M) and Santa Cruz (MCZ 3298
F), Bahia, and Manga, Minas Gerais
(MNRJ 2497 F). The record for Santa
Cruz (MCZ 3298 F) is in fact Santa Cruz
de Cabralia. The specimen was collected
in 1865 by D. Bourget, a French naturalist
who lived in Rio de Janeiro and joined the
Thayer Expedition to Brazil (1865-1866),
led by Louis Agassiz (see Dick, 1977). The
type-locality (Barreiras, Bahia) also marks
the western boundary of M. ibiboboca, a
typical Caatinga coralsnake that has a char-
acteristic immaculate white triad rings and
snout and reduced black head coloration,
similar to M. brasiliensis. While all of the
170 HERPETOLOGICAL MONOGRAPHS
FIG. 23.-Detail of the skull of Micrurus brasilien-
sis (CEPB 2301). A. Dorsal view; B. Lateral view.
Total length of the skull = 22.4 mm.
taxa in the M. frontalis complex have dis-
tributions in tropical, subtropical, and
temperate forests, savannah, and second-
ary growth vegetation, M. ibiboboca has
successfully occupied the semi-arid xero-
morphic vegetation of Caatinga. Within
this same geographic area M. lemniscatus
is characterized by a reduced black head
coloration and a long white band on the
snout, and the resemblance of head pat-
tern and white triad rings among M. brasi-
liensis, M. ibiboboca, and M. lemniscatus
has created taxonomic chaos. The state of
Bahia has the northeasternmost records
for the M. frontalis complex (Brumado
and Santa Cruz de Cabralia), and together
with the records of Barreiras and Desider-
io, it seems that this might represent the
real geographical distribution of this taxon
(M. brasiliensis). These localities (Bahia),
are in a transitional zone between Cerrado
and Caatinga.
Geographical Distribution
In the original description, Roze (1967)
gives no defined geographic range for this
taxon, and which he later proposed (Roze,
1983) to be southeastern Brazil and again
changed to E Bahia and Goias to NE Mato
Grosso and southward to N Minas Gerais
No.
[No. 13
(Roze, 1996). In fact this geographic range
does not reflect reality. All of the speci-
mens that match the original description
of M. brasiliensis (including the holotype
and paratypes) are from central-north Bra-
zil, excluding Mato Grosso. There are no
data available to support the geographic
distribution proposed by Campbell and
Lamar (1989) for M. brasiliensis in Mato
Grosso or Mato Grosso do Sul.
The available specimens are scattered
over a wide range east of the Rio Araguaia
between 10? and 14? S to 41? W and south
to 16? S between 41? and 46? W. M. brasili-
ensis is sympatric with M. frontalis be-
tween 14? and 16? S, in NE Goias, N Mi-
nas Gerais and S-SW Bahia (Fig. 7).
Populations assigned to M. brasiliensis
contact M. ibiboboca in Barreiras (Bahia),
and within this range it is also sympatric
with M. lemniscatus. The Rio Araguaia
may serve as a western natural barrier to
M. brasiliensis.
Micrurus diana Roze
Micrrus frontalis diana Roze 1983:324
Holotype: FMNH 159889 (M).
Type-locality: Serrania de Santiago, De-
partamento de Santa Cruz, Bolivia.
Paratypes: FMNH 195864 (M); FMNH
195886 (M); FMNH 195899 (F); AMNH
120600 (M)-all from the same locality as
the holotype.
Micrurus frontalis diana Campbell and
Lamar 1989:115
Micrurus diana Roze 1994:179
Micrurus diana Roze 1996:152
Roze (1983) described this taxon based
on five specimens from the Serranfa de
Santiago (Bolivia) and since then only two
additional specimens have become avail-
able as museum vouchers. This form was
first described as a subspecies of M. fron-
talis (Roze, 1983), and later elevated to a
full species (Roze, 1994, 1996) without any
further comments or comparisons with re-
lated taxa.
Diagnosis
Micrurus diana is a triadal coralsnake
with a white snout, including the prefron-
tals, preoculars, anterior 3-4 supralabials
1999] HERPETOLOGICAL MONOGRAPHS
A
E
L rals; 7 + 7 supralabials; 7 + 7 infralabials;
_ i
C . ... : . ::-:::
FIG. 24.-Morphological characteristics of the Mi-
crurus diana holotype (FMNH 159899 M). A. Detail
of the head pattern; B. Detail of the first V3 of the
body (gray shading = red); C. Photograph of the
specimen.
(all black bordered or with irregular black
markings) and the anterior 3 of the frontal
and supraoculars. The head is black with
the parietals showing very little reduction
on the sides (Fig. 24). All temporals, pos-
terior 3-4 supralabials are red (immacu-
late). Inferiorly, the chin is white from
mental to anterior genials and 3 anterior
infralabials. Some irregular black markings
may be present, and the remainder is red.
The first triad is always fused with the pos-
terior end of the parietals (fixed character).
The white rings are longer than the black
ones which tend to be semiequal in length
(Table 2). The white rings are mildly
black-tipped, and the red rings are im-
maculate. Triads range from 10 to 11 (Ta-
ble 1).
Redescription of the Holotype
The description of the holotype
(FMNH 195889 M) is as follows: rostral
wider than high; prefrontals longer than
the intemasals. The frontal is as long as its
171
distance from the snout and the parietals
are slightly shorter than their distance
from the snout. There are 1 + 1 tempo-
0 + 2 oculars. It has 215 ventral and 22
subcaudal scales. It has a light snout that
reaches the anterior part of the frontal and
supraoculars, with the internasals and pre-
frontals bordered by black posteriorly. The
parietals are almost completely black and
the coloration fuses with the external ring
of the first triad (Fig. 24). The overall
length of the specimen is 1008.5 mm, with
a head length of 10.2 mm, a SVL of 954.4
mm, and a TL of 54.1 mm. The TL/SVL
ratio is 0.057. Inferiorly, the head is mostly
immaculate with the exception of a medi-
an black line between the first infralabials
and mentals. There are 11 (9 + 2) black
body triads. The black rings are of equal
length separated by longer white rings
(with very few black bordered scales). The
red rings are immaculate.
Variation
As it is true with M. tricolor and M.
brasiliensis, the number of available
vouchers of this taxon is very small. The
males showed a range of 215 to 229 ven-
tral scales, 22 to 24 subcaudal scales, and
10 to 11 triads (n = 5; Table 1). Both fe-
males have 227 ventral and 19 subcaudal
scales, with 10 and 11 triads respectively
(n = 2; Table 1). This taxon can be differ-
entiated from M. frontalis by the fixed
character of the fusion of the anterior
black triad ring with the posterior end of
the parietals, long white triad rings and
lower triad number. Also, it does not have
any combination of characters found in M.
pyrrhocryptus or M. tricolor. Extreme
body lengths (SVL + TL) were 516 and
1008 mm.
Hemipenial Morphology
The hemipenis is weakly bilobed with
short lobes covered by uniform size spines
(Fig. 25). The everted organ is seven sub-
caudals long with lobes formed at the
sixth/seventh subcaudal level. The hemi-
penis of the FMNH 195864 (paratype)
specimen is 16.2 mm long, representing
1.7% of the SVL (SVL = 956 mm) and
172
172 MONOGRAPHS
HERPETOLOGICALMONOGRAPHS
HERPETOLOGICAL
32.4% of the TL (TL = 50.0 mm). The
sulcus spermaticus bifurcates at the sixth
subcaudal level and extends centripetally
to the distal tip of the lobes. Among the
species included in this study M. diana
hemipenes have the lowest bifurcation of
the sulcus spermaticus, preceding the bi-
furcation of the lobes (as opposed to these
features bifurcating at the same point).
The intrasulcar surface is smooth. The
semicapitate line is discretely evident at
the second subcaudal level. Above this line
to the tip of the lobes the spines are fewer
and smaller, and are irregularly distribut-
ed. The sulcate surface is completely cov-
ered by uniform size spines with the base
of the hemipenis naked (Fig. 25).
Cranial Osteology
In M. diana the dorsomedial processes
of the prefrontal bones are strongly devel-
oped and contact each other at midline. At
midline the dorsomedial processes are nar-
row. The anteromedial processes of the
frontals are absent. The braincase (parie-
tals) is rounded at the posterior end. The
suture prefrontals + frontals is anterior to
the maxilla + pterygoid suture. There are
9 to 11 palatine, 4 to 5 pterygoid, and 9 to
10 dentary teeth (Fig. 26).
Geographic Distribution
This species is not restricted to the Ser-
rania de Santiago, Santa Cruz, Bolivia, as
suggested by Roze (1983, 1996). Mr. P.
Bettella (NKR) mentioned (in litt., 1991)
a record for San Ramon, Santa Cruz, for
which we received a color slide, but no
additional data on the scutellation or
where and if it was deposited as a voucher.
This record is about 170 km NW of the
Serrania de Santiago. Another record
NKR 219 (F) is from Serrania Huanchaca
on the extreme NE of the province of San-
ta Cruz, and a third specimen IB 31342
(M) has no specific locality other than the
province of Santa Cruz. These records
conflict with the idea of an isolated pop-
ulation (Fig. 7), and over a larger area this
species may be sympatric with M. fronti-
fasciatus, M. pyrrhocryptus and possibly
with M. lemniscatus.
Micrurus pyrrhocryptus (Cope)
[No. 13
[No.
Elaps pyrrhocryptus Cope 1862:347
Holotype: ANSP 5395
Type-locality: Vermejo River, Argentine
Chaco, Corrected to Rio Bermejo, Chaco,
Argentina by Scrocchi (1990:359).
Elaps marcgravii Boulenger 1896:428.
Elaps pyrrhocryptus Berg 1898:29.
Elaps simonsii Boulenger 1902:338 (Type-
locality: Cruz del Eje, Cordoba, Argen-
tina).
Elaps frontalis Alvarez 1919:159
Elapsfrontalis Serie 1919:417
Micrurus pyrrhocryptus Schmidt 1936:
199
Micrurus lemniscatus frontalis (partim)
Amaral 1944:92
Micrurus frontalis pyrrhocryptus Schreve
1953:5
Micrurus lemniscatus frontalis Ringuelet
and Aramburu 1957:32
Micrurus pyrrhocryptus Hoge and Lanci-
ni 1960:12
Micrurus lemniscatus Abalos 1961:70
Micrurus lemniscatus frontalis Abalos and
Nader 1962:83
Micrurus lemniscatus frontalis Abalos and
Bucher 1970:264
Micrurus lemniscatus frontalis Abalos,
Baez, and Nader 1964:267
Micrurus frontalis pyrrhocryptus Roze
1967:26
Micrurus lemniscatus frontalis Abalos and
Mischis 1975:74
Micrurus pyrrhocryptus Hoge and Ro-
mano 1979:400
Micrurus frontalis pyrrhocryptus Roze
1983:326
Micrurus frontalis pyrrhocryptus Esteso
1985:41
Micrurus frontalis pyrrhocryptus Golay
1985:34
Micrurus frontalis pyrrhocryptus Camp-
bell and Lamar 1989:116
Micrurus pyrrhocryptus Scrocchi 1990:
359
Micrurus pyrrhocryptus pyrrhocryptus
Roze 1994:179
Micrurus pyrrhocryptus pyrrhocryptus
Roze 1996:212
As with M. baliocoryphus this form has
been subject to many taxonomic changes
1999] HERPETOLOGICAL MONOGRAPHS
HERPETOLOGICAL MONOGRAPHS 173
173

A4 B
FIG. 25.-Detail of the hemipenis of Micrurus diana (FMNH 195864-left organ). A. Sulcate view; B.
Asulcate view. Total length of the hemipenis = 16.2 mm.

in the past owing to a lack of understand-
ing of the M. frontalis complex. Unfortu-
nately, the holotype (ANSP 5395) is miss-
ing but Cope's description (1862) is
accurate enough to characterize this taxon.
The main characteristics are the black
head and the triad pattern of a middle
black ring twice as long as the external
ones, and a very low triad number of 7 (6
+ 1). All the scales comprising the red
rings are tipped with black on the poste-
rior margins.
Scrocchi (1990) reviewed most of the
available museum specimens of this taxon
in Argentina but he did not mention the
sex of the specimens in the list of the stud-
ied material, and a description of a M. pyr-
rhocryptus hemipenis is given without ref-
erence to a museum voucher number. It
also appears that Scrocchi did not compare
the single specimen with other individuals
of M. pyrrhocryptus in order to distin-
guish between diagnostic and variable
hemipenial characters. The comparison
174 HERPETOLOGICAL MONOGRAPHS
FIG. 26.-Detail of the skull of Micrurus diana
(FMNH 195864). A. Dorsal view; B. Lateral view.
Total length of the skull = 21.3 mm.
with Roze's (1983) description of the M.
diana hemipenis is meaningless because
both are forms of a bigger and more com-
plex group with unresolved taxonomic is-
sues. This author (Scrocchi, 1990) pre-
sented a list of specimens with mixed
characters between M. pyrrhocryptus and
the other subspecies of M. frontalis. If we
assume that the geographic distribution of
M. pyrrhocryptus is correct, there is no
indication of the presence of this taxon in
the Argentine Mesopotamia and all of the
specimens listed for Corrientes and Mi-
siones deal with identification problems
between M. baliocoryphus and M. altiros-
tris, and not with M. pyrrhocryptus and
M. altirostris.
Diagnosis
Micrurus pyrrhocryptus is a triadal cor-
alsnake with a black snout reaching the
postoculars and 3-4 anterior supralabials.
All scales are white bordered. The head is
black (Fig. 27). The frontal, supraoculars
and parietals may be white bordered in-
cluding along the medial suture of the pa-
rietals. The remainder of the head is red
with all scales black-tipped. Inferiorly, the
chin is red with irregular black markings
on the infralabials, genials, and first ven-
trals. This taxon is characterized by a low
[No. 13
FIG. 27.-Morphological characteristics of a Mi-
crurus pyrrhocryptus specimen (CEPB 2364 M). A.
Detail of the head pattern; B. Detail of the first % of
the body (gray shading = red); C. Photograph of the
specimen.
number of triads. The middle black ring is
at least twice (10-14 dorsal scales) the
length of the external ones (5-7 dorsal
scales). The white rings (always black-
tipped) are of the same length or slightly
shorter than the external black rings (4-6
dorsal scales) (Table 2). Owing to the low
number of triads the red rings are almost
the same length as the entire triads. The
red rings are heavily darkened (black-
tipped) and an intense melanism of the
head and body is characteristic of this tax-
on. The first triad is separated from the
parietals by 5 to 7 dorsal scales. Triads
range from 6 to 11 (Table 1).
Redescription of the Holotype
Since the holotype of M. pyrrhocryptus
(ANSP 5395) is missing from the ANSP
herpetological collection, we present the
specimen CEPB 2364 (M) from Santiago
del Estero (Argentina), as a reference to
which we recognize as a typical M. pyr-
rhocryptus. The rostral is wider than its
length, and the prefrontals are longer that
the internasals. The frontal and parietals
1999] HERPETOLOGICAL MONOGRAPHS
are slightly shorter than their distance
from the snout. There are 1 + 1 tempo-
rals; 7 + 7 supralabials; 7 + 7 infralabials;
0 + 2 oculars. The specimen has 228 ven-
tral and 23 subcaudal scales with 8 (7 +
1) black body triads. The snout and head
are completely black until the parietals
and post-ocular scales. Inferiorly, there are
few black markings on the chin. There is
a distance of 6 dorsal scales between the
black color of the head and the first triad
(Figure 27). The overall length of the
specimen is 1090 mm with a head length
of 25.4 mm, a SVL of 1042.5 mm and a
TL of 47.5 mm. The TL/SVL ratio is
0.046.
Variation
Males showed a range of 218 to 239
ventral scales, 20 to 28 subcaudal scales,
and 6 to 11 triads (n = 196; Table 1). Fe-
males had a range of 221 to 251 ventral
scales, 19 to 25 subcaudal scales, and 8 to
10 triads (n = 19; Table 1). Extreme body
lengths were 224 and 1241 mm. M. pyr-
rhocryptus can be differentiated from the
other taxa by the lowest number of triads
and the longest middle black triad ring.
Hemipenial Morphology
The hemipenis is weakly bilobed with
short lobes covered by uniform size spines
(Fig. 28). The everted organ is six subcau-
dals long with lobes formed at the fifth/
sixth subcaudal level. The hemipenis of
the CEPB 1872 specimen is 14.2 mm
long, representing 1.7% of the SVL (SVL
= 830 mm), and 28.6% of the TL (TL =
49.6 mm). The sulcus spermaticus bifur-
cates at almost the same level of the bi-
furcation of the lobes and extends centrip-
etally to the distal tip of the lobes. The
intrasulcar surface is smooth. The semi-
capitate line (not always evident) runs at
the second subcaudal level. The spines
above this line to the tip of the lobes are
large and uniform. Below the line the
spines have a tendency to be smaller and
irregularly distributed but not as a naked
surface as described by Scrocchi (1990).
The sulcate surface has a uniform pattern
of large spines. The base of the hemipenis
is naked (Fig. 28).
175
A B
FIG. 28.-Detail of the hemipenis of Micrurus
pyrrhocryptus (CEPB 1872-right organ). A. Sulcate
view; B. Asulcate view. Total length of the hemipenis
= 14.2 mm.
