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Title: A Revision of Snakes of the Genus Conophis (Family Colubridae, from Middle America)
Author: Wellman, John
Language: English
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* * * * *
UNIVERSITY OF KANSAS PUBLICATIONS
MUSEUM OF NATURAL HISTORY
Volume 15, No. 6, pp. 251-295, 9 figs.
October 4, 1963
A Revision of Snakes of the Genus Conophis
(Family Colubridae, from Middle America)
BY
JOHN WELLMAN
UNIVERSITY OF KANSAS
LAWRENCE
1963
UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY
Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Theodore H. Eaton, Jr.
Volume 15, No. 6, pp. 251-295, 9 figs.
Published October 4, 1963
UNIVERSITY OF KANSAS
Lawrence, Kansas
PRINTED BY
JEAN M. NEIBARGER. STATE PRINTER
TOPEKA. KANSAS
1963
[Illustration: Union Label]
29-5936
A Revision of Snakes of the Genus Conophis
(Family Colubridae, from Middle America)
BY
JOHN WELLMAN
CONTENTS
PAGE
INTRODUCTION 253
ACKNOWLEDGMENTS 254
MATERIALS AND METHODS 254
GENUS Conophis Peters 255
Key to the Species and Subspecies 257
Analysis of Characters 257
Scutellation 258
Size and Proportions 258
Color Pattern 260
Sexual Dimorphism 260
_C. lineatus_ 262
_C. lineatus dunni_ 262
_C. lineatus lineatus_ 267
_C. lineatus concolor_ 270
_C. nevermanni_ 272
_C. pulcher_ 274
_C. vittatus_ 277
Skull 282
Dentition 288
Vertebrae 288
Hemipenes 289
Food and Feeding 289
Effect of Poison 290
TAXONOMIC RELATIONSHIPS AND EVOLUTION 291
SUMMARY 292
LITERATURE CITED 293
INTRODUCTION
Need for a comprehensive systematic review of the snakes of the genus
_Conophis_ was pointed out by Stuart (1954a, b). Since these snakes
appeared to be of zoogeographic importance in the Central American
region, I undertook the review as set forth on the following pages.
ACKNOWLEDGMENTS
For permission to examine specimens, and for information concerning
specimens in their care, I am grateful to Mr. L. C. Battersby and Miss
Alice G. C. Grandison, British Museum (Natural History); Mr. Charles
M. Bogert and Dr. Richard G. Zweifel, American Museum of Natural
History; Dr. Doris M. Cochran, United States National Museum; Prof.
William B. Davis, Agricultural and Mechanical College of Texas; Dr.
Josef Eiselt, Naturhistorisches Museums, Vienna; Prof. Norman Hartweg
and Prof. Laurence C. Stuart, Museum of Zoology, University of
Michigan; Dr. Robert F. Inger, Chicago Natural History Museum; Dr.
Alan E. Leviton, California Academy of Sciences; Mr. Edmond V.
Malnate, Academy of Natural Sciences, Philadelphia; Prof. George S.
Myers, Stanford University Natural History Museum; Mr. Wilfred T.
Neill, Ross Allen's Reptile Institute; Mr. Neil D. Richmond, Carnegie
Museum; Dr. William J. Riemer, University of Florida Collections;
Prof. Robert C. Stebbins, Museum of Vertebrate Zoology, University of
California; Prof. Hobart M. Smith, University of Illinois Natural
History Museum; and Dr. Ernest E. Williams, Museum of Comparative
Zoology, Harvard.
Prof. William E. Duellman supplied invaluable information and guidance
in my study. I am grateful to Prof. E. Raymond Hall for use of
facilities of the Museum of Natural History and editorial assistance.
I thank Prof. Laurence C. Stuart and Prof. Edward H. Taylor for
information and suggestions. My own field experience in Middle America
came as a result of assisting Professor Duellman in his own researches
supported by a grant from the National Science Foundation (NSF-G
9827). For these things I am deeply grateful. Specimens that I have
seen alive were collected by field companions Dale L. Hoyt and Jerome
B. Tulecke. Finally, I am grateful to my wife, Margaret L. Wellman,
for much help including typing much of the manuscript.
MATERIALS AND METHODS
Of the 325 specimens of the genus _Conophis_ available to me,
representing most of those in museum collections, scale counts were
made in the usual manner on 309. Ventrals were counted following the
system proposed by Dowling (1951:97-99); the anal plate was not
included. The anteroposterior position of the place where reduction
occurs in the number of the dorsal rows of scales is designated by
citing the number of the ventral scale directly beneath that place.
Measurements were taken to the nearest millimeter by means of a
millimeter stick. Body length is the distance from the tip of the
snout to the posterior edge of the anal plate; tail length, from the
latter point to the tip of the tail; and total length, the sum of the
body plus tail.
Descriptions of color are based on preserved specimens. Where
descriptions of the color of living individuals are given, the data
were taken from Kodachrome slides made available to me by William E.
Duellman. Due to the transient nature of the longitudinal dark stripes
in these snakes, no standard terminology has been devised, except that
the posterior continuations of the stripes which on the head pass
through the eye are termed lateral stripes; the posterior
continuations of the median stripe of the head are termed
dorsolateral stripes. A paravertebral stripe is one that is present
on the scale-row on either side of, but not including, the mid-dorsal
(vertebral) scale-row.
In order to reduce confusion in the discussion of variation, the
numbers designating the rows of dorsal scales are written as 1st, 2nd,
whereas the numbers designating the stripes are written as first,
second.
Except in three dried skeletons, teeth were counted on dentigerous
bones _in situ_. Since teeth are often missing, the sockets were
counted in order to obtain an accurate count.
In accounts of the species and subspecies, the observed range of
variation is followed by the mean in parentheses; in some instances
the mean is followed by the standard deviation, also in parentheses.
An example is 65-79 (70.6 ± 3.93).
Each synonymy includes all generic and specific combinations known to
me that have been used for the genus, and, in addition, references to
catalogues, checklists, and reports of collections.
Localities of occurrence that are not plotted on the distribution maps
are recorded in italic type under Specimens Examined. In the list of
Specimens Examined the localities and specimens are listed in the
following order: countries in alphabetical order; states or
departments in alphabetical order in each country; localities in
alphabetical order in each state or department; museum numbers in
numerical order after the abbreviations of names of museums. When more
than one specimen bears a single catalogue number, the number of
specimens is given in parentheses following the museum catalogue
number. Specimens for which data are given only as to country or to
state or department are listed first after the name of that political
unit under "no specific locality."
The abbreviations for the museum collections are:
AMNH American Museum of Natural History
ANSP Academy of Natural Sciences of Philadelphia
BMNH British Museum (Natural History)
CAS California Academy of Sciences
CNHM Chicago Natural History Museum
ERA-WTN E. Ross Allen-Wilfred T. Neill, Ross Allen's Reptile Institute
KU University of Kansas Museum of Natural History
MCZ Museum of Comparative Zoology, Harvard
MVZ Museum of Vertebrate Zoology, University of California
NMW Naturhistorisches Museums Wien, Vienna
SU Stanford University Natural History Museum
TCWC Texas Cooperative Wildlife Collection, Agricultural and
Mechanical College of Texas
UF University of Florida Collections
UIMNH University of Illinois Museum of Natural History
UMMZ University of Michigan Museum of Zoology
USNM United States National Museum
Family COLUBRIDAE
Subfamily Xenodontinae
Genus =Conophis= Peters
_Tomodon_ (part) Duméril, Bibron and Duméril, Erpétologie Générale,
7(pt.2):936, February 7(pt.2):936, February 25, 1854 (_lineatus_
and _vittatus_); Salvin, Proc. Zool. Soc. London, 28:455, 1860
(_pulcher_).
_Psammophis_ (part), Günther, Catalogue of Colubrine Snakes in
the Collection of the British Museum, London, 1858:135
(_lineatus_).
_Conophis_ Peters, Monatsb. Akad. Wiss. Berlin, 1860:519-520,
pl., fig. 3 (_vittatus_); Cope, Proc. Acad. Nat. Sci.
Philadelphia, 13:300, December 28, 1861 (_lineatus concolor_);
Proc. Acad. Nat. Sci. Philadelphia, 18:318-319, February 20,
1867 (_lineatus concolor_); Proc. Acad. Nat. Sci.
Philadelphia, ser. 2, 8:137, 1876 (_pulcher_); Bocourt in
Duméril, Bocourt and Mocquard, Mission Scientifique au Mexique
et dans l'Amerique Centrale, 2:643-644, pl. 38, fig. 5, 1886
(_lineatus lineatus_); Cope, Proc. Amer. Philos. Soc., 23:489,
October 28, 1886; Hoffmann, Klassen und Ordnungen des
Thier-Reichs. Reptilien. Bd. 6, 3:1707, 1890; Cope, Trans.
Amer. Philos. Soc., 18:207, April 15, 1895; Dunn, Bull.
Antivenin Inst. Amer., 2(1):21, 24, April, 1928; Copeia, no.
4:214, December 31, 1937 (_nevermanni_).
_Tachymenis_ (in part), Garman, Bull. Essex Inst., 16:33,
January 9, 1884 (_vittatus_ and _lineatus_).
_Erythrolamprus_ (in part), Ditmars, Bull. Antivenin Inst.
Amer., 2(2):27-29, June.
_Coniophanes_ (in part), Wettstein, Sitz. Akad. Wiss. Wien,
mathem-naturw. kl. 143:37-38, 1934 (_nevermanni_).