Cranial Osteology
In M. pyrrhocryptus the dorsomedial
processes of the prefrontal bones are
strongly developed and contact each other
at midline. At midline the dorsomedial
processes are wide. The anteromedial pro-
cesses of the frontals are absent. The
braincase (parietals) is pointed at the pos-
terior end. The prefrontals + frontals su-
ture is at the same level or anterior to the
maxilla + pterygoid suture. There are 9 to
11 palatine, 4 to 5 pterygoid, and 9 to 11
dentary teeth (Fig. 29).
Remarks
In the Argentine Province of Formosa
the specimen CHINM 2178 (M) has 15
(14 + 1) body triads, which is an extreme
outlier compared to the most common
characteristic of M. pyrrhocryptus (see
Fig. 27), but it has a triad pattern com-
pletely similar to the typical M. pyrrho-
cryptus. However, if we analyze the other
10 specimens available from the same lo-
cality, the triad number ranges from 8 to
10. To interpret the specimen CHINM
2178 (M) as a M. pyrrhocryptus we have
to accept one of three possibilities: 1) this
is an extreme example of variation in M.
176 HERPETOLOGICAL MONOGRAPHS
FIG. 29.-Detail of the skull of Micrurus pyrrho-
cryptus (CEPBc 5977). A. Dorsal view; B. Lateral
view. Total length of the skull = 22.0 mm.
pyrrhocryptus; 2) this is in fact M. balio-
coryphus and its distribution is not re-
stricted to the Argentine Mesopotamia; or
3) this is a M. frontalis. We accept the sec-
ond possibility (= M. baliocoryphus).
Since we pointed out that the distribution
of M. baliocoryphus is not restricted to the
Argentine Mesopotamia these specimens
are the closest to the form present in Par-
aguay, which also lack the white band on
the head. Even with the confirmed pres-
ence of M. frontalis in the Paraguayan
Central province, the triad pattern of the
Formosan specimens are closer to M. bal-
iocoryphus and we did not find any M.
frontalis specimens on the west side of the
Rio Paraguay. In fact, this makes more
sense because another specimen CHINM
3650 (M) from Las Lomitas (-300 km
NW of Formosa) also has 14 triads and a
triad pattern not typical of M. pyrrhocryp-
tus. The sample of M. pyrrhocryptus in-
cluded in this study shows only three spec-
imens with 12, 13, and 14 body triads
respectively, and the majority of specimens
have between 6 and 11 triads, making it
difficult to accept these two specimens
from Formosa as part of this taxon. The
specimen CHINM 712 (M) also shows a
[No. 13
[No.13
triad pattern and number different than
M. pyrrhocryptus. Based on these obser-
vations we tentatively recognize these
three specimens as M. baliocoryphus. If
there is an intergradation zone between
M. baliocoryphus and M. pyrrhocryptus,
the province of Formosa and the adjacent
Paraguayan provinces along Rio Paraguay
are geographically the best candidates so
far. Also, triad, head color patterns, and
scutellation suggest that M. baliocoryphus,
M. tricolor, and M. pyrrhocryptus may be
more similar to each other than to M. fron-
talis and M. altirostris.
Geographic Distribution
In Argentina this taxon has a distribu-
tion to the west of Rio Parana and south
to the province of Rio Negro, and ranges
north to the province of Formosa, and
northwest into Paraguay (Departamento
Boqueron) and Bolivia (Departamento
Santa Cruz), possibly throughout the prov-
inces of Salta and Jujuy. Roze (1983) men-
tioned a specimen of M. pyrrhocryptus
from the Serranmade San Jose de Chiquili-
tos (160 Km W of Serrania de Santiago),
but there is no museum voucher to sup-
port this distribution. However, there are
several specimens at the NKR collection
from the metropolitan area of Santa Cruz
de la Sierra and this could represent the
northernmost record for this species. To
the west the increasing altitudes of the An-
des and to the south the lower tempera-
tures of the high latitudes may act as nat-
ural barries (Fig. 30). Oddly, no specimen
of any taxon was available for the province
of Buenos Aires. This taxon is probably
sympatric with M. baliocoryphus in north-
ern Formosa, Argentina (and possibly with
M. frontalis) and with M. diana and M.
tricolor in Bolivia.
Micrurus tricolor Hoge
Micrurus tricolor Hoge 1956:67
Holotype: IB 16290 (M)
Type-locality: Carandazal, Mato Grosso
do Sul. At the time of this paper the single
state of Mato Grosso was later divided into
Mato Grosso (North) and mato Grosso do
Sul (South). Garandazal is a spelling error.
19991
1999] HERPETOLOGICAL MONOGRAPHS
FIG. 30.-Distribution of Micrurus pyrrhocryptus
(open diamonds) and M. tricolor (black squares). Star
represents the type-locality of M. tricolor (Caran-
dazal, Mato Grosso do Sul-Brazil). Scale represents
250 km.
The correct name of the type-locality is
Carandazal.
Micrurus frontalis tricolor Roze 1983:326
Micurus frontalis tricolor Golay 1985:34
Micrurus frontalis tricolor Campbell and
Lamar 1989:116
Micrurus tricolor Strussmann and Sazima
1993:163
Micrurus pyrrhocryptus tricolor Roze
1994:179
Micrurus pyrrhocryptus tricolor Roze
1996:212
Hoge (1956) described M. tricolor (IB
16290 M) from the S-SW of Mato Grosso
do Sul (Brazil) based on 9 specimens from
Carandazal, Barranco Branco, Guia Lopes
da Laguna, and Taunay. Roze (1967) syn-
onymized it with M. f pyrrhocryptus and
separated them again later (Roze, 1983),
referring to Hoge's original description.
Probably for this 1967 paper, Roze had not
seen the M. tricolor specimens at IB and
177
gave no reasons for synonymizing this
form with M. f pyrrhocryptus, nor to the
later split (Roze, 1983). The synopsis of
Hoge and Romano (1979) failed to address
the problem with M. f tricolor. Scrocchi
(1990) in analyzing a much larger sample
size of M. pyrrhocryptus, decided to syn-
onymize M. f tricolor, as a junior synonym
of M. pyrrhocryptus, based on the overlap
of scale counts (ventrals and subcaudals)
and body triad number. Apparently this
author based his decision on Roze's (1983)
comments without checking Hoge's de-
scription or the specimens at the IB her-
petological collection, as they were not in-
cluded in the list of examined specimens
in his paper (Scrocchi, 1990). Indeed,
Hoge's specimens (1956) share some sim-
ilarities with M. pyrrhocryptus especially
with regard to the triad pattern. However,
Scrocchi's (1990) assumptions may lead to
erroneous interpretations based on the
small sample size of M. tricolor and the
absence of any statistical analysis of the
data.
Strussmann and Sazima (1993) further
confused the situation by resurrecting M.
tricolor as full species based solely on their
ecological opinion. No taxonomic reason-
ing was given for this decision nor did
these authors provide any comparative de-
scription, diagnosis on the basis of unique
characters, or sample sizes compared.
Strussmann and Sazima (1993) added one
more locality to the known distribution of
this taxon (Pocone) but did not mention if
any specimen was collected and/or if it was
deposited as a museum voucher. Finally
Roze (1994, 1996) placed M. tricolor as a
subspecies of M. pyrrhocryptus also with-
out comments or further analysis. The big-
gest problem in accepting Scrocchi's
(1990) and Roze's (1994, 1996) taxonomic
arrangements is that M. tricolor is not an
isolated population in Mato Grosso do Sul,
but it extends into Bolivia where it is sym-
patric with M. pyrrhocryptus and possibly
M. diana.
Diagnosis
Micrurus tricolor is a triadal coralsnake
with a black snout covering all scales with
very little white bordering, which when
178 HERPETOLOGICAL MONOGRAPHS
A
C
FIG. 31.-Morphological characteristics of the Mi-
crurus tricolor holotype (IB 16290 M). A. Detail of
the head pattern; B. Detail of the first % of the body
(gray shading = red); C. Photograph of the specimen.
present is mostly restricted to the supra-
labials. The frontal, supraoculars and pa-
rietals are completely black, which some-
times reaches the first temporals (Fig. 31).
The remainder of the head is red with all
scales black-tipped. Inferiorly, the chin is
red with irregular black markings (when
present) mostly restricted to the infrala-
bials. The distance from the parietals to
the first triad ranges from 7 to 11 dorsal
scales. The triads are composed of a mid-
dle black ring longer (5-7 dorsal scales)
than the external ones (3-4 dorsal scales)
(Table 2). The white rings (immaculate)
are of the same length as the external
black rings. In all red rings the scales are
black-tipped. Triads range from 8 to 12
(Table 1).
Redescription of the Holotype
The holotype of M. tricolor (IB 16290
M) has a completely black snout and head
with the exception of a narrow white mar-
gin on the anterior border of the parietal
scales. Inferiorly, there are irregular black
markings on almost all of the infralabials.
The specimen has 220 ventral and 29 sub-
[No. 13
caudal scales. There are 11 (10 + 1) black
body triads with the middle black ring
twice as long as the external ones. The
white rings (immaculate) are longer than
the external black rings. The red rings are
heavily tipped with black. The overall
length of the specimen is 1146.6 mm with
a head length of 21.3 mm, a SVL of 1065.2
mm, and TL of 81.4 mm. The TL/SVL ra-
tio is 0.076 (Fig. 31).
Variation
Male ventral scales ranged from 220 to
234, 24 to 29 subcaudal scales, and 8 to 11
triads (n = 13; Table 1). Female ventral
scales ranged from 217 to 231, 19 to 26
subcaudal scales, and 9 to 11 triads (n =
6; Table 1). Overall there is not a tendency
to melanism as seen in M. pyrrhocryptus.
It is possible to distinguish M. tricolor
from M. pyrrhocryptus based on lower
number of ventral scales in females, and
higher number of triads in M. tricolor.
Also, in M. tricolor the middle black ring
in considerably shorter, the white rings are
longer, and the external black rings are
shorter than in M. pyrrhocryptus. Ex-
treme body lengths were 436 and 1181
mm.
Hemipenial Morphology
The hemipenis is weakly bilobed with
short lobes covered by uniform size spines
(Fig. 32). The everted organ is seven sub-
caudals long with lobes formed at the
sixth/seventh subcaudals level. The hemi-
penis of the IB 14566 specimen is 16.3
mm long, representing 1.8% of the SVL
(SVL = 926 mm), and 32.5% of the TL
(TL = 50.1 mm). The sulcus spermaticus
bifurcates at almost the same level of the
bifurcation of the lobes extending centrip-
etally to the distal tip of the lobes. The
intrasulcar surface is smooth. The semi-
capitate line is evident at the third subcau-
dal level. In the asulcate surface the spines
are larger and uniform from the second
subcaudal to the tip of the lobes. Below
the second subcaudal there are smaller
and irregularly distributed spines that
reach the base of the hemipenis. The sul-
cate surface is completely covered by large
uniform size spines except the base of the
19991
1999] HERPETOLOGICAL MONOGRAPHS
HERPETOLOGICAL MONOGRAPHS 179

FIG. 33.-Detail of the skull of Micrurustricolor

(IB 53454). A. Dorsal view; B. Lateral view. Total
length of the skull = 24.6 mm.

A B Remarks
FIG. 32.-Detail of the hemipenis of Micrurustri- Additional specimens are also available
color (IB 14566- left organ). A. Sulcate view; B.
Asulcate view. Total length of the hemipenis = 16.3 from the localities of Aquidauana (IB 5535
mm. M, IB 16520 M, IB 19616 M), Corumba
(IB 53454 M), Guaicurus (IB 30555 F, IB
31575 F, IB 32712 M), and Porto Murtin-
ho (IB 19772 F), and together with the
hemipenis which is almost completely na- specimens used in the original description
ked with nine to twelve small spines (Fig. they total 17. A single specimen (COBAP
32). 058 M) from Anastacio (8 km south of
Cranial Osteology Aquidauana) and another from Santa Cruz
(Bolivia-NKR 543M), also have the com-
In M. tricolor the dorsomedial processes bination of characters of M. tricolor (Fig.
of the prefrontal bones are strongly devel- 31), but the taxon remains poorly known.
oped and do not contact at midline. They
are united by connective tissue. At midline Geographic Distribution
the dorsomedial processes are wide. The The distribution of this species in Brazil
anteromedial processes of the frontals are is restricted to S-SW of the State of Mato
absent. The braincase (parietals) is pointed Grosso do Sul and extends into eastern
at the posterior end. The suture prefron- Bolivia in the Departamento Santa Cruz
tals + frontals is anterior to the maxilla + (Fig. 30). This taxon may also have a dis-
pterygoid suture. There are 8 to 9 palatine, tribution in Paraguay through the Depar-
4 to 5 pterygoid, and 9 to 10 dentary teeth tamento de Alto Paraguay and possibly
(Fig. 33). Concepcion.
KEY TO THE SPECIES OF THE MICRURUSFRONTALIS
COMPLEX
1. a. Head (frontal,supraocularsand parietals)completely black. ...................... 2
b. Head with reduction of black on parietals .................................... 5
2. a. Middle black triad ring longer than the external ones ........................... 3
b. Subequal black rings ...................................................... 6
3. a. Immaculatewhite rings .............................................. 4
b. Black-tippedwhite rings; 6 to 11 total triads ................ Micruruspyrrhocryptus
4. a. White snout (scales bordered by black) including anterior chin region; usually with a
180
180 MONOGRAPHS
HERPETOLOGICAL MONOGRAPHS
HERPETOLOGICALT iR
[No. 13
[Nn

white band on the head between frontal and parietals;9 to 16 triads ..............
...................................................... Micrurus baliocoryphus
b. Black snout; head black; neck and chin red; 7 to 11 dorsal scales from parietalsto first
triad;8 to 11 triads ........................................... Micrurus tricolor
5. a. Black reduced anteriorlyon parietals;black snout; chin heavily markedwith black;white
rings shorter than the externalblack ones; middle black ring sometimes longer than the
external ones; tendency towards melanism; 13 to 18 triads ......... Micrurusaltirostris
b. Black reduced posteriorlyon parietals;snout white including anteriorpart of the chin;
white rings (mostly immaculate)longer than any of the black rings; 11 to 14 triads . .
......................................................... Micrurus brasiliensis
6. a. White snout; white rings at least twice as long as the black rings; anterior black triad
ring always fused with the posterior end of parietals;red rings immaculate; 10 to 11
triads ........................................................ Micrurus diana
b. Snout usually black;white rings shorter or the same length as the externalblack rings;
10 to 15 triads .............................................. Micrurusfrontalis

DISCUSSION ditional forms in Rio Grande do Sul (Bra-

A. Taxonomic Arrangement of the
Frontalis Complex
The poor morphological descriptions
had chaotic consequences on the taxonom-
ic arrangements proposed up to this date
(Amaral, 1944; Roze, 1967, 1983, 1994,
1996; Schmidt, 1936; Scrocchi, 1990). Af-
ter Scrocchi's (1990) and Roze's (1994,
1996) latest revisions seven entities were
recognized as part of the Micrurus fron-
talis complex: 1) M. f frontalis; 2) M. f
altirostris; 3) M. f baliocoryphus; 4) M. f
brasiliensis; 5) M. f multicinctus; 6) M. di-
ana; and 7) M. pyrrhocryptus.
We redescribed M. frontalis based on
one of the existing syntypes (MNHN 854
M). In the original paper (Dumeril et al.,
1854) the authors also described other
specimens from Corrientes and Misiones,
Argentina that are missing from the
MNHN collection, together with four oth-
er specimens mentioned in the paper. We
suggest the assignment of the specimen
MNHN 854 (M) as the lectotype of M.
frontalis with Brazil (possibly Lagoa Santa,
Minas Gerais) as type-locality (the second
remaining specimen in this collection
[MNHN 578 M] is a M. isozonus). As de-
fined here, M. frontalis has the widest
range of all recognized taxonomic entities,
and extends over central Brazil, east to Sao
Paulo and Espirito Santo, west to Mato
Grosso and Mato Grosso do Sul, and south
to Paraguay (Fig. 7).
Another distinct form, M. altirostris
might be represented by one or two ad-
zil). Other than the form that matches the
general description of M. altirostris, relat-
ed to head coloration (reduction of black
color on parietal scales) and triad pattern
(semi-equal black rings and shorter white
rings), there is another form that has an
almost completely black head color (pari-
etal scales) with the middle black triad ring
twice as long as the external ones. The sec-
ond form has a longer middle black triad
ring with reduced black head coloration.
However, these characters are not fixed
and due to a very small sample size we
retain them as M. altirostris. Interestingly,
this variation was also noted by Schmidt
(1936), but not by Amaral (1944) whom
apparently had a much larger sample size
of this taxon. The distribution of M. alti-
rostris, as defined here, is restricted to the
east of the Rio Parana in the Brazilian
states of Parana, Santa Catarina and Rio
Grande do Sul, Uruguay, and the Argen-
tine Mesopotamia (Figs. 10, 11, 14).
The name M. f mesopotamicus is not
valid due to the priority of Cope's descrip-
tion (1862) of Elaps baliocoryphus over
Barrio and Miranda's description (1966).
Both holotypes are comparable in all mor-
phological characters examined, and our
arrangement also supports Roze's (1983)
suggestion for changing the type-locality
from Buenos Aires to Villa Federal. The
combination of morphological features re-
lated to scutellation (lower counts of ven-
tral and subcaudal scales in both sexes,
compared to M. pyrrhocryptus) and triad
patterns (higher number than M. pyrrho-
1999] HERPETOLOGICALMONOGRAPHS
HERPETOLOGICAL MONOGRAPHS
cryptus) supports the elevation of this
form to the rank of species, M. baliocor-
yphus. This taxon has a distribution in the
Argentine Mesopotamia, the province of
Formosa, and Paraguay, SW-W of Rio Par-
aguay (Fig. 14).
Two other taxa, M. brasiliensis and M.
diana, are recognized as full species al-
though both are represented by very small
sample sizes. M. brasiliensis is apparently
restricted to the northeast of Goias, north
of Minas Gerais and south of Bahia, while
M. diana is known from the type-locality
and adjacent areas in the province of Santa
Cruz (Fig. 7).
The holotype of M. pyrrhocryptus is
missing from the ANSP herpetological col-
lection, but after the examination of all
other available specimens, we agree with
the current arrangement of this taxon as
proposed by Scrocchi (1990). M. pyrrho-
cryptus is well distributed in Argentina,
reaching western Paraguay and southern
Bolivia (Fig. 30).
M. f tricolor is elevated to a full species
as opposed to a junior synonym of M. pyr-
rhocryptus as proposed by Scrocchi (1990)
and Roze (1996). There is evidence of
sympatry between M. tricolor and M. pyr-
rhocryptus in Bolivia and possibly far
western Brazil (Fig. 30), but a larger sam-
ple size is needed to draw more accurate
conclusions about this geographical distri-
bution.
With the re-examination of most of the
existing holotypes and paratypes, and larg-
er sample sizes it is possible to recognize
these taxa and arrange them into two dif-
ferent groups on the basis of overall mor-
phological similarities: M. frontalis and M.
pyrrhocryptus (details given below). All of
the pertinent forms have been redescribed
in this revision, and a detailed geographi-
cal distribution of each was plotted to aid
in reconstruction of patterns of speciation,
and biogeographic and phylogenetic rela-
tionships within both species groups (if
these are monophyletic), and the remain-
ing South American triad coralsnakes (Jor-
ge da Silva and Sites, in review).
The M. frontalis "group" is now com-
posed of three taxa: 1) M. frontalis; 2) M.
brasiliensis; and 3) M. diana. Micrurus di-
181
181
ana has the lowest number of body triads,
but is grouped with other M. frontalis taxa
on the basis of the scutellation data, and
triad and head pattern. Our molecular
phylogenetic study recovers M. brasiliensis
and M. frontalis as sister taxa, but because
we do not have tissues from M. diana, we
cannot yet be certain of the reality of a
monophyletic M. frontalis "group" (Jorge
da Silva and Sites, in review).
The M. pyrrhocryptus "group" includes
M. pyrrhocryptus, M. altirostris, M. bal-
iocoryphus, and M. tricolor which show
comparable scutellation characteristics in
number of ventral and subcaudal scales,
longer middle black triad ring and lower
number of triads. It is important to em-
phasize that this is a phenotic assessment
but that it is supported by geographic dis-
tribution; the M. pyrrhocryptus group taxa
have juxtaposed ranges in the southern re-
gion of the range of the M. frontalis com-
plex while the three entities of the M.
frontalis group occupy the northern and
eastern regions with a confirmed presence
in Central Paraguay. However, the molec-
ular phylogeny reveals strong support for
a (M. baliocoryphus + M. pyrrhocryptus)
clade (support index-11), and recovers M.
decoratus as the sister group of a clade
with the following topology: ((M. frontalis
+ M. brasiliensis)((M. ibibiboca + M. lem-
niscatus)(M. spixi((M. altirostris(M. balio-
coryphus + M. pyrrhocryptus))))) (Jorge
da Silva and Sites, in review).
For the first time M. lemniscatus was
described for Argentina, in the province of
Misiones (Jorge da Silva and Silva, 1994).
Owing to its unique triad pattern, immac-
ulate white rings and a white band across
the internasal scales with the tip of the
snout black, this taxon was misidentified as
M. f mesopotamicus (= M. baliocoryphus)
by Scrocchi (1990).
With the taxonomic changes suggested
in this study we recognize the South
American triad coral snakes as composed
of the following taxa: M. ancoralis, M. al-
tirostris, M. baliocoryphus, M. brasiliensis,
M. decoratus, M. diana, M. dissoleucus, M.
filiformis, M. frontalis, M. frontifasciatus,
M. hemprichii, M. ibiboboca, M. isozonus,
M. lemniscatus, M. meridensis, M. pyrrho-
182
182 HERPETOLOGICALMONOGRAPHS
cryptus, M. sangilensis, M. spixii, M. suri-
namensis, M. tricolor, and M. tschudii.
B. Species Boundaries and Biological
Diversity in the South American Triads
Ultimately the number of species rec-
ognized in the 'frontalis" complex will de-
pend on what operational criteria are used
to define species, and the extent of geo-
graphic and character sampling required
for their implementation. Species may be
defined on the basis of reproductive inter-
actions (biological or recognition con-
cepts), evolutionary "trajectories", phylo-
genetically diagnosable units, genetic/
demographic "cohesion", or genealogical
concordance (Avise and Wollenberg, 1997;
reviewed by Sites and Crandall, 1997).
The biological and recognition species
concepts (BSC and RSC) emphasize either
the evolution of reproductive barriers to
gene flow between populations (BSC), or
the origin of shared mate-recognition sys-
tems and reproductive "connectivity" with-
in populations (RSC). The evolutionary
species concept (ESC) emphasizes identi-
fication of allopatric populations that are
on separate evolutionary "trajectories"
(Frost and Hillis, 1990; Wiley, 1978), and
is usually tested by the criterion of phe-
notypic distinctiveness (Reichling, 1995).
The phylogenetic species concept (PSC)
defines a species as the smallest diagnos-
able cluster of individual organisms within
which there is a parental pattern of ances-
try and descent. The advantages of this
widely-used concept are that: (1) the
groups defined by the PSC are monophy-
letic units diagnosed by at least one de-
rived character state; (2) the PSC elimi-
nates the concern with reproductive
compatibilities (the BSC and RSC above),
which are usually untestable between al-
lopatric populations; and (3) the PSC fo-
cuses on the geographic and genealogical
histories of populations, irrespective of
modes of reproduction (i.e., unlike the
BSC and RSC, it is not restricted to bisex-
ual populations). Limitations of the PSC
are that: (1) the number of species rec-
ognized depends on the resolving power
of the characters and analytical methods
used, so that different data sets may lead
[No. 13
to different results; (2) given the rapid
growth of molecular character analysis in
phylogenetic studies, one must be cogni-
zant of the possibilities for different gene
genealogies leading to conflicting interpre-
tations of species boundaries; and (3) both
of these points, when compounded with
strongly subdivided metapopulation struc-
tures and recent speciation events, result
in paraphyletic species (Patton and Smith,
1994). These results are possible because
of unequal mutation rates and/or differ-
ential lineages sorting between unlinked
genes, recombination in nuclear gene se-
quences, and differential rates of intro-
gression for different gene loci in cases
where parapatrically distributed popula-
tion hybridize (Sites and Crandall, 1997).
Davis and Nixon (1992) have provided
stringent operational criteria for defining
phylogenetic species that distinguish be-
tween a character (a state fixed in all com-
parable individuals within a species), and
a trait (a state present in a population or
set of populations, but not characteristic of
all individuals [a polymorphism]). Al-
though unambiguous in definition, the ap-
plication of these criteria, and the number
of phylogenetic species identified by their
use, will be heavily dependent on the
number of individuals sampled at a single
locality, the number of localities sampled,
the total number of "markers" (genetic,
morphological, etc.) sampled, and the var-
iability within each marker.
One alternative to these concepts is the
"cohesion" species concept (CSC) first de-
fined by Templeton (1989): a cohesion
species is the most inclusive population of
individuals with the potential for pheno-
typic cohesion (i.e., similarity), through in-
trinsic "cohesion mechanisms" that can be
either genetic (i.e., gene flow) or demo-
graphic (i.e., ecological/selective, or genet-
ic drift). Templeton (1994) has further
elaborated on these ideas to provide a test-
able framework for the recognition of co-
hesion species. The advantage of this
method is that it can be applied as a nest-
ed set of testable null hypotheses, that un-
der the appropriate sampling design, will
incorporate both genetic and demographic
parameters (see also Templeton, 1998;
1999] HERPETOLOGICAL MONOGRAPHS
HERPETOLOGICALMONOGRAPHS 183
183

Templeton et al., 1995). Another alterna- tional approach similar to Reichling

tive is the principle of genealogical con- (1995), who faced similar limitations in
cordance (GC), which relies on either testing species boundaries in the genus
gene-gene phylogenetic concordance, or Pituophis. Although Reichling philosoph-
gene-geographical concordance among co- ically adopted the ESC and attempted to
distributed taxa, to define species (Avise define species on the basis of "indepen-
and Ball, 1990; Avise and Wollenberg, dent trajectories", he used multivariate
1997). Under both of these concepts, the phenotypic clusters as his best interpre-
category of species in bisexual populations tations of species, and we followed that
is likely to remain similar to that currently approach here (Fig. 2). Operationally we
employed by the BSC and RSC, and in- have used multivariate methods to sum-
trinsic reproductive barriers retain prima- marize morphological variation within
cy as a conceptual guide to species dis- and among populations, and then tried to
tinctions. Further, the subspecies category define species on the basis their diagnos-
(or some level of population distinctive- ability by at least two characters (i.e., con-
ness within a species) has a better empir- gruence). However, further comment is in
ical foundation; it will be defined as a order here because the overlap of values
group of populations phylogenetically dis- between many paired combinations of
tinguishable from, but reproductively some species of Micrurus makes species
compatible with, other such groups. The boundaries "fuzzy".
major limitations of either or both CSC Initial species designations were made
and GC principles are that: (1) subjective on the basis of head, neck, and chin col-
taxonomic judgements may still be re- oration characters, and triad patterns
quired; (2) obtaining phylogenies of alleles which seem to be "fixed" across sets of
for independent genes will be difficult (see populations, and in many cases the ma-
also Baum and Shaw, 1995; Doyle, 1995); jority of the snakes from these popula-
and (3) some species may be overlooked tions could also be diagnosed on the basis
as the phylogenetic separation and con- of triad number and/or meristic charac-
cordance may not be evident in the loci ters (ventral scale counts). However, for
sampled in some cases (i.e., very recent di- taxa represented by small sample sizes
vergence events; see details in Avise and (M. brasiliensis, M. diana, and M. tricolor,
Ball, 1990; Avise and Wollenberg, 1997). for example), sexual dimorphism often
Each of the above species concepts has appears to be extreme and/or the total
operational limitations, and these become range of some characters is huge and
especially obvious when the organisms of overlaps with virtually all other taxa (Ta-
interest are difficult to collect; this limits bles 1 and 2). The phenotypic cluster that
the number of museum vouchers avail- includes M. brasiliensis, M. diana, and M.
able for morphological study, increases frontalis, which separates along canonical
uncertainties about geographic distribu- variable 2 from all other species (Fig. 2),
tions, and increases the "waiting time" for reveals that these three overlap extensive-
live specimens from which molecular ly, but two of these are represented by
markers can be collected. All of these is- small sample sizes. Future collecting
sues are the norm for secretive snakes could reveal intergrades between these
such as most species of Micrurus, but samples, and eventually they may be in-
when coupled with the remoteness of terpreted as conspecific. Biochemical
much of northern South America (which (isozymes) and molecular (mtDNA se-
until recently has severely limited field quence) data that will be presented else-
work), and the morphological variability where (Jorge da Silva and Sites, in review)
(in color pattern) and complex taxonomic can clarify some of our interpretations,
history mentioned earlier, they combine and in this context, the following obser-
to present a nearly intractable "species vations are relevant: (1) M. pyrrhocryptus
boundary" problem by any single criteri- is "fixed" for a different allele at the N-
on. We therefore have taken an opera- Acetyl-p-glucosaminidase locus relative to
184 HERPETOLOGICAL MONOGRAPHS
four other frontalis complex species (we
have no molecular data for M. diana or M.
tricolor); (2) M. baliocoryphus and M. pyr-
rhocryptus share an allele that is absent
from the other three frontalis complex
species at the Glucose-6-phosphate isom-
erase locus; (3) M. baliocoryphus is fixed
for a unique allele at the Purine nucleo-
side-phosphorylase locus, relative to the
other four species; and (4) M. frontalis and
M. pyrrhocryptus are both polymorphic
for the same unique allele at the Phospho-
glucomutase locus, relative to the other
three species. However, the extent to
which these data can corroborate our mor-
phologically-based interpretations of spe-
cies boundaries on the criterion of "con-
cordance" from independent characters
(Avise and Ball, 1990; see also Mallet,
1995) is unclear, because sample sizes and
geographic localities represented for the
molecular data are extremely limited for
all five species (n = 1 in most cases). Sim-
ilarly, our assessment of hemipenial and
cranial morphology reveals some charac-
ters in which unambiguous alternative
states are possibly fixed in some popula-
tions, but additional data must be collect-
ed from other triad species of Micrurus
before these can be considered diagnostic.
Therefore, our interpretations of species
boundaries for some taxa in the frontalis
complex must be considered tentative.
Acknowledgments. -We thank the following mu-
seum curators: Darrel Frost and Charles Myers
(AMNH), Ted Daeschler (ANSP), Carlos Grisolia
(CCG), Luiz Augusto da Costa Porto (CEPB), Blanca
B. Alvarez de Avanza (CHC), Ulysses S. Silveira
(COBAP), Frederico Achaval and Raul Maneyro
(CZVR), Gustavo J. Scrocchi (FML), Harold Voris
(FMNH), Giuseppe Puorto (IB), Anibal R. Melgarejo
(IVB), Gustavo R. Carrizo, Gustavo Cuturier, Jorge
Cranwell, and Jos6 M. Gallardo (MACN/CHINM),
Marcos Di Bernardo and Thales de Lema (MCP),
Jos6 P. Rosado (MCZ), Julio C6sar Moura Leite, Re-
nato Be6rnils, and S6rgio Augusto Abrahao Morato
(MHNCI), Jorge D. Williams (MLP), Aida L. Aquino
and Marta Motte (MNHNP), Ivan Ineich and An-
nemarie Ohler (MNHN), Ulysses Caramaschi
(MNRJ), Sergio I. Tiranti (MPHNLP), Paulo Emflio
Vanzolini (MZUSP), Lucindo G. Alvarez and Ingrid
M. Fernandez (NKR), and Gregory Schneider
(UMMZ); for specimen loans and/or permitting the
examination of material deposited in their institu-
tions. We are deeply indebted to FURNAS-Centrais
Eletricas S.A., Serra da Mesa Energia S.A., and Com-
[No. 13
[No.
panhia Hidroel6trica do Sao Francisco-CHESF for
field assistance and for granting collecting permission
on their properties in Goias and Alagoas. We are also
indebted to Ariane C. Almeida, Cristiane K. M. Ko-
lesnikovas, Denise L. C. Martins, Dionei Jos6 da Sil-
va, IErika P. Andriani, Jos6 A. Speroni, Luciana D.
Silva, Luciana L. Casais e Silva, Marisa M. T. Rocha,
Marizane A. Oliveira, Marizete A. Oliveira, Nilmar A.
Dourado, Nilson A. Oliveira, Patricia Q. Torres, Ron-
ilton S. Carlos, S6rgio C. Ferreira, Sylvia R. T. Car-
doso, and Twiggy C. A. Batista, for field work and/or
laboratory assistance. We thank Valdirene Nascimen-
to for the drawings of skulls and hemipenes. We also
acknowledge Gabriel S. Sugliano and Marcus A.
Buononato for preparations of skulls and hemipenes,
and Brian A. Maurer for help with statistics. The his-
torical accounts could not have been completed with-
out the unmatched knowledge and generous assis-
tance of Paulo E. Vanzolini. The reviews provided by
James R. Dixon, Bill Lamar, and Joe Slowinski were
useful in this revision, and the work was in part spon-
sored by CAPES 169/89-3 fellowship (Brazilian Min-
istry of Education), a D Elden Beck Award (BYU-
1994), and a Theodore Roosevelt Memorial Award
(AMNH-1994) granted to NJS, and a NSF Doctoral
Improvement Dissertation Award (DEB 94-12285)
granted to JWS, Jr. and NJS.
LITERATURE CITED
ABALOS,J. W. 1961. Distribuci6n y densidad de las
serpientes venenosas de Santiago del Estero. Acta
Krausi Cuaderno del Instituto Nacional de Micro-
biolgia Buenos Aires 3:67-70.
ABALOS,J. W, AND R. NADER. 1962. La coral Micru-
rus lemniscatus frontalis de Santiago del Estero.
Anales del Instituto Nacional de Microbiologia
(Buenos Aires) 1:83-94.
ABALOS,J. W, AND I. PIROSKY.1963. Venomous Ar-
gentine serpents, ophidism and snake antivenin.