_Historical summary._--In 1854 Duméril, Bibron and Duméril described
and figured _Tomodon lineatum_ from America. In 1860 Peters described
and figured as a new genus and species, _Conophis vittatus_, based on
a specimen that he had obtained from a dealer in Hamburg. The
provenance of this specimen is not known, for it was discovered aboard
a ship near the mouth of the Mississippi River. It was not until 1871
that Cope included _lineatus_ in the genus _Conophis_. Cope (1861)
proposed the name _Conophis vittatus_ (_nec_ Peters, 1860). Later
(1900) he changed its name to _Conophis lineaticeps_. Early
uncertainty of the relationships of the species _lineatus_ caused
Günther (1858) to place it in the genus _Psammophis_. With the
exception of Garman (1884a and 1884b) who placed _lineatus_ in the
genus _Tachymenis_, and Wettstein (1934) who reported five specimens
of _Conophis nevermanni_ as _Coniophanes i. imperialis_, all specimens
reported after 1876 were placed in the genus _Conophis_.
The only previous attempt to review the systematics of this genus was
made by Smith (1941) who based his study primarily on specimens in the
United States National Museum. He examined only 28 specimens,
including none of one species (_nevermanni_).
_Description._--Hemipenis slightly bifurcate having forked sulcus
spermaticus, large spines near base, and smaller spines or papillae on
flounces nearer apices; prediastemal maxillary teeth 8-12, subequal in
length, and followed by short diastema and one enlarged fang or two;
fangs grooved, only one functional at any one time, unless snake is in
process of shedding teeth; teeth 6-10 on palatine, 15 to 19 on
pterygoid, 15 to 21 on dentary; teeth on dentary decreasing in size
posteriorly; large parotid (venom) gland on either side of head in
temporal region; head shields of basically unmodified colubrid type
excepting decurved rostral; rostral concave below and therein modified
for burrowing; internasals and prefrontals paired; nasals divided;
loreal single; preocular one, rarely two; postoculars, two;
supralabials, 7-8, 3rd and 4th or 4th and 5th under eye; infralabials,
8-11, usually 9 or 10; temporals, normally 1 plus 2 plus 3;
chin-shields subequal in length; ventrals, 149-183, rounded and
overlapping; caudals, 55-89, paired and imbricate; anal divided;
dorsal scales smooth and in 19 rows at mid-body with no apical pits
or keels; scale reduction normally involving fusion of 3rd and 4th
rows, resulting in 17 scale-rows near tail; tail length more than 20
per cent of body length; maximum total length exceeding 1.1 meters;
dorsal color pattern consisting of dark stripes, or no darkening, on
paler ground-color; ventral surfaces immaculate pale yellowish or
white, except on specimens having single lateral dark spots on some or
all ventrals; pupil round; diurnal or crepuscular; feeding primarily
on small lizards, sometimes on small mammals or other snakes.
_Distribution._--Semi-arid regions of southern México and Central
America as far south as Costa Rica.
KEY TO THE SPECIES AND SUBSPECIES
Although many juveniles differ greatly in general coloration from the
adults, both the juveniles and the adults of any species or subspecies
can be identified from the following key; juveniles differ from adults
in extent and intensity of dark pigmentation but not in rows of scales
involved.
1. Seven supralabials (3rd and 4th below orbit); 3 to 8 dark
stripes along body 2
Eight supralabials (4th and 5th below orbit); unstriped or with
more than 4 dark stripes along body, or dark with 2 or 4 pale
stripes 3
2. Dark stripes involving no more than one longitudinal
scale-row _C. lineatus lineatus_ (part), p. 267
Dark stripes involving at least two adjacent scale-rows
_C. vittatus_, p. 277
3. Supralabials having black borders above; head and body
generally black with 2 or 4 white lines running length
of body _C. nevermanni_, p. 272
Supralabials immaculate or having dark borders below; head
and body usually pale with dark stripes, or without stripes 4
4. Lateral dark stripe through eye involving upper half of second
scale-row; dark stripe on paravertebral row, at least
posteriorly _C. pulcher_, p. 274
Lateral dark stripe becoming indistinct on body, or restricted
to 4th or 3rd and 4th rows anteriorly, not involving 2nd
scale-row on anterior 1/3 of body (an auxiliary lateral stripe
sometimes present involving 2nd row); no paravertebral stripes 5
5. Stripes disappearing posteriorly (except for small spots of
pigment on scale-row 4 or 7); 1st scale-row unpigmented
_C. lineatus concolor_, p. 270
Stripes present posteriorly; 1st scale-row pigmented 6
6. Lateral stripes narrow on nape, restricted to 4th scale-row
on body _C. lineatus lineatus_ (part), p. 267
Lateral stripes involving 3rd and 4th rows, at least on
nape _C. lineatus dunni_, p. 262
Analysis of Characters
Characters showing inter-specific and intra-specific variation and
that have a wide range of variation were analyzed statistically, when
possible, in order to determine extent of variation. One character
(see table 3) was analyzed for sexual dimorphism, and for it the
coefficient of difference is also given. The statistical terms and
formulae have been adopted from Mayr, Linsley and Usinger (1953).
Dorsal head shields varied individually and were of no taxonomic
importance. Osteological and hemipeneal characters did not show enough
variation to be considered here.
Scutellation
Labials, dorsals, ventrals, and subcaudals were the most useful
scales.
_Labials._--All species usually have eight supralabials except _C.
vittatus_, which has seven. The only other population having a
relatively high frequency of occurrence of seven supralabials is _C.
l. lineatus_. In specimens having eight supralabials, the fourth and
fifth enter the orbit; in specimens having seven supralabials, the
third and fourth enter the orbit (the second and third are fused).
Usually there are ten infralabials, sometimes nine or eleven;
specimens having seven supralabials usually have nine infralabials,
sometimes eight, rarely ten.
_Dorsals._--Although there is no variation in the number of rows of
dorsal scales, there is some in the method of scale reduction. There
are 19 rows of dorsal scales from close behind the head to about
midway on the body where two rows are lost, leaving 17 rows from there
to near the base of the tail. This reduction is accomplished by fusion
of the scales of the 3rd and 4th rows or sometimes by the dropping out
of the 3rd row. The place at which reduction occurs in number of
dorsal scales in relation to the ventral (scale) directly below is
highly variable and of little taxonomic importance (table 1).
TABLE 1.--VARIATION IN THE PLACE OF DOSAL SCALE REDUCTION IN CONOPHIS.
Key to Columns
====================================
Std. Dev. = Standard Deviation
Std. Err. = Standard Error
Coe. Var. = Coefficient of Variation
==============+===========+========+=======+======+======+======
| Number of | | | Std. | Std. | Coe.
Taxon | Specimens | Range | Mean | Dev. | Err. | Var.
--------------+-----------+--------+-------+------+------+------
_l. concolor_ | 45 | 89-114 | 102.5 | 5.57 | 0.83 | 5.43
_l. dunni_ | 36 | 91-111 | 102.1 | 4.59 | 0.77 | 4.50
_l. lineatus_ | 26 | 91-107 | 100.2 | 3.59 | 0.72 | 3.58
_nevermanni_ | 6 | 84- 97 | 93.2 | 4.71 | 1.92 | 5.05
_pulcher_ | 26 | 94-119 | 104.6 | 4.90 | 0.96 | 4.68
_vittatus_ | 170 | 84-118 | 102.3 | 6.60 | 0.16 | 6.45
--------------+-----------+--------+-------+------+------+------
_Ventrals._--The number of ventral scutes varies from 149-183, and
shows no significant variation in the means (table 2).
_Subcaudals._--The number of subcaudal scutes varies from 55 to 89. In
some populations there is no overlap in the range of variation of
males and females. The total variation and sexual dimorphism are
analyzed in table 3.
Size and Proportions
Although considerable variation in size is observable, little
taxonomic use is made of size since sufficient series are not
available to determine age classes. The subspecies attaining the
largest size is _C. lineatus concolor_; all others are smaller and of
about the same size and proportions. The longest specimen, a male of
_C. l. concolor_, has a body length of 893 mm., a tail length of 274
mm., and a total length of 1167 mm.
TABLE 2.--VARIATION IN THE NUMBER OF VENTRALS IN CONOPHIS.
Key to Columns
====================================
Std. Dev. = Standard Deviation
Std. Err. = Standard Error
Coe. Var. = Coefficient of Variation
==============+===========+=========+=======+======+======+======
| Number of | | | Std. | Std. | Coe.
Taxon | Specimens | Range | Mean | Dev. | Err. | Var.
--------------+-----------+---------+-------+------+------+------
_l. concolor_ | 45 | 158-170 | 163.7 | 1.56 | 0.23 | 0.95
_l. dunni_ | 36 | 159-178 | 167.2 | 4.56 | 0.76 | 2.72
_l. lineatus_ | 26 | 157-169 | 163.5 | 3.59 | 0.72 | 2.20
_nevermanni_ | 6 | 173-183 | 176.5 | 4.00 | 1.63 | 2.27
_pulcher_ | 26 | 149-180 | 169.5 | 5.31 | 1.04 | 3.13
_vittatus_ | 171 | 149-180 | 163.7 | 6.33 | 0.15 | 3.87
--------------+-----------+---------+-------+------+------+------
TABLE 3.--SEXUAL DIMORPHISM AS INDICATED BY VARIATION IN THE NUMBER OF
SUBCAUDALS IN CONOPHIS.
Key to Columns
====================================
Num. Spc. = Number of Specimens
Std. Dev. = Standard Deviation
Std. Err. = Standard Error
Coe. Var. = Coefficient of Variation
Coe. Dif. = Coefficient of Difference
====================+=====+====+=======+======+======+======+======+=====
| |Num.| | | Std. | Std. | Coe. | Coe.
Taxon | Sex |Spc.| Range | Mean | Dev. | Err. | Var. | Dif.