Pp. 363-371. In H. L. Keegan and W. V. Mac-
Farlane (Eds.), Venomous and Poisonous Animals
and Nouxious Plants of the Pacific Region. Perga-
mon Press, Oxford.
ABALOS, J. W, E. C. BAEZ, AND R. NADER. 1964.
Serpientes de Santiago del Estero. Acta Zoologica
Lilloana 20:211-283.
ABALOS, J. W, AND E. BUCHER. 1970. Zoo-epide-
miologia del ofidismo en Santiago del Estero. Bole-
tin de la Academia Nacional de Ciencias (Cordoba)
47:259-272.
ABALOS,J. W, AND C. C. MISCHIS. 1975. Elenco sis-
tematico de los ofidios Argentinos. Boletin de la
Academia Nacional de Ciencias (Cordoba) 51:55-
76.
ACHAVAL, F. 1976. Reptiles. Pp. 26-29. In: Lista de
las Especies de Vertebrados del Uruguay (Rep-
tiles). A. Langguth (Ed.). Museo Nacional Historia
Natural Montevideo.
ACHAVAL, F. 1997. Actualizaci6n sistematica y sinoni-
mica de los reptiles del Uruguay com comentarios
y distribuci6n. M.S. Dissertation. Facultad de
Ciencias, Universidad de la Republica, Uruguay.
ACHAVAL, F., A. R. MELGAREJO, AND M. MENEGHEL.
1976. Viboras venenosas del Uruguay. Instituto de
199
19991 HERPETOLOGICAL
HPOGA MONOGRAPHS
O GP1
Investigaciones de Ciencias Biologicas Montevideo.
Pp. 1-6.
ALVAREZ,A. 1919. Flora y Fauna de la provincia de
Santiago del Estero, Sgo. del Estero.
AMARAL,A. 1925. South American snakes in the col-
lection of the United States national museum. Pro-
ceedings U.S. National Museum 67:1-30.
AMARAL,A. 1929. Lista remissiva dos ophidios da re-
giao neotropica. Memorias do Instituto Butantan 4:
127-271.
AMARAL,A. 1944. Nota sobre a ofiologia neotr6pica
e brasfilica. XI. Subespecies de Micrurus lemnisca-
tus (L) e suas afinidades com Micrurus frontalis
(DM & BIBR). Pap6is Avulsos do Departmento de
Zoologia 5:83-94.
AVISE,J. C., AND R. M. BALL,JR. 1990. Principles of
genealogical concordance in species concepts and
biological taxonomy, Survey Evolutionary Biology
7:45-67.
AVISE,J. C., AND K. WOLLENBERG.1997. Phyloge-
netics and the origin of species. Proceedings of the
National Academy of Sciences USA. 94:7748-7755.
BARRIO,A., AND M. E. MIRANDA. 1966. Estudio
comparativo morfol6gico e imunologico entre las
diferentes entidades del genero Micrurus Wagler
(Ophidia: Elapidae) de la Argentina. Mem6rias do
Instituto Butantan 33:869-880.
BAUM, D. A., AND K. L. SHAW. 1995. Genealogical
perspectives on the species problem. Pp. 289-303,
In P. C. Hoch and A. G. Stephenson (Eds.), Ex-
perimental and Molecular Approaches to Plant
Biosystematics. Missouri Botanical Garden, St.
Louis, MO.
BERG, C. 1898. Contribuciones al conocimiento de la
fauna erpetol6gica argentina. Anales del Museo
Nacional de Buenos Aires 6:1-35.
BOULENGER,G. A. 1896. Catalogue of the snakes in
the British Museum (Natural History), Vol. 3. Tay-
lor and Francis, London.
BOULENGER,G. A. 1902. List of the fishes, batrachi-
ans and reptiles collected by the late Mr. P. 0. Si-
mons in the provinces of Mendoza and Cordoba
(Argentina). Ann. Mag. Nat. Hist. 9:175-252.
CADLE, J. E., AND V. M. SARICH.1981. An immu-
nological assessment of the phylogenetic position
of New World coral snakes. Journal Zoology 195:
157-167.
CAMPBELL,J. A., ANDW. W. LAMAR.1989. The Ven-
omous Reptiles of Latin America. Cornell Univer-
sity Press, Ithaca, New York, U.S.A.
COPE, E. D. 1859. Catalogue of the venomous ser-
pents in the museum of the Academy of Natural
Sciences of Philadelphia, with notes on the fami-
lies, genera, and species. Proceedings of the Acad-
emy of Natural Sciences Philadelphia 1859:332-
347.
COPE, E. D. 1862. Catalogues of the reptiles obtained
during the exploration of the Parana, Paraguay,
Vermejo and Uruguay rivers by Capt. Thos. J. Page,
U. S. N., and those procured by Lieut. N. Michler,
U.S. Top. Eng., commander of the expedition con-
ducting the survey of the Atrato river. Proceedings
of the Academy of Natural Sciences Philadelphia
1862:346-359.
185
DAVIS, J. I., AND K. C. NIXON. 1992. Populations,
genetic variation, and the delimitation of phyloge-
netic species. Systematic Biology 41:421-435.
DICK, M. M. 1977. Stations of the Thayer Expedition
to Brazil. 1865-1866. Breviora 444:1-37.
DOWLING,H. G. 1951. A proposed standard system
of counting ventrals in snakes. British Journal Her-
petology 1:97-99.
DOWLING,H. G., AND J. M. SAVAGE.1960. A guide
to snake hemipenes: a survey of basic structure and
systematic characteristics. Scientific Contributions
New York Zoology Society 45:17-28.
DOYLE,J. J. 1995. The irrelevance of allele tree to-
pologies for species delimitation, and non-topolog-
ical alternative. Systematic Botany 20:574-588.
DUMERIL,A. M. C., G. BIBRON,AND A. DUMIERIL.
1854. Erp6tologie generale ou histoire naturelle
compl6te des reptilles. Pp. 1191-1231. Libraire
Encyclop6dique de Roret, Paris.
ESTESO, S. C. 1985. Ofidismo en la Republica Ar-
gentina. Editorial Arpon, C6rdoba.
GOLAY,P. 1985. Checklist and Keys to the Terrestrial
Proteroglyphs of the World. Gilbert Rey, Geneva,
Switzeland.
GREENE,H. W., AND R. W. MCDIARMID.1981. Coral
snake mimicry: does it occur? Science 213:1207-
1212.
GUfNTHER,A. 1858. Catalogue of the Colubrine
Snakes in the Collection of the British Museum.
London.
HOGE, A. R. 1956. Uma nova esp6cie de Micrurus
(Serp. Elap.) do Brasil. Memorias do Instituto Bu-
tantan 27:67-72.
HOGE, A. R., AND A. R. LANCINI.1959. Nota sobre
Micrurus surinamensis nattereri Schmidt e Micru-
rus pyrrhocryptus Cope. Mem6rias do Instituto
Butantan 42/43:373-496.
HOGE, A. R., AND S. A. R. W. D. L. ROMANO.1979.
Sinopse das serpentes peconhentas do Brasil. Me-
morias do Instituto. Butantan 42/43:373-493.
JORGEDA SILVA,JR., N. 1993. The snakes of Samuel
hydroelectric power plant and vicinity, Rondonia,
Brazil. Herpetological Natural History 1:37-86.
JORGEDA SILVA,JR., N. 1995. Molecular systematics
and evolution of venoms in the South American
"triad" coral snakes (Micrurus, Elapidae). Ph.D.
Dissertation. Deparment of Zoology-Brigham
Young University, Provo, UT, U.S.A.
JORGEDA SILVA,JR., N., AND D. J. SILVA.1996. Mi-
crurus lemniscatus. Geographic Distribution. Her-
petological Review 27:34.
LEVITON,A. E., R. H. GIBBS, JR., E. HEAL, AND C.
E. DAWSON. 1985. Standards in herpetology and
ichthyology: Part 1. Standard symbolic codes for
institucional resource collections in herpetology
and ichthyology. Copeia 1985:802-832.
LYDDEKKER,R. 1890. Catalogue of the Fossil Rep-
tilia and Amphibia in the British Museum. Part IV.
Pp. 125.
MALLET,J. 1995. A species definition for the modern
synthesis. Trends in Ecology and Evolution 10:294-
299.
MANLY,B. F. J. 1991. Multivariate Statistical Meth-
ods: a Primer. Chapman and Hall, London.
186 HERPETOLOGICAL MONOGRAPHS
MERREM, B. 1820. Versuch eines Systems der Am-
phibien. Johan Christian Krieger, Marbug.
MOCQUARD, F. 1887. Sur une nouvelle espece
d'Elaps, E. heterochillus. Bull. Soc. Philom. Paris,
s6r. 7.11:39-41.
PESANTES, 0. 1994. A method for preparing hemi-
penis of preserved snakes. Journal of Herpetology
28:93-95.
REICHLING, S. B. 1995. The taxonomic status of the
Louisiana pine snake (Pituophis melanoleucus ruth-
veni) and its relevance to the evolutionary species
concept. Journal of Herpetology 29:186-198.
RINGUELET,R. A., AND R. H. ARAMBURU.1957.
Enumeracion sistematica de los vertebrados de la
provincia de Buenos Aires. Ministerio para Asuntos
Agrarios Buenos Aires, publ. no. 119.
ROUX-ESTEVE,R. 1983. Les specimes-types du genre
Micrurus (Elapidae) conserv6s au Mus6um Nation-
al D'Histoire Naturelle de Paris. Mem6rias do In-
stituto Butantan 46:79-94.
ROZE, J. A. 1967. A checklist of the New World ven-
omous coral snakes (Elapidae) with descriptions of
new forms. American Museum Novitates 2287:1-
60.
ROZE, J. A. 1970. Micrurus. Pp. 196-220. In J. A.
Peters and B. Orejas Miranda (Eds.), Catalogue of
the Neotropical Squamata. U. S. National Museum
Bulletin 297.
ROZE, J. A. 1983. New World coral snakes (Elapidae):
a taxonomic and biological summary. Memorias do
Instituto Butantan 46:305-338.
ROZE, J. A. 1987. Summary of coral snakes (Elapidae)
from Cerro de la Neblina, Venezuela, with descrip-
tion of a new subspecies. Revue francaise de
Aquariologie 14:109-112.
ROZE, J. A. 1989. New species and subspecies of coral
snake, genus Micrurus (Elapidae), with notes on
type specimens of several species. American Mu-
seum Novitates 2932:1-15.
ROZE, J. A. 1994. Notes on taxonomy of venomous
coral snakes (Elapidae) of South America. Bulletin
Maryland Herpetological Society 30:177-185.
ROZE, J. A. 1996. Coral Snakes of the Americas-
Biology, Identification, and Venoms. Krieger Pub-
lishing Company. Malabar, Florida, U.S.A.
ROZE, J. A., AND A. BERNAL-CARLO. 1987. Las ser-
pientes corales venenosas del g6nero Leptomicru-
rus (Serpentes, Elapidae) de Suramerica con des-
cripci6n de una nueva subespecie. Bolletin del
Museo regionale di Scienze Naturale di Torino 5:
573-608.
ROZE, J. A., AND N. JORGEDA SILVA, JR. 1990. Coral
snakes (Serpentes, Elapidae) from hydroelectric
power plant of Samuel, Rond6nia, Brazil, with de-
scription of a new subspecies. Bulletin Maryland
Herpetological Society 26:169-176.
SAS INSTITUTE,INC. 1990. Base SAS software for
personal computers, version 6.04. SAS Institute,
Inc., Cary, North Carolina, U.S.A.
SAVAGE, J. M. 1997. On the terminology for the de-
scription of hemipenes of squamate reptiles. Her-
petology Journal 7:23-25.
SAVAGE,J. M., AND J. B. SLOWINSKI. 1992. The col-
ouration of the venomous coral snakes (family
[No.
[No. 13
Elapidae) and their mimics (families Aniliidae and
Colubridae). Biological Journal Linnean Society 45:
235-254.
SAVITZKY,A. H. 1978. The origin of the New World
proteroglyphous snakes and its bearing on the
study of venom delivery systems in snakes. Ph.D.
Dissertation, University of Kansas, Lawrence.
SCHMIDT, K. P. 1936. Preliminary account of the cor-
al snakes of South America. Field Museum Natural
History Publication, Zoology Series 20:189-203.
SCOTT, N. J., AND J. W. LOVETT.1975. A collection
of reptiles and amphibians from the Chaco of Par-
aguay. University Connecticut Occasional Papers,
Biological Science 2:257-266.
SCROCCHI, G. J. 1990. El g6nero Micrurus (Serpen-
tes: Elapidae) en la Republica Argentina. Bolletin
del Museo regionale di Scienze Torino 8:343-368.
SERIE, P. 1919. Notas sobre las serpientes venenosas
de la Argentina. Abstracts of the I1 Reunion Na-
cional de la Sociedad Argentina de Ciencias Na-
turales 418-420.
SHREVE, B. 1953. Notes on the races of Micrurus
frontalis (Dum6ril, Dum6ril, and Bibron). Breviora
16:1-6.
SITES,J. W, JR., AND K. A. CRANDALL.1997. Testing
species boundaries and biodiversity studies. Con-
servation Biology 11:1289-1297.
SLOWINSKI, J. B. 1995. A phylogenetic analysis of the
New World coral snakes (Elapidae: Leptomicrurus,
Micruroides, and Micrurus) based on allozymic and
morphological characters. Journal of Herpetology
29:325-338.
STANGERUP, H. 1984. The Road to Lagoa Santa.
Marion Boyars, New York, U.S.A.
STRUSSMANN,C., AND I. SAZIMA.1993. The snake
assemblage of the Pantanal at Pocone, western Bra-
zil: faunal composition and ecological summary.
Studies on Neotropical Fauna and Environment
28:157-168.
TEMPLETON, A. R. 1989. The meaning of species and
speciation: a genetic perspective. Pp. 3-27. In D.
Otte and J. A. Endler (Eds.), Speciation and Its
Consequences. Sinauer, Sunderland, Massachu-
setts, U.S.A.
TEMPLETON,A. R. 1994. The role of molecular ge-
netics in speciation studies. Pp. 455-477. In B.
Schierwater, B. Streit, G. P. Wagner, and R. De
Salle (Eds.), Molecular Ecology and Evolution: Ap-
proaches and Applications. Birkhauser Verlag, Bas-
el, Switzerland.
VANZOLINI, P. E. 1953. On the type locality of some
Brazilian reptiles and amphibians collected by H.
H. Smith and described by E. D. Cope. Copeia
1953:124-125.
WIED-NEUWIED, M. 1820. Ueber die Cobra Coral
oder Cobra Coraes der Brasilianer. Nova Acta
Acad. Caesar. Leop. Carol., Halle 10:105-110.