--------------------+-----+----+-------+------+------+------+------+-----
_lineatus concolor_ | [M] | 22 | 68-74 | 70.3 | 2.14 | 0.46 | 3.04 |
| | | | | | | | 1.97
| [F] | 16 | 56-65 | 61.8 | 2.18 | 0.55 | 3.53 |
| | | | | | | |
_lineatus dunni_ | [M] | 14 | 67-80 | 74.5 | 3.86 | 1.03 | 5.18 |
| | | | | | | | 0.95
| [F] | 16 | 60-72 | 67.1 | 3.91 | 0.97 | 5.82 |
| | | | | | | |
_lineatus lineatus_ | [M] | 11 | 67-73 | 69.8 | 6.17 | 1.85 | 8.84 |
| | | | | | | | 0.60
| [F] | 9 | 60-66 | 62.4 | 6.17 | 2.06 | 9.89 |
| | | | | | | |
_nevermanni_ | [M] | 3 | 82-89 | 85.3 | .... | .... | .... |
| | | | | | | | ....
| [F] | 2 | 71-76 | 73.5 | .... | .... | .... |
| | | | | | | |
_pulcher_ | [M] | 7 | 70-79 | 74.3 | 3.11 | 1.17 | 4.19 |
| | | | | | | | 0.93
| [F] | 11 | 65-71 | 68.2 | 3.42 | 1.08 | 5.01 |
| | | | | | | |
_vittatus_ | [M] | 95 | 59-76 | 67.8 | 3.33 | 0.34 | 4.91 |
| | | | | | | | 1.28
| [F] | 58 | 55-66 | 60.0 | 2.75 | 0.36 | 4.58 |
--------------------+-----+----+-------+------+------+------+------+-----
Color Pattern
This is the primary feature used to separate species and subspecies in
this genus. The color pattern consists of three black or deep brown
stripes on the dorsal part of the head, one mid-dorsally, and one on
each side of the head passing through the eye. On the body, there are
usually dark longitudinal stripes on a pale tan or white background.
There may be as few as three in _vittatus_, and as many as 13 in _l.
dunni_; except that there is none in _C. l. concolor_. There are two
pairs of primary dark stripes. The first is the body stripe that is
the posterior extension of the stripe which on the head passes through
the eye and is termed the lateral stripe. The other primary stripe is
the posterior continuation of the mid-dorsal head stripe. Usually it
is split into two dorsolateral stripes on the body. Stripes may be
present on the scale-row to either side of the primary stripe. These
stripes are usually dark brown or black and are the secondary stripes.
Finally, additional stripes may be present that are paler brown and
bear no direct relationship to the primary stripes. These are
auxiliary stripes.
Every stripe originates either as broad continuous stripe or as a row
of spots or dashes, forming a discontinuous stripe, which in some
specimens becomes continuous posteriorly. The stripes are usually
black or deep brown, although auxiliary stripes are sometimes paler.
The dorsal ground color is pale brown, tan, olive, or white; usually
the ground color is palest ventrally and darkest dorsally.
In some specimens of _Conophis_ the lateral tips of the ventrals are
spotted, one spot on each end of each ventral. Otherwise, the ventrals
are immaculate white.
In some species there is considerable ontogenetic change in color
pattern, although the juveniles bear the basic color characteristics
of the adults. For example, juveniles of the sympatric species _C.
lineatus dunni_ and _C. pulcher_ can be separated on the basis of
which scale-rows are darkly pigmented. _C. l. dunni_ has eight stripes
in juveniles and as many as 13 in adults. Juveniles show a greater
contrast between the black stripes and the pale ground color than do
adults. With increased age (size) the stripes in some populations
become paler and are split; simultaneously the ground color becomes
darker.
Sexual Dimorphism
Sexual dimorphism is evident in all species and subspecies of
_Conophis_. Differences always exist in the number of subcaudals and
in the tail/body ratio; males have more subcaudals and relatively
longer tails than do females (table 3). Otherwise, there is little
sexual dimorphism in these snakes. Males and females cannot be
differentiated by any feature of coloration.
Formulation of a biological concept of the species as defined by Mayr
(1942) is difficult when most of the data primarily relied upon are
from preserved specimens. Nevertheless, a total view of variation was
attempted so that differences within and between populations could be
recognized. Differences, between populations, that seem to be part of
a continuous or internal cline (Huxley, 1942) are not used for
characterizing subspecies.
[Illustration: FIG. 1. Patterns of dorsal coloration at
mid-body of adults of all species and subspecies of the genus
_Conophis_ except _C. lineatus concolor_. A. _C. lineatus
dunni_ (UMMZ 107339) from Santa Rosa, Guatemala. B. _C.
lineatus dunni_ (UMMZ 116537) from 1.5 mi. N Matagalpa,
Nicaragua. C. _C. lineatus dunni_ (ANSP 3480) from "San Jose,"
Costa Rica. D. _C. l. lineatus_ (KU 23253) from Río Blanco,
20 km. WNW Piedras Negras, Veracruz, México. E. _C. nevermanni_
(ANSP 22424) "San Jose," Costa Rica. F. _C. pulcher_ (UIMNH
33646) from Soconusco, Chiapas, México. G. _C. vittatus_ (KU
39626) from Atencingo, Puebla, México. H. _C. vittatus_ (TCWC
9473) from 1 mi. S Colotlipa, Guerrero, México. I. _C.
vittatus_ (UMMZ 82653) from "vicinity of" Salina Cruz, Oaxaca,
México. Approximately × 3/4.]
=Conophis lineatus= (Duméril, Bibron and Duméril)
_Tomodon lineatum_ (in part) Duméril, Bibron and Duméril,
Érpétologie Genérale, 7(pt. 2):936-938, February 25, 1854.
_Diagnosis._--No dark pigmentation posterior to nape; lateral dark
stripe anteriorly passing through eye and posteriorly involving 4th or
3rd and 4th scale-rows only; first scale-row darkly pigmented; no
paravertebral dark stripe; six to thirteen (or no) dark stripes at
mid-body; usually eight (sometimes seven) supralabials immaculate
white or having dark ventral margins.
_Variation._--The variation in this species is discussed more
completely in the descriptions of the subspecies. One hundred and
seven specimens have 157 to 178 (164.8) ventrals. Eighty-eight of
these snakes having complete tails have 56 to 80 (68.0) subcaudals;
the number of ventrals plus subcaudals varies from 222 to 247 (233.5)
in 87 of these. On 107 specimens the reduction from 19 to 17 dorsal
scale-rows takes place between ventrals 89 and 114 (101.8). Sexual
dimorphism is evident in the number of subcaudals; there are, on the
average, fewer subcaudals in females than in males of each subspecies.
The largest specimen is a male _C. l. concolor_ (USNM 46345) from
Chichén Itzá, Yucatán, México, having a body length of 893 mm., a tail
length of 274 mm. and a total length of 1167 mm. The smallest is a
juvenile _C. l. dunni_ (MCZ 49749) from Tegucigalpa, Honduras, having
a body length of 162 mm., a tail length of 51 mm. and a total length
of 213 mm.
The greatest variation is in coloration. Dark color, or lack thereof,
has been used to separate the subspecies of _C. lineatus_. The
ground-color is pale brown, pale olive or white, either with no
stripes on the body or with eight to thirteen dark stripes at
mid-body. Specimens having dark stripes on the body always have black
or dark brown pigmentation on the first, 4th and 7th dorsal
scale-rows. In some there is dark pigmentation on the 2nd, 3rd, 8th
and 10th rows of scales. The stripes appear on the nape or farther
posteriorly, usually on the anterior third of the body, either as a
series of spots or dashes that form a continuous stripe farther
posteriorly or as a continuous stripe.
The ventrals usually have more or less conspicuous dark spots
laterally on those specimens having dark stripes present on the
dorsum; spots are absent on all specimens having no dorsal stripes and
on some specimens having dorsal stripes. Except for the dark lateral
spots (when present) the ventrals are immaculate white. Usually the
dorsal ground-color is pale tan, especially on the striped forms. The
ground-color is usually palest on the lower dorsal scale rows and
darkest dorsally.
Three populations are separable as subspecies; one has no stripes on
the body and occurs in the Yucatán Peninsula. The other two have
stripes on the dorsum and vary clinally in coloration from the north
(Veracruz, México) to south (Costa Rica) (Fig. 2). Reasons for
separating these widespread, variable snakes into two subspecies are
that they are discontinuous in distribution (the population in
Veracruz is disjunct from the one that extends from Guatemala to Costa
Rica), and that these populations have distinctly different color
patterns.
[Illustration: FIG. 2. Selected locality records for the
subspecies of _Conophis lineatus_.]
=Conophis lineatus dunni= Smith
_Psammophis lineatus_, Günther, Catalogue of Colubrine Snakes
in the Collection of the British Museum, p. 135, 1858.
_Conophis lineatus_, Cope, 3rd Ann. Rept. Peabody Acad. Sci.,
p. 82, 1871; Proc. Acad. Nat. Sci. Philadelphia, 23:204,
October 24, 1871; Journ. Acad. Nat. Sci. Philadelphia, ser. 2,
8:137, 1876; Bull. U. S. Natl. Mus., 32:77, 1887; Günther,
Biologia Centrali-Americana, p. 165, March, 1895; Boulenger,
Catalogue of the Snakes in the British Museum (Natural
History), 3:122-123, 1896; Werner, Arch. Naturges., 90, abt. A,
12:143, 1925; Schmidt, Zool. Ser. Field Mus. Nat. Hist.,
12:199-200, November 21, 1928; Amaral, Mem. Inst. Butantan,
4:212, 1929; Werner, Zool. Jahrb., 57:184, 1929; Stuart, Occas.
Papers Mus. Zool. Univ. Michigan, 292:5, June 29, 1934; Dunn,
Copeia, no. 4:214, December 31, 1937.
_Conophis lineatus similis_ Smith, Journ. Washington Acad.
Sci., 31:123-124, March 15, 1941 (Type.--United States National
Museum, No. 79963; type locality.--Managua, Nicaragua; _nec_
Bocourt _in_ Duméril, Bibron and Mocquard, Mission Scientifique
au Mexique et dans l'Amerique Centrale, 2:647-648, 1886);
Cochran, Bull. U. S. Natl. Mus., 220:167, 1961.
_Conophis lineatus dunni_ Smith, Proc. U. S. Natl. Mus.
92:394-395, November 5, 1942; Savage, Trans. Kansas Acad. Sci.,
50:483-486, December 31, 1949; Taylor, Univ. Kansas Sci. Bull.,
34(pt. 1):145, October 1, 1951; Neill and Allen, Publ. Res.
Div. Ross Allen's Rept. Inst., 2:56, November 10, 1959;
Herpetologica, 16:146-148, fig. 2, September 23, 1960.
_Conophis pulcher pulcher_, Stuart, Misc. Publ. Mus. Zool.
Univ. Michigan, 69:79, June 12, 1948; Contr. Lab. Vert. Biol.
Univ. Michigan, 45:24, May, 1950; Contr. Lab. Vert. Biol.
Univ. Michigan, 49:14, August, 1951; Contr. Lab. Vert. Biol.
Univ. Michigan, 65:19-20 (part), March, 1954.
_Conophis pulcher plagosus_, Mertens, Zool. Anz., 148:93,
February, 1952; Abhand. Senken. Naturw. Gesell., 487:61-62,
December 1, 1952.
_Conophis lineatus nevermanni_, Taylor, Univ. Kansas Sci.
Bull., 37(pt. 1):563-565, fig. 16, October 15, 1955.
_Type._--United States National Museum, no. 79963, obtained by Lt. H.
C. Kellers. Type locality: Managua, Nicaragua. There are also three
paratypes; one a topotype (USNM 79964), one from "Nicaragua" (USNM
25237), and one from Esparta, Costa Rica (USNM 37758).
_Diagnosis._--Lateral dark stripe anteriorly passing through eye and
posteriorly involving 3rd and 4th scale-rows; 1st scale-row darkly
pigmented; no paravertebral dark stripe, although vertebral row
sometimes darkly pigmented; six to thirteen stripes at mid-body; eight
supralabials immaculate or having dark ventral margins.
_Variation._--Thirty-six specimens have 159 to 178 (167.2 ± 4.56)
ventrals. Thirty of these snakes having complete tails have 60 to 80
(70.5 ± 5.36) subcaudals; the number of ventrals plus subcaudals
varies from 224 to 247 (237.6). In 36 specimens the reduction from 19
to 17 dorsal scales takes place between ventrals 91 and 111 (102.1 ±
4.59). Sexual dimorphism is evident in the number of subcaudals; 16
females have 60 to 72 (67.1), and 14 males have 67 to 80 (74.5)
subcaudals. The largest specimen (ERA-WTN BH-300) is a female from
Augustine, British Honduras, having a body length of 732 mm., a tail
length of 183 mm. and a total length of 915 mm. A juvenile (MCZ 49794)
from Tegucigalpa, Honduras, has a body length of 162 mm., a tail
length of 51 mm. and a total length of 213 mm.
The greatest variation is in coloration. The ground-color is pale
brown or white with dark stripes of black or deep brown present
dorsally and laterally. Some specimens from Costa Rica have as many as
13 dark stripes at mid-body (fig. 1, C). In these snakes the first
row of dorsal scales bears a series of large, slightly elongated, dark
spots; on the 2nd row a narrow dark brown stripe on the middle of the
scales; on the 3rd a black stripe on the dorsal one-third to one-half
of the scales; on the 4th and the 7th rows black stripes on the medial
half of the scales of each row; on the 8th and 10th (vertebral) rows
dark brown stripes on the medial third of the scales of each row. A
specimen from Guatemala (UMMZ 107339) shows the greatest reduction of
stripes and dark pigmentation (fig. 1, A); it has only eight stripes
at mid-body: on the first row of dorsal scales a discontinuous stripe
is formed by a series of dashes; the 3rd row bears a series of small
black spots near the base and tip of each scale; the 4th and 7th rows
bear continuous black stripes on the medial third to fourth of the
scales of each row; the 8th row has extremely small dark spots near
the tips of some scales.
The primary stripes, characteristic of the species _lineatus_, are
those on the 1st, 4th and 7th rows of dorsal scales; these are the
most prominent stripes. In some specimens these primary stripes begin
as spots or dashes on the nape and become continuous stripes
posteriorly; in others they are continuous for the length of the body.
The stripe on the 1st row is most variable; usually it consists of
only a discontinuous series of dashes for most of its length. The
secondary stripes are those on the 3rd and 8th rows; of these, only
the one on the 3rd scale-row is present on the nape. The stripe on the
3rd row in combination with the dark stripe on the 4th row is the
posterior continuation of the dark stripe that on the head passes
through the eye; this stripe is characteristic of _C. lineatus dunni_.
Both secondary stripes usually begin anteriorly as a series of spots
or dashes and become continuous stripes posteriorly; occasionally near
the base of the tail they fuse with the primary stripes on the 4th and
7th rows. In some specimens in Costa Rica indistinct stripes are
present on the 10th (posteriorly the 9th) rows, and in some specimens
in Honduras, Nicaragua, and Costa Rica similar indistinct stripes are
present on the 2nd row.
Usually there are more or less conspicuous dark spots laterally on the
ventrals, but in some specimens there are no spots. Except for the
dark lateral spots (when present) the ventrals are immaculate white.
The dorsal ground-color is a pale brown or brownish white in preserved
specimens on the 1st, 2nd, 3rd and 4th rows of scales where dark
stripes or spots are not present. The ground-color of the dorsum
between the 5th rows on each side is a somewhat darker shade of pale
to medium brown.
Never is more than the lower one-third of each of the supralabials
brown. In many specimens little or no brown is present on the lower
margins of these scales. Some of the specimens having brown on the
supralabials also have dusky markings of tan or gray on the chin and
infralabials. Specimens from the northern part of the range
(Guatemala) less frequently have dark chins and supralabials than do
specimens from the southern part of the range (Costa Rica). There is,
nevertheless, at any one locality considerable variation in the amount
of dark pigmentation present on the chin and supralabials, thereby
indicating that the slight geographic trend in this character is not
significant.
Probably the most common pattern of dorsal coloration consists of
eight or ten dark stripes (fig. 1, B). In snakes having this pattern
the stripes on the 1st, 3rd, 4th and 7th rows are always present and
prominent, although those on the 1st and 3rd rows sometimes are
present as discontinuous rows of dashes. The ground-color from the
venter to the 7th row is usually pale brown, and that dorsally between
the 7th rows on each side is usually a darker, medium brown. A series
of spots or dashes or a continuous stripe is sometimes present on the
8th row of scales.
Snakes having a larger number of dark stripes and more dark
pigmentation occur in the southern part of the range. There seems to
be a cline from paler snakes having fewer stripes in the north to
darker snakes in the south.
[Illustration: FIG. 3. Patterns of dorsal coloration at
mid-body of juveniles of two sympatric species of _Conophis_.
A. _C. lineatus dunni_ (MCZ 49794) from Tegucigalpa, Honduras.
B. _C. pulcher_ (MCZ 49791) from Tegucigalpa, Honduras.
Approximately × 1.]
In juveniles, there are six or eight black stripes boldly contrasting
with a white or pale tan ground-color (fig. 3, A). The first pair of
stripes is on the 1st scale-row; the second pair, on the 3rd and 4th
scale-rows; the third pair, on the 7th row; the fourth pair (when
present), on the 8th row. Ontogenetic change in coloration consists of
the splitting of the second pair of dark stripes in the juvenile.
Additional stripes may form later on the 2nd and/or 10th rows of
dorsal scales.
_Remarks._--Savage (1949:483-486) stated that his specimen of _C. l.
dunni_ (from Honduras) resembled _l. lineatus_ in having secondary
stripes on the 2nd and 8th rows and dark pigmentation throughout the
length of the 2nd row. As can be seen from the preceding discussion of
variation, a specimen having this color pattern is clearly within the
observed range of variation of _l. dunni_. The specimen in no way
represents an intergrade between _C. l. dunni_ and _l. lineatus_.
A specimen in the British Museum (Natural History), catalogued in 1853
(no. 53.2.4.16), has the locality listed as "México." Since this
specimen is of _C. l. dunni_ and this subspecies occurs only south of
México, the locality must be considered erroneous; possibly the
locality as recorded referred only to the fact that the specimen came
from tropical Middle America.
The absence of paravertebral stripes, the presence of a lateral
dark stripe on the nape involving the 3rd and 4th rows of scales,
and the darkly pigmented 1st scale-row, in combination with the
characteristics of the genus, distinguish _C. l. dunni_ from all other
snakes in México and Central America. The only sympatric species of
this genus, _C. pulcher_, differs in that it has paravertebral stripes
(though never a vertebral dark stripe). _Conophis pulcher_ has a
lateral dark stripe that includes the upper half of the second
scale-row on the anterior part of the body; stripes of _C. l. dunni_
never include more than the 3rd and 4th rows. Even as juveniles the
paravertebral row is not darkly pigmented in _C. l. dunni_ as it is in
_C. pulcher_.
_Distribution._--Semi-arid habitats from sea level to elevations of
1000 m. from the Cuilco Valley in western Guatemala, El Peten and
British Honduras southeastward to northeastern and southern Honduras,
western Nicaragua and northwestern Costa Rica (fig. 2).