APPENDIX
Specimens Examined
Micrurus frontalis: BRAZIL (620). Distrito Fed-
eral (4): IB 20541 (M), IB 20542 (M); IB 20543 (F);
IB 33895 (M)-Brasilia. Espirito Santo (3): IB 8808
(M)-Baixo Guandu; IB 32714 (M)-Fundao; IB
1999] HERPETOLOGICAL MONOGRAPHS
50684 (M)-Sao Gabriel da Palha. Goids (116):
CEPB 2076 (M)-Anapolis; CEPB 604 (M), CEPB
1968 (M)-Aparecida de Goiania; IB 44323 (M)-
Apor6; CEPB 614-Aragoiania; CEPB 1915 (M)-
Bela Vista de Goias; CEPB 1873 (M)-Cacu; IB
55133 (M)-Caiap6nia; IB 46422 (M)-Campo Ale-
gre de Goias; IB 53848 (M)-Corumbd de Goias; IB
32187 (M)-Catalao; CEPB 446 (M)-Goianapolis;
CEPB 770 (F), CEPB 1657 (M), CEPB 1874 (M),
CEPB 2432 (F), CEPB 2433 (M), IB 5114 (M), IB
5277 (M), IB 6874 (M), IB 45519 (M)-Goiania;
CEPB 345 (M)-Goias; CEPB 1763 (M), CEPB
2140 (M)-Guap6; CEPB 1587 (M)-Hidrolandia;
IB 51595 (M)-Ipameri; CEPB 304 (M)-Itumbiara;
CEPB 1656 (M), CEPB 1658 (F), CEPB 2284 (F),
CEPB 7404 (F)-Jataf; CEPB 336 (F), IB 48813 (F),
IB 48951 (M)-Niquelandia; IB 46337 (M), IB
46570 (M), IB 48059 (M)-Ouvidor; IB 40205 (M)-
Padre Bernardo; CEPB 1659 (M), CEPB 2285 (M),
CEPB 2363 (M), IB 26661 (M)-Pontalina; CEPB
1770 (M)-Portelandia; CEPB 1762 (M), CEPB
2365 (M)-Senador Canedo; CEPB 1813 (M)-Ser-
ranopolis; CEPBc s/n (F), CEPBc 11611 (F), CEPBc
13444 (F), CEPBc 13647 (F), CEPBc 14121 (M),
CEPBc 14149 (F), CEPBc 14174 (F), CEPBc 14180
(F), CEPBc 14260 (F), CEPBc 14268 (F), CEPBc
14269 (M), CEPBc 14279 (M), CEPBc 15817 (F),
CEPBc 16575 (M), CEPBc 16761 (F), CEPBc 17243
(F), CEPBc 18421 (M), CEPBc 18467 (F), CEPBc
19063 (F), CEPBc 19156 (M), CEPBc 19552 (F),
CEPBc 19618 (M), CEPBc 19787 (F), CEPBc 19942
(F), CEPBc 20448 (M), CEPBc 20666 (F), CEPBc
20667 (M), CEPBc 21265 (F), CEPBc 21490 (M),
CEPBc 21625 (M), CEPBc 21833 (F), CEPBc 22262
(M), CEPBc 22365 (F), CEPBc 22881 (F), CEPBc
23381 (M), CEPBc 23753 (M), CEPBc 23828 (M),
CEPBc 23698 (F), CEPBc 24137 (F), CEPBc 24180
(M), CEPBc 24380 (M), CEPBc 25309 (F), CEPBc
25530 (F), CEPBc 25875 (M), CEPBc 26796 (M),
CEPBc 27133 (F), CEPBc 27158 (F), CEPBc 27544
(F), CEPBc 27573 (F), CEPBc 27617 (F), CEPBc
27728 (M), CEPBc 27841 (F), CEPBc 29076 (M),
CEPBc 29183 (F), CEPBc 29487 (M), CEPBc 29584
(F), CEPBc 29905 (M), CEPBc 29913 (F), CEPBc
29958 (F), CEPBc 30644 (M), CEPBc 30712 (M),
CEPBc 31131 (F), CEPBc 31460 (M), CEPBc 31569
(M), CEPBc 31619 (F), CEPBc 31670 (M), CEPBc
32338 (F), CEPBc 35018 (F), CEPBc 36862 (M),
CEPBc 47275 (F)-Serra da Mesa. Mato Grosso
(5): IB 43217 (M)-Alto do Araguaia; IB 19950
(M)-Barra do GarVas;IB 44342 (M)-Chapada dos
Guimaraes; IB 17992 (M)-Lago Itaci (not plotted in
map); IB 7828 (F)-Tres Pontes. Mato Grosso do
Sul (51): IB 46761 (F)-Amambai; IB 42338 (M)-
Bandeirantes; IB 42426 (F)-Bataguassu; IB 29544
(M)-Bonito; IB 33345 (F) Brazilandia; IB 43814
(M), IB 50715 (M)-Caarap6; IB 28952 (M), IB
31152 (M), IB 33271 (F), IB 42844 (M), IB 44140
(M), IB 50376 (F), IB 51596 (M), IB 53091 (M),
COBAP 050 (F), COBAP 054 (F), COBAP 057 (M),
COBAP 418 (M), MZUSP 10186 (M), MZUSP 10187
(M)-Campo Grande; IB 33256 (M)-Casa Verde;
IB 43731 (M)-Cassilandia; IB 6751 (M), MZUSP
8640 (F)-Coxim; IB 5796 (M), IB 42134 (M), IB
187
42175 (M), IB 43225 (F), MHNCI 426 (M)-Dour-
ados; IB 32797 (M), IB 32935 (F), IB 44294 (M), IB
44512 (M)-Iguatemi; IB 17568 (F)-Itaum; IB
40196 (M)-Maracajui; IB 40491 (M)-Nioaque; IB
32353 (M), IB 46334 (M), IB 49170 (M)-Paranaiba;
IB 46800 (M), IB 16324 (M), IB 19421 (M), IB
21975 (M), IB 27528 (M), IB 28772 (M), IB 41368
(F), IB 41832 (M), IB 46341 (M)-Ponta Pora; IB
12926 (M)-Rio Brilhante; MZUSP 10185 (M)-
Terenos; IB 50350 (M)-Tres Lagoas. Minas Gerais
(121): IB 46347 (M)-Andrelandia; IB 951 (F)-Ar-
a9uai; IB 4819 (M), IB 6097 (M), IB 53092 (F)-
Araguari; IB 48959 (M)-Araxa; IB 50209 (M)-Ar-
cos; IB 42403 (M)-Barbacena; IB 31797 (M), IB
31799 (F)-Barao de Cocais; IB 42921 (M)-Belo
Horizonte; IB 40092 (F)-Borda da Mata; IB 17997
(M)-Botelho; IB 44155 (M), IB 44510 (M)-Bra-
sopolis; IB 42268 (M)-Camanducaia; IB 44489
(F)-Cambul; IB 8282 (M)-Cambuquira; IB 50137
(M)-Campo do Meio; IB 51540 (M)-Cascalho
Rico; IB 41389 (M)-Cassia; IB 16922 (F)-Congon-
has do Campo; IB 21735 (M)-Conselheiro Lafaiete;
IB 43214 (F)-Cristina; IB 32379 (M), IB 33075
(M), IB 33081 (M), IB 33318 (M), IB 33929 (M), IB
40161 (M)-Cruzflia; IB 32825 (M)-El6i Mendes;
IB 42074 (M), IB 42169 (M), IB 42901 (M), IB
44515 (M), IB 44541 (M)-Extrema; IB 33905 (M),
IB 41112 (F)-Formiga; IB 42172 (M)-Furnas; IB
53098 (M)-Guirinhata; IB 40091 (F), IB 41036
(F)-Ibia; IB 40650 (M)-Itabirito; IB 4655 (M), IB
8984 (F)-Itanhandu; IB 5914 (M)-Itajuba;
MZUSP 8060 (F)-Itamb6 do Mato Dentro; IB
53331 (M)-Itutinga; IB 54839 (M), IB 55310 (M)-
Jacutinga; IB 22736 (F), IB 31796 (M), IB 32274
(M), IB 33230 (M), IB 33237 (F), IB 33334 (F), IB
37382 (F), IB 37542 (M), IB 40144 (M), IB 40169
(M), IB 40184 (M), IB 40204 (F), IB 40409 (M), IB
40468 (M), IB 42294 (F), IB 45924 (F)-Juiz de
Fora; IB 53162 (F)-Lavras; IB 22101 (M)-Macha-
do; IB 4643 (M), IB 5172 (M)-Mariana; IB 34270
(M), IB 53090 (M)-Monte Carmelo; IB 460 (F), IB
19508 (M)-Monte Santo de Minas; IB 54315 (M)-
Monte Siao; IB 48040 (M)-Monte Verde; IB 8870
(M), IB 12235 (M), IB 15510 (F), IB 15562 (M), IB
28956 (F), IB 32193 (M), IB 32819 (M)- Ouro Fino;
IB 851 (M)- Ouro Preto; IB 40158 (M), IB 47660
(M)-Paracatu; IB 32823 (F), IB 32977 (F)-Passos;
IB 50139 (M), IB 51607 (F)-Patrocfnio; IB 16146
(M)-Perd6es; IB 27884 (M)-Po9os de Caldas; IB
34389 (F)-Pouso Alegre; IB 42366 (F), IB 42938
(F)-Prata; IB 2123 (M)-Santa Barbara; IB 18397
(M)-Santa Rita do Sapucaf; IB 49031 (M)-Santa
Rosa da Serra; IB 19487 (M), IB 29735 (F), IB 33094
(M), IB 32295 (F), IB 34334 (M)-Sao Sebastiao do
Paraiso; IB 47412 (M), IB 50714 (F), IB 53078 (F),
IB 53083 (M)-Tres Cora6ces; IB 42319 (M)-Tres
Pontas; IB 4939 (M), IB 46758 (M)-Tupaciguara;
IB 1288 (M), IB 30213 (M), IB 42463 (F)-Uberaba;
IB 42995 (M), IB 43931 (M), IB 53243 (M), MHNCI
360 (F)-Uberlandia; IB 606 (M); IB 24932 (M)-
Varginha; IB 43017 (M), IB 49100 (M)-Vicosa. Sao
Paulo (320): IB 43203 (M)-Aguaf; IB 51317 (M)-
Agua Branca; IB 3196 (F)-Aguas de Lind6ia; IB
46342 (M)-Aguas de Santa Barbara; IB 548 (M), IB
188 HERPETOLOGICAL MONOGRAPHS
4492 (M), IB 16955 (M), IB 21667 (M), IB 26723
(M), IB 31863 (M), IB 32301 (M), IB 40159 (M), IB
40201 (M), IB 53096 (M)-Agudos; IB 22038 (M),
IB 32423 (M), IB 33444 (M), IB 49229 (M)-Altin-
6polis; IB 24880 (M), IB 41072 (M), IB 41226 (M),
IB 41319 (M), IB 41321 (F), IB 42728 (M), IB 46093
(M), IB 53570 (M), IB 53620 (M), IB 53691 (M), IB
54632 (M)-Amparo; IB 7722 (F)-Aracatuba; IB
40366 (M)-Ara9oiaba da Serra; IB 24141 (M), IB
28571 (M)-Araraquara; IB 27453 (M), IB 37524
(M), IB 43786 (M), IB 44514 (M), IB 46760 (M), IB
49165 (F), IB 50133 (M), IB 50326 (M), IB 50726
(M), IB 52161 (M)-Assis; IB 42316 (M)-Atibaia;
IB 410 (M), IB 1741 (M)-Batatais; IB 7483 (M), IB
30233 (M), IB 40140 (M), IB 40896 (M), IB 41405
(M)-Bauru; IB 47356 (M)-Boa Esperan9a do Sul;
IB 21973 (M), IB 40215 (M), IB 41125 (M); IB
45912 (M)-Boituva; IB 7344 (M), IB 47649 (M)-
Botucatu; IB 32300 (M), IB 32581 (F), IB 40403 (M),
IB 40450 (M), IB 42118 (M), IB 42332 (M), IB
42340 (M), IB 42662 (M), IB 42720 (M), IB 42930
(M), IB 44417 (M), IB 44516 (M), IB 45626 (M), IB
47652 (M), IB 48810 (F), lB 50131 (M), IB 50135
(M), IB 51681 (M), IB 51853 (M), IB 54108 (M)-
BraganVa Paulista; IB 42379 (M), IB 42391 (M), IB
42895 (M), IB 45497 (M), IB 50501 (M)-Brotas; IB
43491 (M), IB 53714 (M)-Buri; IB 46678 (M)-
Cagapava; IB 33888 (M)-Caieiras; IB 23245 (M), IB
26982 (M), IB 29610 (M), IB 42106 (M), IB 42387
(M), IB 42637 (M), IB 42990 (M), IB 50675 (M)-
Campos do Jordao; IB 46099 (M)-Campos Novos
Paulista; IB 4841 (M)-Capao Bonito; IB 9682
(M)-Casa Branca; IB 7750 (M)-Cerrado (Sao Si-
mao County); IB 1302 (M), IB 6975 (M), IB 32971
(M)-Cerqueira Cesar; IB 33054 (M), IB 46387 (M),
IB 53084 (M)-Descalvado; IB 50953 (M)-Duarti-
na; IB 1348 (M)-Espirito Santo do Pinhal; IB 50970
(M), IB 51591 (M)-Fernand6polis; IB 2586 (M), IB
24303 (M), IB 29304 (M), IB 30012 (M), IB 50646
(M)-Franca; IB 1637 (M), IB 4815 (M)-Ibate; IB
36905 (M)-13 km N of Ilha Solteira reservoir; IB
29092 (M), 30677 (M), IB 32022 (M), IB 33885 (M),
IB 34441 (M), IB 42428 (M), IB 42461 (M); IB
42980 (M), IB 43657 (M), IB 44080 (M), IB 44313
(M), IB 44352 (M), IB 46356 (M), IB 47639 (M), IB
53169 (M)-Ibiuna; IB 27660 (M)-Iep8; IB 42451
(M)-Itapecerica da Serra; IB 374 (M), IB 7756 (M),
IB 18894 (M), IB 43986 (M), IB 50882 (M), IB
53082 (M)-Itapetininga; IB 33074 (M), IB 33313
(M)-Itapeva; IB 19415 (M)-Itapira; IB 37364
(M)-Itarar6; IB 40306 (M), IB 53260 (M)-Itatinga;
IB 33532 (M), IB 40021 (M), IB 42356 (M), IB
42627 (M), IB 45644 (M), IB 47654 (M), IB 48304
(M), IB 49669 (M), IB 50060 (F), IB 50515 (M), IB
508770 (M), IB 50887 (M), IB 51153 (M), IB 51714
(M), IB 52056 (M), IB 52315 (M), IB 53168 (M), IB
53964 (M), IB 54712 (M), IB 55260 (M), IB 55324
(M), MHNCI 1221 (M)-Itu; IB 50129 (F) Jacaref;
IB 33436 (F)-Jaguariuna; IB 52055 (M), IB 54820
(M)-Jales; IB 41292 (M)-Jeriquara; IB 581 (M)-
Leme; IB 51315 (M)-Lins; IB 3381 (M)-Lobo; IB
50136 (M)-Luiz Ant6nio; IB 1630 (M), IB 32034
(F), IB 31387 (M), IB 32309 (M), IB 32506 (M), IB
33254 (M), IB 33266 (M), IB 40132 (M), IB 41161
[No.
[No. 13
(M), IB 42405 (M), IB 43114 (M), IB 43223 (M), IB
43609 (M), IB 43638 (M), IB 43904 (M), IB 43933
(M), IB 44165 (M), IB 44449 (F), IB 45521 (M), IB
46683 (M), IB 47630 (M), IB 50377 (M), IB 53086
(M)-Mairinque; IB 6552 (M)-Mandurf; IB 32399
(F)-Marflia; LB 53592 (M)-Matao; IB 4778 (M)-
Mirassol; IB 27545 (M), IB 29967 (M), IB 31496
(M)-Mogi-Gua,u; IB 29736 (M)-Monte Apraziv-
el; IB 46688 (M)-Nipoa; IB 13152 (M)-Nova
Granada; IB 40191 (M)-Orlandia; IB 1338 (M), IB
5822 (M), IB 7516 (M), IB 40391 (M), IB 54977
(M)-Paraguacu Paulista; IB 8467 (M), IB 50910
(M)-Penaopolis; IB 50128 (F)-Piedade; IB 34139
(M), IB 40187 (M), IB 53085 (M)-Pilar do Sul; IB
7155 (M), IB 29741 (M), IB 32870 (M), IB 42275
(M)-Pindamonhangaba; IB 28925 (M)-Piquete; IB
44494 (M)-Piracaia; IB 30167 (M)-Piraj6; IB
45968 (M), IB 51600 (M)-Porto Feliz; IB 6266
(M)-Presidente Prudente; IB 32402 (M)-Presi-
dente Wenceslau; IB 24137 (M), IB 29964 (M), IB
40163 (M), IB 40259 (M), IB 41284 (M), IB 43006
(M)-Quata; IB 26846 (M), IB 45910 (M)-Ran-
charia; IB 43785 (M), IB 51594 (M)-Ribeirao Preto;
IB 42380 (M)-Rincao; IB 3173 (M), IB 30298 (M),
IB 34413 (M), IB 43906 (M)-Rio Claro; IB 51132
(M), IB 51592 (M)-Salto de Pirapora; IB 42326
(M)-Santa F6 do Sul; IB 51154 (M)-Santana de
Parnafba; IB 20791 (M)-Santo Anastacio do Pinhal;
IB 45642 (M), IB 45908 (M)-Sao Carlos; IB 4534
(M)-Sao Jos6 do Rio Pardo; IB 30974 (M)-Sao
Jos6 do Rio Preto; IB 43801 (M), IB 44369 (M), IB
51801 (M)-Sao Miguel Arcanjo; IB 8822 (M), IB
42390 (M)-Sao Paulo; IB 9321 (M)-Sao Pedro; IB
40165 (M), IB 41783 (M), IB 51608 (M)-Sao
Roque; IB 7484 (F), IB 7606 (M), IB 24149 (M), IB
46572 (M), IB 48049 (M)-Sao Simao; IB 50686
(M)-Sarapuf; IB 7803 (M), IB 34402 (M), IB 37436
(M), IB 46797 (M), IB 49451 (M)-Serra Negra; IB
8974 (M), IB 45978 (M)-Socorro; IB 6095 (M), IB
6125 (M), IB 16789 (M), IB 22228 (M), IB 23049
(M), IB 33490 (M), IB 33517 (F), IB 33518 (M), IB
40151 (M), IB 40154 (M), IB 40253 (M), IB 42674
(M), IB 43748 (M), IB 44481 (M), IB 45439 (M), IB
45957 (M), IB 45959 (M), IB 46034 (M), IB 46335
(M), IB 46340 (M), IB 46357 (M), IB 50045 (M), IB
50134 (M), IB 51680 (M), IB 53095 (M)-Sorocaba;
IB 4918 (M), IB 33483 (M)-Tamba6; IB 50986
(M)-Tapirf; IB 43934 (M)-Tatui; IB 44435 (M)-
Taubat6; IB 32976 (M)-Valentim Gentil; IB 42170
(M), IB 42220 (M), IB 42305 (M), IB 43932 (M), IB
50711 (M), IB 53171 (M), IB 53745 (M)-Votoran-
tim; IB 53079 (M), IB 54619 (M) Votuporanga; IB
37277 (M), IB 37479 (M)-Xavantes. PARAGUAY
(13). Departamento Amambay (1): MNHNP
2685-P. N. Cerro Cora; Departamento Central
(9): IB 10110 (M)-Bafa de Asunci6n; AMNH 76575
(M), AMNH 77028 (M), AMNH 78988 (M), AMNH
78989 (M), MNHNP 5142 (F)-Asunci6n; MNHNP
2689-Nueva Italia; MNHNP 5141 (M)-San Lor-
enzo; MNHNP 5139 (M)-Aregua (Lago Ypacarai);
Departamento Cordillera (1) MNHNP 3405-Ita-
curubi; Departamento Paraguari (1): MNHNP
2600-Acahay; Departamento San Pedro (1):
MNHNP 2599-Lima.