_Specimens examined._--Total of 41 specimens, as follows: BRITISH
HONDURAS: _Cayo District_: Augustine, ERA-WTN BH-300; _Mountain Pine
Ridge, 10 mi. E Augustine_, ERA-WTN BH-298.
COSTA RICA: _no specific locality_, AMNH 17309. "_Cartago_," BMNH
71.11.22.15. _Puntarenas_: 32 km. N Barranca, KU 35630; Esparta, USNM
37758. "_San José_," ANSP 3480, 12232.
EL SALVADOR: _Morazan_: El Divisadero, CNHM 10999. _San Miguel: San
Pedro_, MCZ 57061.
GUATEMALA: _El Petén_: Sojio (Toocog), AMNH 69969, 69986.
_Huehuetenango_: flood plain Río Cuilco, W of Finca Canibal, 18 km. N
Tacaná, UMMZ 98283. _Santa Rosa_: Santa Rosa, UMMZ 107339.
HONDURAS: _no specific locality_, AMNH 32814, UF 7657. _Cortes:
Cofradía_, SU 8422; _Gracias_, CNHM 28560; _Hacienda de Santa Ana, W
San Pedro Sula_, CNHM 5297; San Pedro Sula, UMMZ 68695(2); _near San
Pedro Sula_, MCZ 27563. _Francisco Morazan: Potrero de Melio, Escuela
Agricola Pan-americana_, MCZ 49987; Tegucigalpa, MCZ 49784, 49786,
49789-90, 49792, 49794.
MÉXICO: _no specific locality_, BMNH 53.2.4.16.
NICARAGUA: _no specific locality_, UMMZ 65633, USNM 25237. _Leon_: El
Polvón, MCZ 5645, 5696. _Managua_: Managua, USNM 79963-64; _3 mi. SW
Managua_, KU 42315; _8 mi. WNW Managua_, KU 42314; _1 mi. N Sabana
Grande_, KU 42311-13. _Matagalpa_: 1.5 mi. N Matagalpa, UMMZ 116537.
=Conophis lineatus lineatus= (Duméril, Bibron and Duméril)
_Tomodon lineatum_ (in part) Duméril, Bibron and Duméril,
Érpétologie Genérale, 7(pt. 2):936-938, atlas, pl. 73,
February 25, 1854; Bocourt, Journ. de Zool., 5:406-407, 1876.
_Tomodon lineatus_, Jan, Arch. Zool. Anat. Fis., Genoa,
2(2):234, March 1863; Elenco sistematico degli ofidi. Milano,
p. 57, 1863; Muller, Reisen in den Vereinigten Staaten,
Canada, und México. Bd. 3. Beitrage zur Geschichte, Statistik,
und Zoologie von Mexiko. 3:607, 1865; Jan and Sordelli,
Iconographie Generale des Ophidiens, Milano. liv. 19, pl. 6,
fig. 3, December, 1866; liv. 50, pl. 2, fig. 34, November,
1881.
_Tachymenis lineata_ (in part), Garman, Bull. Essex Inst., 16:
33, January 9, 1884; Mem. Mus. Comp. Zool., 8:60-61, July,
1884.
_Conophis lineatus_, Bocourt _in_ Duméril, Bocourt and
Mocquard, Mission Scientifique au Mexique et dans l'Amerique
Centrale, 2:643-644, pl. 38, fig. 5, 1886; Cope, Trans. Amer.
Philos. Soc., 18:218, pl. 28, fig. 2, (hemipenis), April 15,
1895; Boulenger, Catalogue of the Snakes in the British Museum
(Natural History), 3:122-123 (part), 1896; Cope, Ann. Rept. U.
S. Natl. Mus. for 1898, pp. 1094-1095, 1242, pl. 26, fig. 2,
(hemipenis), 1900; Amaral, Mem. Inst. Butantan, 4:212, 1929;
Mittleman, Copeia, no. 2:122, June 30, 1944.
_Conophis lineatus lineatus_, Smith, Journ. Washington Acad.
Sci., 31:122, March 15, 1941; Proc. U. S. Natl. Mus., 92:395,
November 5, 1942; Proc. U. S. Natl. Mus., 93:407, October 29,
1943; Smith and Taylor, Bull. U. S. Natl. Mus., 187:43,
October 5, 1945; Shannon and Smith, Trans. Kansas Acad. Sci.,
52:505, December 31, 1949; Smith and Taylor, Univ. Kansas Sci.
Bull., 33(pt. 2):351, March 20, 1950; Werler and Smith, Texas
Journ. Sci. 4(4):565, December 30, 1952; Fugler and Dixon,
Herpetologica, 14:186, December 1, 1958.
_Type._--Museum National d'Histoire Naturelle, Paris, no. 3738. Type
locality.--"México," restricted to Veracruz, Veracruz, México, by
Smith and Taylor (1950:351). Little is known about the type specimen,
and nothing, concerning its collector or the locality at which it was
collected. Smith (1941:122) assumed that the specimen illustrated by
Bocourt in Duméril, Bocourt, and Mocquard (1886:pl. 38, fig. 5) was
the type of _C. l. lineatus_. I have also made this assumption
concerning the identity of the type specimen of this species,
especially because of the many inconsistencies appearing in the plate
accompanying the description by Duméril, Bibron and Duméril (1854:pl.
73), and by Jan and Sordelli (1866:pl. 6). Neither show the nape nor a
regular number of dorsal scales by which accurate determination of
color pattern can be made and by means of which _C. l. dunni_ and _C.
l. lineatus_ can be separated.
_Diagnosis._--Lateral dark stripe anteriorly passing through eye and
posteriorly involving fourth scale-row only; first scale-row darkly
pigmented; no paravertebral stripe; no dark pigment on vertebral row;
six or eight dark stripes at mid-body, secondary stripes often present
posteriorly; usually eight (sometimes seven) supralabials immaculate
or having dark ventral margins.
_Variation._--Twenty-six specimens have 157 to 169 (163.5 ± 3.59)
ventrals. Twenty of these snakes having complete tails have 60 to 73
(66.5 ± 4.26) subcaudals; the number of ventrals plus subcaudals
varies from 224 to 238 (230.1) in nineteen of these. In 26 specimens
the reduction from 19 to 17 dorsal scale-rows takes place between
ventrals 91 and 107 (100.2 ± 3.59). Sexual dimorphism is evident in
the number of subcaudals; nine females have 60 to 66 (62.4), and 11
males have 68 to 73 (69.8) subcaudals. The largest specimen (AMNH
19643) is a male from "México," having a body length of 626 mm., a
tail length of 168 mm. and a total length of 786 mm. No small
juveniles have been examined; the smallest specimen (AMNH 19618) is a
male from Veracruz, México, having a body length of 325 mm., a tail
length of 90 mm. and a total length of 415 mm.
The greatest variation is in coloration. In preserved specimens the
ground-color is white, tannish-white, or often pale blue, with dark
stripes of black or deep brown present dorsolaterally and laterally.
Secondary stripes of paler brown are sometimes present, but the pale
browns have faded badly on many specimens. Normally four black stripes
are present at mid-body--a lateral pair on the 4th row of dorsal scales
and a dorsolateral pair on the 7th row (fig. 1, D). The lateral pair
is the posterior continuation of the stripe that on the head passes
through the eye; it continues on the nape as a narrow stripe on the
4th row only. In a few specimens the lateral stripe broadens to
include the upper third of the 3rd row posterior to the nape. In some
specimens both the dorsolateral and lateral dark stripes are present
on the nape as a row of elongated spots or dashes that become
continuous stripes of even width one-third to one-half of the distance
posteriorly along the body; in other specimens the stripes are
continuous on the nape. Posterior to the place of dorsal
scale-reduction from 19 to 17 rows by the fusion of the 3rd and 4th
rows, the lateral and dorsolateral stripes are moved downward by one
row. In some specimens secondary black or dark brown stripes are
present in the form of a series of dashes on the 5th and 8th rows;
posterior to the place of scale reduction, these dashes are on the 4th
and 7th rows. These dashes form a continuous stripe near the base of
the tail. On the tail the secondary and primary stripes on adjacent
rows sometimes fuse into a single broader stripe.
Usually the 1st row of dorsal scales is dark brown; in some specimens
the brown on the 1st or 7th row has faded in preservative. A few
specimens have small black spots on the moderate brown background of
the 1st row; in others the 1st row is only a somewhat darker brown
than the ground-color. The 2nd row sometimes is a medium brown, and
appears to be an additional stripe.
The ventrals usually have more or less conspicuous dark spots
laterally; in some specimens there are no spots. Except for the
lateral spots (when present) the ventrals are immaculate white. The
dorsal ground-color is pale brownish-white, white or pale blue between
the 4th and 7th rows of dorsal scales and dorsally between the 7th
rows on each side. Stripes are never present on the uniformly pale
colored 8th, 9th and vertebral scale-rows.
Usually there are eight supralabials on each side; however, seven of
the 27 specimens examined have seven supralabials on each side, and
three others have seven on one side, and eight on the other. Never is
more than the lower third of the supralabials dark brown. In many
specimens little or no brown is on the supralabials. There is little
or no brown on the chin.
Variation in coloration and in number of supralabials appears to be of
no geographic significance.
Although no juveniles have been collected, I expect that juveniles
resemble adults in coloration. Probably there would be a greater
contrast between the dark stripes and the pale ground-color in
juveniles.