1999] HERPETOLOGICAL MONOGRAPHS
Micrurus altirostris: ARGENTINA (84). Corri-
entes (6): MACN n/d (M), IB 55595 (M), IB 55596
(M), IB 55597 (M)-Ituzaig6; CHINM 2005 (M)-
Santo Tom6; CHINM 3537 (M)-Torrent. Entre
Rios (7): CHINM 2861 (M), CHINM 3379 (M),
CHINM 3571 (M)-Conc6rdia; CHINM 1710
(M)-Chajari; CHINM 3633 (M)-Federaci6n;
MACN 087 (M), MACN 1117 (M)-Santa Elena.
Misiones (71): CHINM 2243 (F)-Ap6stoles; FML
974 (M)-Arroyo Fortaleza, Ruta 21, 20 km from San
Pedro; CHINM 3002 (M), CHINM 3003 (M)-Ca-
piovi; CHINM 734 (M)-Corpus; CHINM 1361 (F),
CHINM 2884 (M), CHINM 2890 (M), CHINM
2930 (M), CHINM 2607 (M), CHINM 2644 (M)-
Eldorado; MACN 35077 (M), MACN 35078 (M),
MACN 35302 (F)-Guarany, San Vicente (not plot-
ted in map); CHINM 2963 (F) Jardin America;
CHINM 729 (M), CHINM 1980 (M), CHINM 1985
(M), CHINM 2695 (M)-Leandro N. Alem; CHINM
2301 (M), MACN 35348 (M)-Liberdad; CHINM
1941 (M), CHINM 1942 (M), CHINM 1946 (M),
CHINM 1953 (M), CHINM 1992 (M), CHINM
1993 (M), CHINM 2307 (M), CHINM 2610 (F),
CHINM 2613 (M), CHINM 2668 (M), CHINM
2885 (M), FML 1985 (M) Obera; CHINM 732 (F),
CHINM 733 (F) Picada San Javier; FML 609
(M)-Picada Vieja, Obera; CCG 007 (M) = MZUSP
10810, CCG 010 (M) = MZUSP 10811, CCG 011
(F) = MZUSP 10813, CHINM 1605 (M), CHINM
1689 (F), CHINM 1923 (M), CHINM 2167 (M)-
Posadas; CHINM 1928 (M)-Puerto Azara; MACN
3115- (M), MACN 3115b (M), MACN 3115c (F),
MACN 3115d (M), MACN 3115e (M), MACN 3115f
(M), MACN 3115g (F), MACN 3115h (M), MACN
3157 (F)-30 km from Puerto Bemberg (not plotted
in map); CHINM 727 (M), CHINM 1510 (M),
CHINM 1538 (M), CHINM 1539 (M), CHINM
2166 (M)-Puerto Esperanza; CHINM 2217 (M)-
Puerto Leone; CHINM 1847 (M), CHINM 1920
(M), CHINM 2400 (M)-Puerto Mineral; CHINM
1802 (M)-Puerto Paranaf; FML 777 (M), CHINM
2961 (M), CHINM 2962 (F)-Puerto Piray; MACN
3191 (F)-Rfo Uruana (not plotted in map); MACN
014A (M), MACN 014B (M)-San Ignacio; CHINM
1893 (M)-San Javier. BRAZIL (960). Parana
(409): MHNCI 1630 (M), MHNCI 2330 (M),
MHNCI 2331 (M)-Agudos do Sul; IB 6503 (M),
MHNCI 6503 (M), MHNCI 7020 (M), MHNCI
7628 (F)-Almirante Tamandare; MHNCI 2962
(M)-Ant6nio Olinto; IB 53087 (M), MHNCI 263
(F), MHNCI 4347 (M)-Apucarana; MHNCI 4112
(M)-Arapotf; IB 4644 (M), IB 5115 (M), IB 5116
(M), IB 5117 (M), IB 5931 (M), IB 6515 (M), IB
6783 (M), IB 7050 (F), IB 7132 (F), IB 7296 (M),
IB 7342 (M), IB 8167 (F), IB 9578 (F), IB 14352
(M), IB 15730 (M), IB 15769 (M), IB 15797 (F), IB
15992 (M), IB 16058 (M), IB 16600 (M), IB 16451
(F), IB 16884 (M), IB 16921 (F), IB 19004 (M), IB
22230 (M), IB 27103 (M), IB 29316 (M), IB 29908
(M), IB 31779 (F), IB 31905 (M), IB 33507 (M), IB
33632 (M), IB 34398 (M), IB 34427 (M), IB 37259
(M), IB 37262 (M), IB 37365 (M), IB 37506 (M), IB
40649 (M), IB 42206 (M), IB 43810 (F), IB 44134
(F), IB 44220 (F)-Araucaria; IB 4644 (M), IB 6795
189
(M), IB 7802 (M), IB 7326 (M), IB 22321 (M), IB
23978 (M), IB 31905 (M), MHNCI 2531 (M)-Balsa
Nova; MHNCI 1199 (M), MHNCI 1692 (M),
MHNCI 1753 (F), MHNCI 3840 (M)-Bocaiuiva do
Sul; IB 55210 (F)-Cambara; MHNCI 234 (M),
MHNCI 1709 (M), MHNCI 2846 (M), MHNCI
2861 (F), MHNCI 3855 (M)-Campina Grande do
Sul; IB 5930 (F), IB 16096 (M), IB 30007 (M),
MHNCI 232 (M), MHNCI 2127 (M)-Campo do
Tenente; IB 5957 (M), IB 7789 (M), IB 41230 (M),
IB 45727 (F), MHNCI 2663 (M), MHNCI 3103 (M),
MHNCI 3109 (F), MHNCI 4731 (F), MHNCI 4845
(M), MHNCI 4846 (M), MHNCI 4847 (F), MHNCI
4848 (F), MHNCI 6186 (M)-Campo Largo;
MHNCI 3235 (F)-Candido de Abreu; MHNCI 783
(M)-Cantagalo; CEPB 2313 (M), IB 42218 (F), IB
42499 (M), MHNCI 4389 (M), MHNCI 4884 (M),
MHNCI 6255 (M)-Cascavel; MHNCI 1724 (F)-
Castro; IB 40898 (M)-Serro Azul; MHNCI 3338
(F)-Chopinzinho; MHNCI 1367 (F), MHNCI 1781
(F), MHNCI 3180 (M), MHNCI 3988 (M), MHNCI
3989 (F), MHNCI 4991 (F)-Colombo; MHNCI
2974 (M), MHNCI 7399 (M)-Contenda; IB 1600
(M), IB 4818 (M), IB 5966 (M), IB 1764 (M), IB
7714 (M), IB 16291 (F), IB 24699 (M), IB 24700
(M), IB 24701 (M), IB 28323 (M), IB 28415 (M), IB
28416 (M), IB 29050 (M), IB 29069 (F), IB 29178
(M), IB 31807 (M), IB 31822 (M), IB 31824 (M), IB
41038 (F), IB 42574 (F), IB 42989 (M), IB 43204
(F), IB 43644 (F), IB 43796 (M), IB 45838 (M), IB
46802 (F), IB 48223 (M), IB 48228 (M), IB 50022
(M), IB 50576 (M), IB 50695 (M), IB 50713 (F), IB
50954 (M), IB 50969 (M), IB 50971 (M), IB 50989
(M), IB 51579 (M), IB 51597 (M), IB 51598 (M), IB
52054 (M), IB 52057 (M), IB 52059 (F), IB 52060
(M), IB 52061 (M), IB 52386 (M), IB 53257 (M), IB
54804 (M), MHNCI 225 (M), MHNCI 242 (M),
MHNCI 517 (F), MHNCI 1046 (M), MHNCI 1348
(M), MHNCI 1595 (F), MHNCI 1795 (F), MHNCI
2976 (M), MHNCI 3667 (M), MHNCI 3747 (F),
MHNCI 6113 (F), MHNCI 6158 (F)-Curitiba;
MHNCI 4169 (F)-Dois Vizinhos; MHNCI 1061
(F)-Eneas Marques; IB 31806 (M), MHNCI 146
(M), MHNCI 542 (M), MHNCI 928 (M), MHNCI
948 (M), MHNCI 962 (M), MHNCI 1060 (M),
MHNCI 1408 (M), MHNCI 1440 (M), MHNCI
1441 (F), MHNCI 1443 (M), MHNCI 1532 (M),
MHNCI 1621 (F), MHNCI 1622 (M), MHNCI 1777
(M), MHNCI 2198 (M), MHNCI 2393 (M), MHNCI
2907 (M), MHNCI 3748 (F)-Francisco Beltrao; IB
15765 (F), IB 16828 (M), IB 29142 (M), IB 33105
(M), MHNCI 147 (F), MHNCI 949 (M), MHNCI
3301 (F), MHNCI 3202 (M), MHNCI 3303 (M),
MHNCI 3716 (M), MHNCI 5138 (M)-Guarapuava;
IB 7694 (M), MHNCI 2204 (M)-Imbituva;
MHNCI 1162 (F), MHNCI 1301 (M), MHNCI 1302
(F), MHNCI 1303 (M)-Ipiranga; CEPB 2308 (M),
IB 7156 (M), IB 16144 (M), IB 17302 (M), IB 30519
(M), IB 31212 (M), IB 31340 (M), IB 46338 (M), IB
46111 (M), MHNCI 1045 (M), MHNCI 1258 (M),
MHNCI 1337 (M), MHNCI 1613 (M), MHNCI
2499 (F), MHNCI 3749 (M), MHNCI 4208 (F)-
Irati; IB 46253 (F), MHNCI 3352 (M)-Jaguarialva;
IB 9073 (M), IB 40200 (M), MHNCI 1460 (M),
190 HERPETOLOGICAL MONOGRAPHS
MHNCI 1492 (M), MHNCI 1785 (F), MHNCI 2966
(F)-Lapa; IB 52058 (M)-Londrina; MHNCI 1442
(F), MHNCI 1693 (F), MHNCI 1799 (F), MHNCI
4160 (M)-Mandirutuba; MHNCI 5096 (M),
MHNCI 5103 (M), MHNCI 5111 (F)-Mangueirin-
ha; MHNCI 1368 (M)-Marmeleiro; IB 48157
(M)-Maua da Serra; MHNCI 230 (F), MHNCI
1274 (M), MHNCI 1634 (F)-Morretes; MHNCI
1472 (M), MHNCI 1533 (M), MHNCI 3559 (M)-
Ortigueiras; MHNCI 5068 (F)-Palmas; MHNCI
6156 (M)-Palmeira; MHNCI 1422 (M), MHNCI
1467 (F), MHNCI 1488 (M), MHNCI 1701 (F),
MHNCI 2023 (F), MHNCI 2030 (F), MHNCI 2031
(M), MHNCI 2171 (F), MHNCI 2206 (F), MHNCI
2662 (F), MHNCI 2782 (M), MHNCI 4168 (M)-
Pato Branco; IB 33269 (M), IB 37547 (M)-Pien;
MHNCI 4204 (M)-Pinhais; (not plotted in map); IB
43119 (M), MHNCI 5014 (M), MHNCI 5030 (M),
MHNCI 5041 (F), MHNCI 5042 (M), MHNCI 5043
(M), MHNCI 5082 (F), MHNCI 5086 (F), MHNCI
5118 (M), MHNCI 5132 (M), MHNCI 5142 (M),
MHNCI 5143 (F)-Pinhao; IB 9313 (M), IB 43000
(F)-Piraf do Sul; IB 7634 (F), IB 7698 (M), IB
18124 (M), IB 27088 (M), MHNCI 1112 (M),
MHNCI 2686 (M), MHNCI 2687 (F), MHNCI 2977
(M), MHNCI 2978 (M), MHNCI 3108 (F), MHNCI
3673 (F), MHNCI 3990 (M), MHNCI 4332 (F)-
Piraquara; IB 33435 (M), IB 33642 (M), IB 42328
(F), MHNCI 236 (M), MHNCI 927 (F), MHNCI
1263 (F), MHNCI 1428 (M), MHNCI 1629 (M),
MHNCI 1630 (M), MHNCI 1834 (M), MHNCI
1894 (F), MHNCI 2400 (M), MHNCI 3745 (M),
MHNCI 3750 (M)-Ponta Grossa; MHNCI 6172
(F)-Pranchita; IB 5733 (M), IB 5786 (M), IB 7859
(M), MHNCI 3319 (F)-Prudent6polis; MHNCI
235 (F), MHNCI 239 (F), MHNCI 1078 (F),
MHNCI 1369 (M), MHNCI 2561 (F), MHNCI 4872
(F), MHNCI 6103 (M), MHNCI 6104 (F)-Quatro
Barras; MHNCI 3374 (M)-Realeza; MHNCI 53088
(M)-RenascenVa; MHNCI 3224 (F), MHNCI 3225
(M), MHNCI 3227 (M)-Reserva; IB 10359 (M), IB
44289 (M), MHNCI 2092 (F), MHNCI 2354 (M)-
Rio Branco do Sul; IB 49376 (M)-Sao Jer6nimo da
Serra; IB 24192 (M), IB 24193 (M), IB 24259 (M),
IB 26884 (M), IB 26885 (F), IB 26886 (M), IB 31231
(M), IB 31424 (M), IB 32884 (M), IB 33073 (F), IB
31101 (F), IB 33526 (M), 33637 (M), IB 40145 (M),
IB 40405 (M), IB 43857 (F), IB 44622 (M), IB 46386
(F), MHNCI 226 (M), MHNCI 227 (M), MHNCI
228 (M), MHNCI 229 (F), MHNCI 233 (M),
MHNCI 306 (M), MHNCI 1524 (F), MHNCI 2018
(M), MHNCI 2975 (F), MHNCI 3746 (M), MHNCI
4946 (M), MHNCI 6147 (M), MHNCI 6241 (M)-
Sao Jose dos Pinhais; MHNCI 1006 (M)-Sao Ma-
teus do Sul; IB 43143 (M)-Santa Helena; IB 4777
(M), IB 8379 (M), IB 9514 (M), IB 28532 (M),
MHNCI 2532 (F), MHNCI 2533 (M), MHNCI 2534
(M), MHNCI 2535 (M)-Teixeira Soares; IB 27441
(M), IB 27820 (F), IB 28236 (F), IB 45313 (M), IB
45423 (M), IB 45466 (M), 45475 (F), IB 45480 (M),
IB 45508 (F), IB 45635 (M), IB 46030 (M), IB 46036
(M), IB 46565 (M), IB 47057 (M), IB 47656 (M), IB
46759 (M), IB 49683 (M), IB 50645 (M), IB 50716
(M), IB 53089 (M), IB 53093 (M), IB 53165 (M), IB
[No.
[No. 13
55464 (M), MHNCI 237 (M), MHNCI 1963 (M),
MHNCI 2448 (F), MHNCI 2845 (M), MHNCI 3156
(M), MHNCI 3455 (F), MHNCI 3493 (M), MHNCI
4161 (F), MHNCI 4238 (F), MHNCI 4239 (F)-
Telemaco Borba; MHNCI 1487 (F), MHNCI 3220
(M), MHNCI 3221 (M), MHNCI 6261 (M)-Tibaji;
MHNCI 956 (M)-Tijucas do Sul; MHNCI 231
(M)-Tomasina; IB 42372 (M)-Uniao da Vit6ria.