In life an adult from three miles northwest of Lerdo de Tejada,
Veracruz, México (UMMZ 114484), had black stripes on the 4th and 7th
rows of dorsal scales, and black spots on a brown background on the
1st row. The 2nd row had a medial, pale to medium brown auxiliary
stripe on a brownish-white background. Posterior to the nape the 3rd
row was medium brown. The area between the 4th and 7th rows and the
dorsum between the 7th row of scales on each side was a pale
brownish-white. Posterior to the place of scale-reduction the primary
stripes were displaced downward by one row to the 3rd and 6th rows and
secondary stripes originated as elongated spots on the 4th and 7th
rows. Near the tail the secondary stripes were broad and continuous.
The head was white or tannish-white with three dark brown or black
stripes.
_Remarks._--In his diagnosis of _C. l. lineatus_, Smith (1941:122)
states: "lateral dark stripe ... very narrow posterior to nape,
extending along fourth scale row; posteriorly a stripe along third and
eighth (farther posteriorly the seventh) scale rows; a narrow dark
stripe along sixth scale row, continuous throughout length of
body...." I fail to find a dark stripe on the 6th row throughout the
length of the body. In all specimens that I have seen, there is a dark
stripe on the 7th row anteriorly and on the 6th row posteriorly. In
many specimens the stripes on the 3rd and 8th (posteriorly the 7th)
scale-rows are absent or present so far posteriorly that the 8th row
is never involved.
The dark brown on the first scale-row and the presence of a lateral
dark stripe on the 4th row of dorsal scales only, in combination with
the characteristics of the genus, distinguish _C. l. lineatus_ from
all other snakes in México.
_Distribution._--Semi-arid habitats on the coastal plain of Veracruz,
México, from Tecolutla to Lerdo de Tejada and Piedras Negras (fig. 2).
_Specimens examined._--Total of 27, as follows: MÉXICO: _no specific
locality_, AMNH 19614-15, 19621-24, 19642-43, NMW 16827. _Veracruz: no
specific locality_, AMNH 19618-20, CAS 73640, NMW 16829; _4 km. S
Alvarado_, KU 58124; _14 mi. N Alvarado_, UIMNH 46978; 6 mi. SE Boca
del Río, UIMNH 28023; Etiopa, 2 mi. S Tecolutla, UIMNH 3847; _ca._ 30
mi. E Jalapa, AMNH 81948; 3 mi. NW Lerdo de Tejada, UMMZ 114484-85;
Paso del Macho, USNM 109708; Río Blanco, 20 km. WNW Piedras Negras, KU
23253; Veracruz, AMNH 19612, UF 8990; _W side Veracruz_, AMNH 19616;
_2 mi. W Veracruz_, AMNH 19617, 19619.
=Conophis lineatus concolor= Cope
_Conophis vittatus_ Cope, Proc. Acad. Nat. Sci. Philadelphia,
13:300, December 28, 1861 (_nec_ Peters, 1860; type.--United
States National Museum, no. 4941; type locality--"Petén,"
Guatemala); Journ. Acad. Nat. Sci. Philadelphia, ser. 2,
8:137, 1876; Bull. U. S. Natl. Mus., 32:76, 1887.
_Conophis concolor_ Cope, Proc. Acad. Nat. Sci. Philadelphia,
18:318-319, February 20, 1867; Journ. Acad. Nat. Sci.
Philadelphia, ser. 2, 8:137, 1876; Bocourt _in_ Duméril,
Bocourt and Mocquard, Mission Scientifique au Mexique et dans
l'Amerique Centrale, 2:648, 1886; Müller, Verh. Ges. Basel,
8:263, 1887; Cope, Bull. U. S. Natl. Mus., 32:77; 1887; Ann.
Rept. U. S. Natl. Mus. for 1898, p. 1095, 1900; Schmidt and
Andrews, Zool. Ser. Field Mus. Nat. Hist., 20:178, October 31,
1936; Andrews, Zool. Ser. Field Mus. Nat. Hist., 20:358,
December 28, 1937; Smith, Occas. Papers Mus. Zool. Univ.
Michigan, 388:7, October 31, 1938; Taylor and Smith, Univ.
Kansas Sci. Bull., 25:253, July 10, 1939; Smith, Zool. Ser.
Field Mus. Nat. Hist., 24:31, January 30, 1939; Cochran, Bull.
U. S. Nat. Mus., 220:167, 1961; Neill and Allen,
Herpetologica, 17:44-46, fig. 3, April 15, 1961.
_Conophis lineatus_ (in part), Günther, Biologica
Centrali-Americana, p. 165, March, 1895; Gaige _in_ Pearse,
_et al._ Carnegie Inst. Washington Publ., 457:302, February 5,
1936.
_Conophis lineaticeps_ Cope, Ann. Rept. U. S. Natl. Mus. for
1898, pp. 1094-95, 1900 (Substitute name for _Conophis
vittatus_ Cope, 1861, _nec_ Peters, 1860).
_Conophis lineatus concolor_, Smith, Journ. Washington Acad.
Sci., 31:122-123, March 15, 1941; Proc. U. S. Natl. Mus.,
92:395, November 5, 1942; Proc. U. S. Natl. Mus., 93:407,
October 29, 1943; Smith and Taylor, Bull. U. S. Natl. Mus.,
187:43, October 5, 1945; Univ. Kansas Sci. Bull., 33(pt.
2):352, March 20, 1950.
_Types._--Two in the United States National Museum, no. 12368 (two
specimens). Type locality: "Yucatán," restricted to Chichén Itzá,
Yucatán, México by Smith and Taylor (1950:352).
_Diagnosis._--Dark stripes either absent posterior to the nape, or
present as a row of small spots on fourth or seventh scale-row; no
dark stripe on first scale-row; eight supralabials having dark ventral
margins.
_Variation._--Forty-five specimens have 158 to 170 (163.7 ± 1.56)
ventrals. Thirty-eight of these snakes having complete tails have 56
to 74 (66.7 ± 4.77) subcaudals; the number of ventrals plus subcaudals
varies from 222 to 245 (230.6). In 45 specimens the reduction from 19
to 17 dorsal scales takes place between ventrals 89 and 114 (102.5 ±
5.57). Sexual dimorphism is evident in the number of subcaudals; 16
females have 56 to 65 (61.8), and 22 males have 68 to 74 (70.3)
subcaudals. The longest specimen (USNM 46395) is a male from Chichén
Itzá, Yucatán, having a body length of 893 mm., a tail length of 274
mm., and a total length of 1167 mm. A juvenile (AMNH 38833) from
Chichén Itzá, Yucatán, has a body length of 194 mm., a tail length of
50 mm., and a total length of 244 mm.
The venter is immaculate white or pale yellow and the dorsum of the
body is immaculate pale gray to pale olive. Some specimens have small
dark brown spots on the tips of the scales of the 4th or of the 7th
row, but never on both. Only on the nape are spots present on both the
4th and the 7th rows; these spots are the posterior continuations of
the dark stripes on the head and on many specimens do not reach the
nape. Posterior to the place of scale reduction from 19 to 17 rows by
the fusion of the 3rd and 4th rows of scales, the dark spots (when
present) are on the 3rd or 6th row of scales.
The coloration of juveniles is the same as that of adults. Color in
life is thought not too different from that of preserved specimens,
for notes on the color of living individuals (Neill and Allen,
1961:44) agree with what I have observed on preserved snakes.
_Remarks._--The specimen from "Petén" (USNM, no. 4941) is the only
specimen that has a controversial history. As can be seen from the
synonymy of the species, the relationship of this specimen with the
rest of the genus has been interpreted in several ways. Smith
(1941:122-123) stated that the above specimen was catalogued as being
from El Salvador; however, the locality was presumed by him to be El
Petén, Guatemala, due to the presence in the bottle of a piece of
paper inscribed "_Conophis vittatus_, Petén, J. M. Dow." This specimen
is the one mentioned by Cope (1861:300, 1876:76, and 1900:1094-95),
and in the first paper is ascribed to Guatemala. In 1900 this specimen
was named _C. lineaticeps_ by Cope who thought the specimen differed
significantly from _C. concolor_ (Cope, 1867:318-319). This specimen
has the coloration normal for _C. l. concolor_ as far posteriorly as
mid-body; beyond mid-body the dark lines, typical of _C. l. lineatus_
or of _C. l. dunni_, are present. It is likely that this specimen is
an intergrade between _C. l. concolor_ and _C. l. dunni_, the other
subspecies present in Guatemala.
The only specimen not from the Yucatán Peninsula is allegedly from
Patuca, Honduras (USNM 20271). It was obtained in the 1870's. Possibly
more collecting will verify the presence of _C. l. concolor_ in
northern Honduras. This individual may be merely a genetically
aberrant specimen from an area where normal specimens are _C. l.
dunni_. Neill and Allen (1961:44-45) suggested that the specimen from
Patuca implies widely overlapping distributions for _C. l. dunni_ and
_C. concolor_. The occurrence of _C. l. concolor_ in Honduras needs to
be verified before this assumption is made. There can, therefore, at
present be no objection to the view that intergradation between the
subspecies _C. l. dunni_ and _C. l. concolor_ could occur through a
relatively broad area of El Petén and British Honduras.
Neill and Allen (1961:44-45) further suggest that the present range of
_C. l. dunni_ extends "presumably still farther northward toward the
Méxican state of Veracruz where _C. l. lineatus_ exists." Actually the
presence of the subspecies _C. l. dunni_ and _C. l. lineatus_ as
presently disjunct populations implies merely that they were
presumably a continuous population at some time in the past.
The characteristics of the genus in combination with the reduction of
dark coloration posterior to the head distinguish this snake from all
other snakes in México and Central America.
_Distribution._--The Yucatán Peninsula: eastern Campeche, all of
Yucatán, probably in Quintana Roo, and the northern third of British
Honduras. A record for northeastern Honduras is questioned (fig. 2).