Rio Grande do Sul (441): IB 29969 (M), IB 31151
(M)-Alegrete; IB 30904 (M), IB 33166 (M), IB
43765 (M), MCP 2223 (F), MCP 4577 (M)-Bage;
MCP 1519 (M), MCP 1690 (F), MCP 1744 (F), MCP
3245 (M)-Barao do Triunfo; MCP 3969 (M), MCP
3870 (F), MCP 3959 (M), MCP 3965 (M), MCP
3970 (M), MCP 3977 (M), MCP 4089 (F), MCP
4138 (M), MCP 4141 (M)-Barra do Ribeiro; MCP
3194 (M)-Barracao; MCP 3594 (F), MCP 3595 (M),
MCP 3596 (M), MCP 3597 (M), MCP 3741 (M),
MCP 3767 (M), MCP 3787 (M), MCP 3846 (F),
MCP 3859 (M), MCP 3886 (M), MCP 3887 (M),
MCP 3888 (M), 4027 (F), MCP 4031 (M)-Butia;
MCP 3375 (M)-Cacapava do Sul; IB 52173 (M), IB
53450 (M)-Cacequi; IB 23816 (M), IB 28558 (M),
IB 29165 (M), IB 40284 (M)-Cachoeira do Sul; IB
44652 (M)-Cambara do Sul; MCP 1131 (F), MCP
1920 (M), MCP 1939 (F)-Campo Bom; MCP 2373
(M), MCP 4146 (M)-Canoas; IB 6072 (M), IB
16643 (M), IB 18689 (M), IB 23418 (F), IB 50699
(M), IB 50717 (M), IB 50875 (M), IB 50876 (M), IB
50878 (M), IB 50879 (M), IB 51056 (M), IB 51057
(M), IB 51058 (M), IB 51059 (M), IB 51060 (F), IB
51061 (M), IB 51163 (M), IB 51580 (F), IB 51601
(F), IB 53081 (M)-Carazinho; IB 53080 (M)-Ca-
tuipe; IB 11122 (M), IB 11256 (M), IB 11432 (M),
IB 11614 (M), IB 15758 (M), IB 16604 (M), IB
18570 (M), IB 40108 (F), IB 41293 (M)-Cruz Alta;
MCP 3731 (M), MCP 4553 (M)-Eldorado do Sul;
MCP 1931 (M), MCP 1932 (F), MCP 1933 (M),
MCP 2662 (M)-Estancia Velha; IB 48234 (M)-Er-
echim; MCP 4334 (M), MCP 4474 (M)-Frederico
Westphalen; MCP 3072 (M)-Garruchos, Barreiro;
IB 40089 (M), IB 40141 (M)-Gaurama; MCP 2609
(M), MCP 2610 (M), MCP 4616 (M)-General Ca-
mara; MCP 1593 (M), MCP 1597 (M), MCP 1954
(M), MCP 2350 (M)-Glorinha; IB 49281 (M), IB
51271 (M), MCP 198 (M), MCP 1530 (M); MCP
1850 (M), MCP 1880 (M), MCP 2072 (F), MCP
4599 (M)-Gravatai; IB 32352 (M), IB 33003 (M),
MCP 1256 (M), MCP 1533 (F), MCP 2176 (M),
MCP 2191 (M), MCP 2240 (M), MCP 2244 (M),
MCP 2300 (M), MCP 2453 (M), MCP 2493 (M),
MCP 2738 (M), MCP 3775 (M)-Guafba; IB 33261
(M), IB 37339 (M), IB 41785 (F), IB 42320 (M), IB
42583 (F), IB 43728 (F), IB 53097 (M)-Harmonia;
MCP 4549 (M), MCP 4558 (M), MCP 4658 (M)-
Ibiruba; IB 6491 (M), IB 7846 (M), IB 7939 (M), IB
7954 (M), IB 7955 (M), IB 23621 (M), IB 40186 (M),
MCP 3385 (M), MCP 3386 (M), MCP 3387 (M),
MCP 3388 (M), MCP 3390 (F), MCP 3391 (M),
MCP 3392 (M), MCP 3393 (M)-Ijuf; IB 11615
(M)-Itapevi (not plotted in map); IB 34423 (F), IB
34425 (M)-Itatiba do Sul; IB 46229 (M)-Lageado;
IB 23668 (M)-Lagoa Vermelha; MCP 2825 (F)-
Marcelino Ramos; MCP 2485 (M), MCP 2736 (M),
1999]
1999] HERPETOLOGICALMONOGRAPHS
HERPETOLOGICAL MONOGRAPHS
MCP 2872 (M), MCP 3035 (M), MCP 3180 (M)-
Mariana Pimentel; IB 28245 (M), IB 31083 (M), IB
33246 (M), IB 37256 (M), IB 40142 (M), IB 40192
(M), IB 41209 (M), IB 45831 (M), MCP 4359 (M)-
Montenegro; MCP 4598 (M)-Nova Hartz; MCP
2226 (M)-Nova Santa Rita; IB 27300 (M), IB 33689
(M), IB 50246 (M)-Novo Hamburgo; IB 46086 (M),
MCP 1781 (F), MCP 5028 (M), MCP 5258 (M)-
Osorio; MCP 2534 (M), MCP 2535 (M)-Parobe;
MCP 3013 (F)-Passo Fundo; IB 1799 (M)-Pelota;
CEPB 2310 (M), IB 1251 (M), IB 6527 (M), IB 6818
(M), IB 7428 (M), IB 7649 (M), IB 7842 (M), IB
18690 (M), IB 20941 (M), IB 30241 (M), IB 32096
(M), IB 32105 (M), IB 32489 (M), IB 32639 (M), IB
32847 (M), IB 42199 (M), IB 42202 (F), IB 42204
(M), IB 42208 (F), IB 42325 (M), IB 44491 (F), IB
44640 (F), IB 45724 (M), IB 46336 (M), IB 49129
(M), IB 53244 (M), IB 53370 (M), MCP 178 (M),
MCP 179 (M), MCP 192 (M), MCP 940 (F), MCP
1098 (M), MCP 1159 (M), MCP 1171 (M), MCP
1223 (F), MCP 1244 (M), MCP 1310 (F), MCP 1706
(M), MCP 1727 (M), MCP 1851 (M), MCP 1994 (F),
MCP 1999 (M), MCP 2004 (M), MCP 2146 (M),
MCP 2187 (M), MCP 2228 (F), MCP 2229 (M),
MCP 2275 (M), MCP 2500 (M), MCP 2555 (M),
MCP 2744 (M), MCP 2806 (M), MCP 3042 (M),
MCP 3294 (M), MCP 3371 (M), MCP 3571 (F),
MCP 3650 (M), MCP 3726 (F), MCP 3763 (M),
MCP 3889 (M), MCP 3896 (M), MCP 3923 (M),
MCP 4190 (M), MCP 4195 (M), MCP 4222 (M),
MCP 5036 (M), MCP 5259 (M)-Porto Alegre; MCP
3296 (M)-Porto Xavier; IB 7176 (M), IB 7242 (M),
IB 27103 (M), IB 27750 (M)-Restinga Seca; IB
24143 (M)-Rondinha; IB 13640 (M), IB 32838
(M)-Rosario do Sul; MCP 3338 (M)-Sananduva;
IB 16055 (M)-Santa Barbara do Sul; IB 7358 (M),
IB 16140 (M)-Santa Rosa; IB 22301 (F), IB 22342
(M), IB 23837 (F), IB 26620 (F), IB 26621 (M), IB
26622 (M), IB 26623 (F), IB 26624 (F), IB 26625
(M), IB 26626 (M), IB 26627 (M)-Santana do Liv-
ramento; IB 8187 (F)-Santo Angelo; IB 22390 (M),
IB 33370 (F), IB 41793 (M), IB 42378 (M)-Sao
Borja; IB 42246 (M), MCP 1246 (M), MCP 1441
(M), MCP 2003 (M), MCP 2494 (M), MCP 2495 (F),
MCP 2496 (M), MCP 2497 (M), MCP 2783 (M),
MCP 3670 (M), MCP 3966 (M), MCP 3967 (M),
MCP 5585 (M), MCP 5586 (M)-Sao Jer6nimo; IB
5068 (M), IB 5214 (M), IB 5503 (M), IB 6734 (M),
IB 6794 (M), IB 6824 (M), IB 6924 (M), IB 6969
(M), IB 7407 (M), IB 7878 (M), IB 8131 (M), IB
8248 (M), IB 9087 (M)-Sao Leopoldo; IB 32444
(M)-Sao Paulo das Missoes; IB 51881 (M), IB
51915 (M), IB 52062 (F), IB 52383 (M), IB 53258
(M), MCP 2486 (M), MCP 4196 (M), MCP 4218
(M), MCP 4342 (M), MCP 4482 (M)-Sao Sebastiao
do Cal; IB 6348 (M), IB 7668 (M), MCP 1416 (F),
MCP 1439 (M)-Sertao; IB 52063 (M)-Tapes;
MCP 2720 (M)-Taquara; IB 52414 (M), MCP 1552
(M), MCP 1624 (M), MCP 1868 (F), MCP 1963 (M),
MCP 1952 (M), MCP 2169 (M), MCP 2291 (M),
MCP 2307 (M), MCP 2311 (F), MCP 2384 (M),
MCP 2591 (M), MCP 2726 (M), MCP 2766 (F),
MCP 2796 (F), MCP 3052 (M), MCP 3118 (F), MCP
3255 (M), MCP 3334 (M), MCP 3342 (M), MCP
191
3443 (M), MCP 3345 (F), MCP 3346 (M), MCP
3353 (M), MCP 3380 (M), MCP 3540 (F), MCP
3703 (M), MCP 3729 (F), MCP 3884 (M), MCP
3931 (M), MCP 3968 (M), MCP 4112 (M), MCP
4115 (M), MCP 4168 (M), MCP 4169 (M), MCP
4179 (M), MCP 4187 (M), MCP 4188 (M), MCP
4189 (M), MCP 4194 (M), MCP 4197 (M), MCP
4203 (M), MCP 4204 (F), MCP 4206 (M), MCP
4212 (M), MCP 4233 (M), MCP 4241 (M), MCP
4249 (M), MCP 4686 (M), MCP 4737 (M), MCP
5025 (M), MCP 5026 (M), MCP 5027 (M), MCP
5029 (M), MCP 5030 (M), MCP 5031 (M), MCP
5032 (M), MCP 5033 (M), MCP 5034 (M), MCP
5035 (M)-Torres; IB 54478 (M), IB 54788 (M)-
Tuparendi; IB 23602 (F), IB 6840 (M), IB 8171 (M),
IB 43110 (F), IB 43219 (M), IB 43224 (F), IB 43736
(M), IB 43795 (M), IB 43798 (M), IB 43812 (M), IB
43836 (M), IB 43843 (M), IB 43928 (M), IB 43929
(M), IB 43943 (M), IB 43946 (M), IB 44217 (M), IB
44347 (M), IB 48023 (M), IB 48141 (M), IB 50698
(F), IB 51582 (F), IB 53100 (F), MCP 53170 (M)-
Uruguaiana; MCP 2049 (M)-Vacaria; IB 46656 (M),
MCP 199 (M), MCP 200 (M), MCP 237 (M), MCP
240 (F), MCP 241 (F), MCP 245 (F), MCP 306 (M),
MCP 0307 (M), MCP 309 (F), MCP 1112 (M), MCP
1113 (M), MCP 1245 (F), MCP 1280 (F), MHNCI
1286 (M), MCP 1343 (M), MCP 1419 (M), MCP
1504 (M), MCP 1570 (F), MCP 1573 (M), MCP
1964 (M), MCP 2046 (M), MCP 2186 (M), MCP
2492 (M), MCP 2594 (M), MCP 2602 (M), MCP
3235 (F), MCP 3547 (M), MCP 3628 (M), MCP
3772 (M), MCP 3958 (M), MCP 4467 (F), MCP
4483 (M), MCP 4546 (M), MCP 4665 (M), MCP
4711 (M), MCP 5088 (M), MCP 5257 (M), MCP
5583 (M), MCP 5584 (M)-Viamao. Santa Catarina
(110): MCP 2980 (M), MCP 2982 (M)-Anita Gar-
ibaldi; MHNCI 3146 (M), MHNCI 3147 (M),
MHNCI 3148 (M)-Ararangua; IB 5284 (M)-Blu-
menau; IB 50587 (F)-Cagador, IB 29282 (M), IB
29476 (M), IB 29518 (M), 29764 (F)-Campo Er8;
IB 4664 (F), IB 4836 (M), IB 40097 (M)-Capinzal;
MHNCI 4083 (M), MHNCI 6299 (M), MHNCI
6300 (M), MHNCI 6301 (M), MHNCI 6302 (M),
MHNCI 6306 (F)-Catanduvas; IB 47055 (M)-Flo-
rianopolis; IB 53883 (M)-Fraiburgo; MHNCI 5883
(F)-Herval D'Oeste; IB 53869 (M)-Igara (not plot-
ted in map), MCP 2917 (F), MCP 2919 (M)-Ipira;
IB 16084 (M)-Ipumirim; MCP 2876 (M), MCP
2879 (M)-Ita; MCP 180 (F)-Itapeva (not plotted
in map); IB 26664 (M)-Itapiranga; IB 16468 (M)-
JoaVaba; IB 27161 (M)-Lajes; IB 28260 (M), IB
29439 (M), IB 29970 (M), IB 30310 (M), IB 30386
(M), IB 30463 (M), IB 31036 (M), IB 31256 (M), IB
31477 (M), IB 31480 (M), IB 31548 (M), IB 31568
(M), IB 31641 (M), IB 31688 (M), IB 31710 (M), IB
31743 (M), IB 31748 (F), IB 31785 (M), IB 31786
(F), IB 31788 (M), IB 31798 (F), IB 31813 (M), IB
31842 (F), IB 31862 (M), IB 31940 (M), IB 32085
(M), IB 32122 (M), IB 32195 (M), IB 32351 (M), IB
32356 (M), IB 32376 (F), IB 32456 (M), IB 32525
(M), IB 32716 (M), IB 32719 (M), IB 32724 (M), IB
32726 (M), IB 32734 (M), IB 32750 (M), IB 32763
(M), IB 32778 (F), IB 32793 (M), IB 32796 (M), IB
32824 (M), IB 32830 (M), IB 32940 (M), IB 32946
192 HERPETOLOGICAL MONOGRAPHS
(M), IB 33005 (M), IB 33030 (M), IB 33033 (M), IB
33602 (M), IB 33663 (M), IB 33678 (F), IB 33619
(M), IB 33694 (M), IB 33697 (M), IB 34372 (M), IB
34374 (F), IB 37478 (M), IB 42368 (M), IB 42560
(M), MCP 2928 (F), MCP 2929 (M), MCP 2930 (M),
MCP 2931 (M)-Peritiba: MCP 2900 (F), MCP 2901
(F)-Piratuba; IB 5273 (M)-Rio do Peixe; IB 27905
(M), IB 27906 (F), IB 44777 (F), IB 46482 (M), IB
46566 (M), IB 53094 (M), IB 54901 (F), IB 54929
(F)-Sao Joaquim. URUGUAY (45) Departamento
Artigas (8): CZVR 327 (M)-Empalme a Barnab6
Rivera (not plotted in map); CZVR 279 (F), CZVR
562 (M), CZVR 1205 (M)-Arroyo Catalan Chico
(not plotted in map); CZVR 3365 (F), CZVR 4106
(M)-Colonia Palma; CZVR 3994 (F), CZVR 4023
(M)-Estancia Washington. Departamento Cane-
lones (9): CZVR 957 (M), CZVR 3780 (M)-Alrede-
dores de Atlantida; CZVR 1211 (F), CZVR 1213 (M),
CZVR 3310 (F), CZVR 3311 (F), CZVR 3312 (F),
CZVR 4908 (M)-La Palmita, Ruta 8; CZVR 1669
(M)-Paso Escobar. Departamento Colonia (2):
CZVR 2516 (M)-Nueva Palmia; CZVR 2863 (M)-
Puerta Gorda. Departamento Flores (3): CZVR
2864 (M)-Estancia Ariztegui; CZVR 4920 (M)-
Grata del Palacio; CZVR 4798 (F)-Ruta 3. Depar-
tamento Maldonado (2): CZVR 132 (M)-Balneario
Solis; CZVR 2865 (F)-Paso de La Horqueta. De-
partamento Melo (1): IB 8754 (M)-Melo; Depar-
tamento Rio Negro (3): CZVR 1203 (M)-Las Ca-
fiadas, 10 Km from Estancia Francia (not plotted in
map); CZVR 1484 (M)-Paso de Navarro, over Rio
Negro; CHINM 2680 (F)-Arroyo Salsinpedes
Grande (not plotted in map). Departamento Rivera
(6): CZVR 849 (M), CZVR 851 (F), CZVR 853 (M),
CZVR 872 (F), CZVR 1832 (M)-Minas de Cufnapiru
(not plotted in map); CZVR 1476 (M)-Tranqueras.
Departamento Salto (1): CZVR 4245 (M)-Pueblo
Lavalleja. Departamento San Jose (4): CZVR 556
(M), CZVR 991 (F), CZVR 1212 (F), CZVR 1389
(F)-Sierra de Mahoma (not plotted in map). De-
partamento Soriano (4): CZVR 441 (M)-San Mar-
tin (not plotted in map); CZVR 4927 (M), CZVR
5024 (F), CZVR 5070 (M)-Zona Tala. Departa-
mento Tacuarembo (1): CZVR 2866 (M)-Serra dos
Tambores. Departamento Treinta y Tres (1): CZVR
261 (M)-Santa Clara de Olimar. PARAGUAY (8).
Departamento Itapua (4): MNHNP 3478 (M)-
Encarnaci6n; MNHNP 4624 (M), MNHNP 4625
(M), MNHNP 4626 (F)-Yacireta. Departamento
Paraguari (4): MNHNP 2688 (M), MNHNP 2496
(M), MNHNP 2497 (M), MNHNP 3397 (M)-
Parque Nacional de Ybycui.
Micrurus baliocoryphus: ARGENTINA (99).
Corrientes (44): CHINM 2484 (F)-Alvear; MLP
005 (F)-Bella Vista; CHC 058 (F), CHC 218 (M),
MACN 6427 (M)-Corrientes; CHC 131 (M)-El
Sombrero; CHC 008 (M), CCG 008 (M)-Emped-
rado; CHINM 3102 (M), CHINM 3103 (M),
CHINM 3105 (M) Felipe Yofre; CHINM 2870
(M), CHINM 2871 (M), FML 1986 (M)-General
Paz; CHC 376 (M), CHINM 739 (M)-Goya; CHC
135 (M)-Itati; CHC 126 (M)-Ituzaing6; CCG 002
(M) = MZUSP 10808-Paso de La Patria; CCG 279
(F)-Ramada Paso (not plotted in map); CHC 127
No.