_Specimens examined._--Total of 48, as follows: BRITISH HONDURAS:
_Belize District_: 13.0 mi. W, 1.5 mi. S Belize, ERA-WTN BH-1562.
GUATEMALA: _El Petén, no specific locality_, USNM 4941.
HONDURAS: _Colón_: Patuca, USNM 20271.
MÉXICO: _Campeche_: Champotón, UMMZ 73063-66; Encarnación, CNHM
106462. _Yucatán: no specific locality_, BMNH 80.7.13.30; Chichén
Itzá, AMNH 38826, 38833, CNHM 20610-11, 26986-87, 36299-300, 36303-04,
36307, 36316, MCZ 7422, 28748, UMMZ 68236, 73060-62, 80806, USNM
46395; Kantunil, CNHM 36301, 36305-06, 36308-09, 36312-13; _Libré
Union_, CNHM 36298, 36302, 36310-11, 36314; Mayapán, CNHM 40720;
Mérida, CNHM 19411, 19413, NMW 16828; Progreso, CNHM 40721; Tekom,
CNHM 49374; Yokdzonot, CNHM 36315.
=Conophis nevermanni= Dunn
_Coniophanes imperialis imperialis_, Wettstein, Sitz. Akad.
Wiss. Wien. mathem-naturw. Kl., 143:37-38, 1934.
_Conophis nevermanni_ Dunn, Copeia, no. 4:214, December 31,
1937; Smith, Proc. U. S. Natl. Mus., 92:395, November 5, 1942;
Savage, Trans. Kansas Acad. Sci., 50:484, December 31, 1949;
Taylor, Univ. Kansas Sci. Bull., 34(pt. 1): 145-146, October
1, 1951.
_Type._--Academy of Natural Sciences of Philadelphia, no. 22423,
obtained by Emmet R. Dunn from Prof. Manuel Valerio. Type locality:
Río Poas de Aserri (a few miles south of San José), Costa Rica.
_Diagnosis._--Head and body dark brown or black above with two or four
white stripes along body; usually two white lines on head immediately
above eye passing from canthus rosetralis posteriorly to connect with
white stripe on 6th row of dorsal scales; eight supralabials with
black margins above.
_Variation._--Six specimens have 173 to 183 (176.5 ± 4.00) ventrals.
Five of these snakes having complete tails have 71 to 89 (80.6 ± 7.15)
subcaudals; the number of ventrals plus subcaudals varies from 250 to
263 (257.0). In the six specimens the reduction from 19 to 17 dorsal
scales takes place between ventrals 84 and 97 (93.2 ± 4.71). Sexual
dimorphism is evident in the number of subcaudals; two females have 71
and 76 (73.5), and three males have 82 to 89 (85.3) subcaudals. The
longest specimen (ANSP 22424) is a female from San José, Costa Rica,
having a body length of 660 mm., a tail length of 168 mm. and a total
length of 828 mm.
The dorsal coloration (fig. 1, E) varies from a black ground-color
with two or four narrow white stripes to a dark brown ground-color
with a series of black stripes and four white stripes. In the black
specimens there are no dark stripes. The darkest specimen (NMW
16838:1) has only two white stripes; these more or less continuous
stripes are on the ventral third of the 2nd row of scales and
occasionally on the dorsalmost part of the first scale-row. The venter
is immaculate white except for black on the tips of the ventral
scales. The dorsum above the 2nd scale-row is uniform black. There are
no white stripes on the head.
The palest specimen (NMW 16838:2) has four dorsal white stripes; the
lateral pair of these stripes is on the ventral half of the 2nd and
the dorsal third of the 1st scale-rows; the dorsolateral pair is on
the dorsal two-thirds of the 6th and the ventral third of the 7th rows
of scales. This latter stripe is the posterior continuation of the
white stripe on the head, which originates immediately posterior to
the rostral scale and passes posteriorly along the canthus rostralis
and along the lateral margin of the supraocular scale to the nape.
Posterior to the place of scale reduction, the dorsolateral white
stripe is displaced ventrally one scale-row. Except for black flecks
or spots on the lateral margins of the ventrals, the venter is
immaculate white. The dorsum above the lateral white stripes is brown
and black; there is a pair of dorsolateral white stripes. The dorsal
half of the 2nd, most of the 3rd, 4th and 5th rows of scales are
black; the dorsal margin of the 3rd, both margins of the 4th, and the
ventral margin of the 5th rows are paler brown. The dorsal two-thirds
of the 7th, all but the dorsal most part of the 8th, and the middle
two-thirds of the 10th scale-rows are black; the areas between are a
medium brown.
Only six specimens are available on which to base a description of the
variation in this species. Furthermore, there are no juveniles, notes
on the colors of living individuals, or photographs of this species.
[Illustration: FIG. 4. Selected locality records for _Conophis
pulcher_ and _Conophis nevermanni_.]
_Remarks._--Taylor (1955:563-565) hesitantly referred a specimen (KU
35630) from 32 kilometers north of Barranca, Puntarenas Province,
Costa Rica, to _Conophis lineatus nevermanni_. This specimen, a
female, has 169 ventrals and ventral scale-reduction taking place
opposite the 109th ventral; both of these characters are well out of
the range of _C. nevermanni_. Furthermore, the ventral margins of the
supralabials are brown, and the pale dorsal stripes are tan and too
wide for _C. nevermanni_ (compare figs. 1, C and E). The specimen
definitely is _C. lineatus dunni_, and corresponds well with another
specimen from Costa Rica (ANSP 12232).
The dark brown or black dorsum with two or four white stripes and the
presence of eight supralabials having dark brown dorsal margins, in
combination with the characters of the genus, serve to distinguish
_Conophis nevermanni_ from other Central American snakes.
_Distribution._--Pacific coastal plain of northwestern Costa Rica and
the Meseta Central of central Costa Rica (fig. 4).
_Specimens examined._--Total of six, as follows: COSTA RICA:
_Guanacaste_: Bebedero, Río Tenorio, NMW 16838(5). "_San José_," ANSP
22424.
=Conophis pulcher= Cope
_Tomodon lineatus_ (in part), Salvin, Proc. Zool. Soc. London,
28:455, 1860.
_Conophis pulcher_ Cope, Proc. Acad. Nat. Sci. Philadelphia,
20(5):308, 1869; Journ. Acad. Nat. Sci. Philadelphia, ser. 2,
8:137, 1876; Bocourt _in_ Duméril, Bocourt and Mocquard,
Mission Scientifique au Mexique et dans l'Amerique Centrale,
2:646-648, pl. 38, fig. 6, 1886; Ferrai-Perez, Proc. U. S.
Natl. Mus., p. 196, September 28, 1886; Cope, Bull. U. S.
Natl. Mus., 32:77, 1887; Trans. Amer. Philos. Soc., 18:194,
April 15, 1895; Ann. Rept. U. S. Natl. Mus. for 1898, p. 1095,
1900; Alvarez del Toro, Reptiles de Chiapas, pp. 154-155,
1960.
_Tomodon pulcher_, Bocourt, Journ. de Zool., p. 408, 1876.
_Conophis pulcher_ var. _similis_ Bocourt _in_ Duméril,
Bocourt and Mocquard, Mission Scientifique au Mexique et dans
l'Amerique Centrale, 2:647-648, pl. 38, fig. 6, 1886
[Type.--Museum National d'Histoire Naturelle, Paris, no. 6090;
type locality.--unknown, restricted to Tonalá, Chiapas, by
Smith and Taylor (1950:326)].
_Conophis lineatus_, Günther, Biologia Centrali-Americana, p.
165, March, 1895; Boulenger, Catalogue of the Snakes in the
British Museum (Natural History), 3:122-123, 1896; Stuart,
Occas. Papers Mus. Zool. Univ. Michigan, 292:5, June 29, 1934;
Slevin, Proc. California Acad. Sci. 4th Ser., 23:409, December
29, 1939.
_Conophis pulcher pulcher_, Smith, Journ. Washington Acad.
Sci., 31:121, March 15, 1941; Proc. U. S. Natl. Mus., 92:395,
November 5, 1942; Stuart, Contr. Lab. Vert. Biol. Univ.
Michigan, 65:19-20 (part), March, 1954; Contr. Lab. Vert.
Biol. Univ. Michigan, 68:63, November, 1954; Cochran, Bull. U.
S. Natl. Mus., 220:167, 1961.
_Conophis pulcher plagosus_ Smith, Journ. Washington Acad.
Sci. 31:121-122, March 15, 1941 (Type.--United States National
Museum, no. 109707; type locality: Tonalá, Chiapas); Smith and
Taylor, Univ. Kansas Sci. Bull., 33(pt. 2):326, March 20,
1950; Stuart, Contr. Lab. Vert. Biol. Univ. Michigan,
65:19-20, March, 1954; Cochran, Bull. U. S. Natl. Mus.,
220:167, 1961.
_Conophis pulcher similis_, Smith, Proc. U. S. Natl. Mus.,
92:395, November 5, 1942; Proc. U. S. Natl. Mus., 93:408,
October 29, 1943; Smith and Taylor, Bull. U. S. Natl. Mus.,
187:43-44, October 5, 1945; Univ. Kansas Sci. Bull., 33(pt.
2):43-44, March 20, 1950; Maldonado-Koerdell, Inst. Mexicanos
Recursos Nat. Renov. pp. 132-133, 1953.
_Types._--Three in the United States National Museum, nos. 6751 (2
specimens) and 6803, obtained by Henery Hague. Type locality: "Petén,"
or "Verapaz," Guatemala. There is much doubt about localities for many
of Hague's specimens collected in the 1860's (Stuart, 1948:10). Since
_Conophis pulcher_ is found predominantly in semi-arid environments,
the types might have come from the semi-arid Cahabón, Negro, or Salamá
river basins--all places near the sugar plantation that Hague managed
at San Jerónimo, Baja Verapaz. Possibly the types were obtained from
as far away as the Motagua Valley or the southeastern highlands of
Guatemala, both of which areas Hague is known to have visited.