[No. 13
(M)-Sconza Cue (not plotted in map); CHINM 728
(M), CHINM 737 (M), CHINM 738 (M), CHINM
740 (M), CHINM 742 (F), CHINM 3053 (M)-Sa-
ladas; CCG 003 (M) = MZUSP 10809-San Luis del
Palmar; CHINM 2878 (M), CHC 217 (M), CHC 259
(M)-San Miguel; CHINM 2003 (M), CHINM 2004
(M), CHINM 2452 (F), CHINM 2959 (M), CHINM
3159 (M), CHINM 3211 (M), MLP 669 (M)-San
Roque; CHINM 1720 (F), CHINM 2008 (F)-Santa
Lucia; CCG 014 (M) = MZUSP 10819, CCG 015
(M) = MZUSP 10820, CHC 121 (M)-Santa Rosa;
CHINM 726 (F)-Sauce. Entre Rios (39): CHINM
2661 (M)-Bandera (not plotted in map), CHINM
1627 (F), CHINM 1710 (M)-Chajari; CHINM 2606
(M)-Conscripto Bernardi; CHINM 1512 (F),
CHINM 1523 (M), CHINM 1525 (F), CHINM 1526
(F), CHINM 1783 (M), CHINM 1852 (M), CHINM
1967 (M), CHINM 1994 (F), CHINM 2007 (M),
CHINM 2009 (M), CHINM 2030 (M), CHINM
2140 (M), CHINM 3076 (F)-El Cimarr6n; CHINM
2426 (M), CHINM 2665 (M), CHINM 3572 (M)-
La Paz; CHINM 1522 (M), CHINM 3391 (M),
MACN 8629 (M)-Parana; MACN 27465 (M)-Pie-
dras Blancas (not plotted in map); CHINM 3267
(M)-Rosario del Tala; CHINM 1524 (M), MACN
1012 (M), MACN 6747 (M)-Santa Elena; CHINM
3564 (M), CHINM 3657 (M)-Viale; CHINM 730
(M), CHINM 1521 (M), CHINM 1823 (M), CHINM
2397 (F), CHINM 2580 (M), CHINM 2642 (M),
CHINM 3081 (M), MLP 527 (M)-Villa Federal;
MACN 11909 (M)-Villaguay. Formosa (3):
CHINM 712 (M)-Clorinda; CHINM 2178 (M)-
Formosa; CHINM 3650 (M)-Las Lomitas. Mi-
siones (13): CHINM 2499 (M), CHINM 2705 (M)-
Ap6stoles; CHINM 2420 (M) Concepci6n de la Si-
erra; CHINM 735 (M)-Garupa; CHINM 1981
(M) Obera; CHINM 1518 (M)-Posadas; CHINM
2487 (M)-Puerto Azara; MACN 2066A (M), MACN
2066b-Puerto Laudera (not plotted in map);
CHINM 743 (M)-San Ignacio; CHINM 2788 (M),
FML 1247 (M)-San Jos6; MLP 748 (M)-Wanda.
PARAGUAY (6). Departamento Central (1):
MNHNP 2689 (M)-Nueva Italia. Departamento
Neembucu (1): MNHNP 5044 (M)-Estancia Ya-
card. Departamento Presidente Hayes (4):
MNHNP 2500 (F)-Ruta Transchaco, 17 km N;
MNHNP 5143 (M)-2 km W de la Ruta Transchaco
a General Diaz; MNHNP 5144 (F)-40 km W de la
Ruta Transchaco a General Diaz; MNHNP 5222
(M)-Ruta Militar.
Micrurus lemniscatus carvalhol (23)-ARGENTI-
NA (5) Misiones: CCG 016 (M)-Ap6stoles; CHINM
2403 (M)-Puerto Azara; CHINM 1473 (M),
CHINM 3382 (M), CCG 006 (M)-Posadas. BRA-
ZIL (17). Mato Grosso do Sul (5): IB 43646-An-
t6nio Joao; IB 17513-Itaum; MZUSP 10135-Ni-
oaque; IB 19671, IB 46010-Ponta Pora: Parana
(12): IB 16235-Araucaria (not plotted in map);
MHNCL 3105 (F)-Guaira; IB 5976-Guarapuava;
IB 4341-Imbaui; IB 5266, IB 5267-Job Eugenio
(not plotted in map); MHNCI 2188 (M)-Maringa;
MHNCI 2779 (M); MHNCI 4309 (F) Paranaval,
PR; MHNCI 4932 -So Pedro do Parana; IB 9946-
Uniao da Vit6ria. PARAGUAY (2). Departamento
1999]
1999] HERPETOLOGICAL MONOGRAPHS
Amambay (1): MHNCI 4541 (F)-Estancia Arroyo
Blanco, Capitan Bado. Departamento Caaguazu
(1): MNHNP 5145 (M)-Campo 9.
Micrurus brasiliensis: BRAZIL (15). Bahia (10):
UMMZ 108880 (M) (Holotype), UMMZ 108881 (F),
MNRJ 2494 (M), CEPB 2301 (M), IB 51310 (M)-
Barreiras; IB 54848 (F), IB 55385 (M)-Desid6rio;
IB 50142 (M), 54921 (M)-Brumado; MCZ 3298
(F)-Santa Cruz de Cabralia; Goias (2): IB 9152
(M)-Cana Brava; IB 55883 (M)-Guarani de Goias;
Minas Gerais (2): UMNZ 108878 (M)-Janudria;
MCZ 3298 (F)-Manga; Tocantins (1): AMNH
90361 (M)-Santa Isabel do Morro, Ilha do Bananal.
Micrurus diana: Bolivia (7). Departamento de
Santa Cruz (7): NKR 219 (F)-Serrania Huanchaca;
FMNH 159889 (M) (holotype); FMNH 195864 (M);
FMNH 195864 (M); FMNH 195899 (F); AMNH
120600 (M)-Serrania de Santiago; IB 31342 (M)-
Santa Cruz de La Sierra.
Micrurus pyrrhocryptus: ARGENTINA (229).
Catarmaca (14): CHINM 2873 (M), MACN 2906
(M), MACN 540A (M), MACN 540B (M)-Catar-
maca; FML 508 (M)-Finca Chacabuco (not plotted
in map); FML 873 (M), FML 1645 (M)-Hualfin;
CHINM 2530 (F)-Huillapina; FML 1184 (M)-La
Calera (not plotted in map); FML 1519 (F)-La
Puerta, 25 km from Catarmaca; FML 1645 (M)-Los
Nascimientos (not plotted in map); MACN 1714
(M)-Pomancillo; CHINM 1982 (M)-Sijan; FML
1320 (M)-Sumalao (not plotted in map). Chaco
(41): CHC 151 (M)-Brasail; CHINM 736 (M)-
Charata; CHINM 1853 (M)-Chorotis; CHINM 705
(F), CHINM 716 (M), CHINM 2496 (M)-Colonia
Benitez; CHINM 707 (M)-Colonia Elisa; CHC 023
(M), CHC 042 (M), CHC 043 (M), CHC 107 (F),
CHC 244 (M)-Colonia Las Mercedes; CHINM 709
(M)-Corzuela; CHINM 715 (M)-Enrique Urien;
CHC 344 (M)-Fuerte Esperanza (not plotted in
map); CHINM 721 (M)-General Obligado;
CHINM 2702 (M)-General Pinedo; CHINM 2567
(M), CHINM 2609 (M)-General Vedia; MACN
32073 (M)-between Haumonia and Villa Berthet;
CHINM 2778 (M), CHINM 2833 (M)-Itin;
CHINM 713 (M)-Juan Jose Castelli; CHINM 720
(F)-La Escondida; CHINM 704 (M)-La Verde;
CHINM 718 (M), CHINM 2302 (M)-Las Palmas;
CHINM 2018 (M), CHINM 2874 (M)-Machagai;
CHINM 725 (M), MACN 28768 (M)-Quitilipi;
CHINM 1472 (M), MLP 242 (M), CCG 290 (M)-
Resistemcia; CHINM 2555 (M)-Roque Saenz Pefia;
CHINM 2427 (M), CHINM 2467 (M)-Villa Angela,
Estancia Suiza; CHINM 1927 (M) CHINM 2148
(M), CHINM 2916 (M), CHINM 2937 (M)-Villa
Berthet. Cordoba (25):CHIMN 2171 (M)-Agua de
Ram6n; MACN 17535 (M)-Aniscate, Villa Naza-
reth; FML 2118 (M), MACN 24675 (M)-Ascochin-
ga; CHINM 706 (M)-Cafiada de Luque; MACN
21452 (M)-Cruz Grande (not plotted in map),
CHINM 701 (M), CHINM 708 (F)-Chaiacea;
CHINM 3258 (M)-Dean Funes; MACN 10251
(M)-Huerta Grande; CHINM 1878 (M)-La Posta;
MACN 8923 (M)-Los Cocos; MACN 19491 (M),
MACN 34370 (F)-Los Hoyos; CHINM 2954 (M)-
Lucio V. Mansilla; MLP 322 (M)-Rio Ceballos (not
193
plotted in map); FML 1167 (M), FML 1686 (M)-
San Esteban; MACN 33552 (M)-Villa Huidobro;
FML 1524 (M) Villa Maria; MACN 17535 (M)-
Villa Nazareth (not plotted in map); MACN 35546
(M)-Tanti; MLP 244 (M), MLP 245 (M), MLP 247
(M)-Tanti Viejo (not plotted in map). Formosa
(19): CHINM 2177 (M), CHINM 2915 (M)-Com-
andante Fontana; CHINM 702 (F), CHINM 703
(M)-Estanislao del Campo; CHINM 1840 (M),
CHINM 1866 (M), CHINM 1921, CHINM 1926,
CHINM 1997 (M), CHINM 2101 (M), CHINM
2330 (M), CHINM 2474 (M), CHINM 2806 (M),
CHINM 3044 (M)-Formosa; FML 2341a (M), FML
2341b (F)-20 km from Formosa; FML 1162 (M)-
Fortin Lavalle; CHC 197 (F)-Laguna Yema (not
plotted in map); CHINM 1517 (F)-Pirane. Jujuy
(1): CHINM 3398 (M)-Yuto. La Pampa (4):
MPCNLP 213 (M)-12 km SW of Lihuel Lalel (not
plotted in map); MPCNLP 222 (M)-Pulches (not
plotted in map); MPCNLP 150 (M)-25 km SE of
Puelen; MPCNLP 171 (F)-Santa Isable. La Rioja
(12): MACN 7511 (M), MACN 7512 (M), MACN
7513 (F), MACN 9993 (M), MACN 15181 (M),
MACN 24980 (M)-Aimogasta; MACN 9990 (M)-
Asha, close to Aimogasta not plotted in map);
CHINM 1907 (M)-Chilecito; MLP 333 (M)-Fa-
matina; MLP 255 (M)-La Rioja (not specific loca-
tion); CHINM 2261 (M)-Patqufa; MACN 24972
(M)-Quebrada de Duranznillo (not plotted in map).
Mendoza (2): MLP 256 (M)-Chacra de Coria (not
plotted in map); CCG 009 (F)-Mendoza. Neuquen
(5): CHINM 3532 (M)-Ciudad de Neuquen;
CHINM 3646 (M)-El Medanito (100 km NW):
CHINM 2716 (M), CHINM 2757 (M)-Portezuelo
Grande; MACN 33233 (F)-between Semillosa and
Chalac6. Rio Negro (4): MACN 24888 (M)-Allen;
MACN 30013 (M)-Chelfor6; MACN n/d-Coronel
J. J. Gomez; MLP n246 (M)-Rio Negro (no specific
location). San Juan (1): FML 1217 (M)-Vallecito.
San Luis (3): CHINM 2359 (M), CHINM 2574
(M)-Aerodromo de San Luis; MLP 649 (M)-Mer-
lo. Salta (18): FML 1576 (M)-Camino a La Isla
(not plotted in map); FML 1417 (M)-Cerrillos;
CHINM 2170 (M)-Coronel Conejo; CHINM 2540
(M)-Coronel Mollinedo; FML 358 (M)-Coronel
Montes; CHINM 2927 (M)--El Galp6n; CHINM
2829 (M)-Embarcarci6n; FML 062 (M), FML 130
(F), FML 153 (M)-Hickmann; CHINM 717 (M)-
Las Lajitas; CHINM 2568 (F)-Lumbrera; CHINM
1660 (M)-Padre Lozano; CHINM 2168 (M)-Rio
Piedras (not plotted in map); CHINM 722 (M), MLP
252 (F)-Salta; CHINM 2181 (M)-Tartagal; MACN
n(M)-Urundel. Santa Fe (29): MACN 27962
(M)-18 km of Antonio Pini; CHINM 2579 (F)-
Avellaneda; CHINM 2837 (M)-Ceres; CHINM
2907 (M)-E1 Nochero; CHINM 2701 (M)-Hersi-
lia; CHINM 2560 (M), CHINM 2899 (M), CHINM
3096 (M)-Atahualpa, Estancia Fortin; CHINM
3133 (M)-Esteban Rams; CHINM 2701 (M)-Flo-
rencia; MACN 23588 (M), MACN 26515 (F)-Gato
Colorado, 40 km NE of El Nochero; CHINM 1738
(M), CHINM 1786 (M), CHINM 2834 (M), CHINM
3100 (M)-La Lucila; CHINM 1480 (M)-Las Av-
ispas; CHINM 3651 (M)-Logrofio; CHINM 3061
194 HERPETOLOGICAL MONOGRAPHS
(M)-Monte Fiore; CHINM 2031 (M)-Moussy (not
plotted in map); CHINM 1479 (M)-Nare; CHINM
2533 (M)-Puerto Iturralde (not plotted in map);
CHINM 3658 (M)-Reconquista; CHINM 2224 (F),
CHINM 2529 (M), CHINM 3155 (M)-Tostado;
CHINM 2984 (M)-Vera; CHINM 710 (F), CHINM
711 (M)-Villa Ana. Santiago del Estero (30): FML
1506 (M)-Ahl Veremos; CHINM 2585 (M),
CHINM 3652 (M)-Afiatuya; CHINM 1850 (M),
CHINM 2664 (M)-Averfas; MACN 29618 (M)-
Bobadal; FML 1452 (M)-E1 Cadillo; CHINM 3070
(M), FML 2512 (M)-Frfas; FML 1968 (M)-Figu-
eroa; CHINM 1736 (M)-Haase; CHINM 1667
(M)-La Banda; FML 1532 (F)-La Gramilla;
CHINM 1354 (M), CHINM 1355 (M)-Los Juries;
FML 1684 (M)-Monte Quemado; CHINM 2812
(M)- Ojo de Agua; CHINM 2172 (M), MACN
27239 (M)-Otumpa; FML 1617 (M), FML 1643
(M)-Pellegrini (not plotted in map); CCG 001 (F),
CEPB 2364 (M), FML 1557 (M), MLP 254 (M)-
Santiago del Estero; CHINM 2476 (F), CHINM
2477 (M)-Tiun Punco; FML 1684 (M)-Turena
(not plotted in map); CHINM 2169 (M), CHC 030
(M)-Urutau. Tucuman (21): FML 2372 (M)-A1-
deretes (not plotted in map); CHINM 1713 (M)-
Atahona (not plotted in map); FML 1246 (F)-Bi-
[No. 13
furcaci6n Raco, Las Tipas; FML 519 (M), FML 1701
(M), FML 2042 (F)-Burruyacui; FML 054 (F)-El
Cardillal; FML 519 (M)-La Ramada; FML 1316
(F), FML 1453 (M), FML 1919 (M)-Leales (not
plotted in map); FML 808 (M)-Monteagudo; FML
166 (F)-Raco; CHINM 2749 (F), CHINM 3042
(M)-San Miguel de Tucuman; FML 1164 (M)-Tafi
Viejo; FML 436 (F), FML 2046 (M)-Ticucho; FML
060 (M), FML 312 (M), FML 518 (M)-Trancas.
PARAGUAY (5). Departamento Boqueron (5):
MNHNP 2498 (F), MNHNP 2499 (F), MNHNP
3544 (F)-Parque Nacional Teniente Enciso;
MNHNP 2686 (M)-Base Aeronaval, Pozo Hondo;
MNHNP 4018 (M)-10 km SE de Pedro Pefia.
Micrurus tricolor: BRAZIL (18). Mato Grosso do
Sul (18): IB 16290 (M)-holotype-Carandazal;
Paratypes: IB 4802 (F)-Barranco Branco; IB 14297
(F), IB 14298 (M)-Guia Lopes da Laguna; IB 6142
(F), IB 14299 (F), IB 14223 (F), IB 14253 (M), IB
14567 (M)-Taunay; COBAP 058 (M)-Anastacio;
IB 5535 (M), IB 16520 (M), IB 19616 (M)-Aqui-
dauana; IB 53454 (M)-Corumba; IB 30555 (F), IB
31575 (M), IB 32712 (M)-Guaicurus; IB 19772
(F)-Porto Murtinho. BOLIVIA (1). Departamento
Santa Cruz (1): NKR 543 (M)-1 km N de Gutier-
rez.