_Diagnosis._--Paravertebral stripes present at least posteriorly (fig.
1, F); eight or ten stripes at mid-body; lateral dark stripe passing
through eye anteriorly and including at least upper one-half of second
scale-row from neck region posteriorly to place of scale reduction
near mid-body; eight supralabials immaculate or having dark ventral
margins.
_Variation._--Twenty-six specimens have 161 to 182 (169.5 ± 5.31)
ventrals. Eighteen of these snakes with complete tails have 65 to 79
(70.6 ± 3.93) subcaudals; the number of ventrals plus subcaudals
varies from 231 to 251 (239.3). In 26 specimens the reduction from 19
to 17 dorsal scales takes place between ventrals 94 and 119 (104.6 ±
4.90). Sexual dimorphism is evident in the number of subcaudals;
eleven females have 65 to 71 (68.2), and seven males have 70 to 79
(74.3) subcaudals. The longest specimen (AMNH 58364) is a female from
El Zamarano, Honduras, having a body length of 703 mm., a tail length
of 164 mm. and a total length of 867 mm. The smallest juvenile (MCZ
49793) from Tegucigalpa, Honduras, has a body length of 162 mm., a
tail length of 46 mm. and a total length of 208 mm.
The dorsal ground-color is pale brown or white; black or dark brown
stripes are present dorsally and laterally. Normally ten stripes are
present at mid-body; the first pair on the first row of dorsal scales;
the second pair on the upper half of 2nd and lower part of 3rd rows;
the third pair on 4th row; the fourth pair on 7th and sometimes part
of 8th rows; the fifth pair (paravertebral stripes) on the 9th row.
Posterior to the place of reduction from 19 to 17 rows by the fusion
of the 3rd and 4th rows, the third, fourth and fifth pairs of stripes
are displaced downward one row. Sometimes the second and third pairs
of stripes are fused resulting in only eight stripes at mid-body. On
some specimens the fourth and fifth pairs of stripes are close
together, but in none are they fused so as to result in a pattern of
six stripes at mid-body.
The paravertebral stripes begin anteriorly on the nape or at any point
on the anterior one-third of the body and continue as discrete stripes
onto the base of the tail. Anteriorly these stripes are always broken
into a series of dashes; posteriorly the stripes are continuous. In
specimens in which the paravertebral stripes do not begin on the
anterior-most part of the body, there is no paravertebral pigmentation
anteriorly.
In addition to the paravertebrals, the other dorsal dark stripes are
variable. In some specimens the stripes are present anteriorly and
gradually disappear near mid-body (the first dark stripe only on three
specimens). In other specimens the stripes are present anteriorly as
dashes and become continuous at mid-body; in others the stripes are
continuous throughout. Posteriorly continuous stripes are of uniform
width; anteriorly sometimes they are wide on the tip of each scale and
narrow on the base (fig. 1, F). The variation in continuity and width
described above is found in all of the dorsal dark stripes.
The ventrals usually have more or less conspicuous dark spots
laterally; in some specimens there are no spots. Except for the dark
lateral spots, when present, the ventrals are immaculate white.
Usually the dorsal ground-color is a pale tan, especially between the
first and second, and the third and fourth dark stripes. The areas
between the second and third dark stripes and across the dorsum
between the fourth stripes on each side are pale brown. In some
specimens the dorsum between the paravertebral stripes is still paler
brown.
Never is more than the lower third of the supralabials brown. Many
specimens have little brown, and others none. In most of those
specimens having brown on the supralabials, the chin and infralabials
are dusky tan or gray. There is little or no brown on the supralabials
or the chin in the northern part of the range (Chiapas), whereas the
greatest amount of brown on the labials and chin is found on some
specimens from the southern part of the range (Honduras). Since there
is considerable variation in the amount of brown on the chin and
labials of specimens from single localities, the slight geographic
trend in this character seemingly is not significant.
In juveniles six black or dark brown stripes boldly contrast with a
white or pale tan ground-color. At mid-body the first pair of dark
stripes is on the 1st scale row; the second pair on the 3rd and 4th
rows; the third pair on the 7th, 8th and at least the lower half of
the 9th rows (fig. 3, B). Ontogenetic change in coloration consists of
the splitting of the second and third pairs of dark stripes in the
juvenile. The first stripe does not split. Consequently adults have
ten dark stripes.
In life an adult from Tonalá, Chiapas, had black stripes. The
ground-color below the second stripe, and between the third and fourth
dark stripes was tan. The area between the second and third dark
stripes was reddish-brown, as was the dorsum between the fourth pair
of dark stripes, except that the 10th scale-row was paler.
Three excellent photographs of this species have been published under
the name _Conophis lineatus_ (Ditmars, 1931:pls. 26 and 27).
_Remarks._--Smith (1941:121-122) described _C. pulcher plagosus_ from
Tonalá, Chiapas, and characterized the subspecies by its having "(1)
the ventrals completely unspotted; (2) secondary lines on
paravertebral rows not continuous posteriorly; (3) all other lines on
body also somewhat spotted in appearance; (4) dusky markings on chin
and supralabial border very dim (less distinct than in _p. pulcher_ or
any member of the _lineatus_ series)." Although all Chiapan specimens
lack ventral spots, specimens from Guatemala have no spots, small
spots, or large spots. Even in specimens from Tegucigalpa, Honduras,
the southernmost limit of the range, the spotting varies from a few
inconspicuous spots to many large spots. Paravertebral rows were
continuous posteriorly in all specimens examined by me. Likewise, all
other stripes were continuous bands of uniform width posteriorly,
having appeared anteriorly as rows of spots or dashes. The amount of
brown on the chin and labials has been shown previously not to be
geographically significant. The absence of characters of adequate
significance to separate populations precludes the naming of
subspecies in this species.
Mertens (1952a:93, and 1952b:61-62) designated three specimens from El
Salvador as _C. pulcher plagosus_. In the latter paper, Mertens, on
the basis of a description of a specimen of "_C. lineatus_" from
Divisadero, El Salvador, given by Schmidt (1928:200), referred that
specimen also to _C. pulcher plagosus_. I have examined this specimen
and refer it to _C. lineatus dunni_. Although I have not seen Merten's
specimens, on the basis of the excellent descriptions given by Mertens
(1952b:61-62), I refer the three Salvadoranean specimens to _C.
lineatus dunni_.
The presence of paravertebral stripes in combination with the
characteristics of the genus distinguish _Conophis pulcher_ from all
other snakes in southern México and Central America. The only
sympatric species of this genus, _C. lineatus dunni_, differs in that
it lacks paravertebral stripes, although it may have a single
vertebral stripe. _Conophis lineatus dunni_ has lateral dark stripes
that are present on the 3rd and 4th scale-rows, never on the anterior
third of the body as in _C. pulcher_. Even in juveniles the third pair
of dark stripes includes the lower part of the 9th scale-row in _C.
pulcher_, whereas the dorsal most dark stripe of _C. lineatus dunni_
never includes more than the lower part of the 8th scale-row.
_Distribution._--Pacific coastal region of Chiapas, México,
southeastward into Guatemala; southeastern highlands and the dry
valley of central and eastern Guatemala; Caribbean lowlands of
Honduras southward to the region of Tegucigalpa, Honduras (fig. 4).
_Specimens examined._--Total of 27, as follows: GUATEMALA: _no
specific locality_, CNHM 22912, NMW 16830. _Jutiapa_: Hacienda Mongoy,
UMMZ 106725. _El Progreso_: El Progreso, CAS 67000; _El Rancho_, UMMZ
106724; _San Antonio_, CAS 66999. "Peten," USNM 6751(2), 6803.
_Sacatepequez_: Dueñas, BMNH 64.1.26.17, 64.1.26.126-127. _Zacapa_:
Pepesca, AMNH 72555-56.
HONDURAS: _no specific locality_, AMNH 58364. _Cortes_: San Pedro
Sula, CNHM 5295-96. _Francisco Morazan: El Zamarano_, AMNH 70189;
Tegucigalpa, MCZ 49785, 49787-88, 49791, 49793, 49795.
MÉXICO: _Chiapas_: _Soconusco_, UIMNH 33646-47; Tonalá, USNM 109707.
=Conophis vittatus= Peters
_Tomodon lineatum_ (in part), Duméril, Bibron and Duméril,
Érpétologie Genérale, 7(pt. 2):936-938, February 25, 1854.
_Conophis vittatus_ Peters, Monatsb. Akad. Wiss. Berlin, pp.
519-520, pl., fig. 3, October, 1860; Cope, Proc. Amer.
Philos. Soc., 11:162, 1870; Bocourt _in_ Duméril, Bocourt and
Mocquard, Mission Scientifique au Mexique et dans l'Amerique
Centrale, 2:644-646, pl. 38, fig. 7, 1886; Günther, Biologia
Centrali-Americana, p. 165, March, 1895; Boulenger, Catalogue
of the Snakes in the British Museum (Natural History),
3:123-124, 1896; Cope, Amer. Nat., 30:1024, 1896; Ann. Rept.
U. S. Natl. Mus. for 1898, pp. 1094-1095, 1232, 1900; Gadow,
Proc. Zool. Soc. London, 2:225, 1905; Amaral, Mem. Inst.
Butantan, 4:211, 1929; Gadow, Jorullo, p. 55, 1930; Smith,
Zool. Ser. Field Mus. Nat. Hist., 24:31-32, January 30, 1939;
Taylor and Smith, Univ. Kansas Sci. Bull., 25:252-253, pl. 23,
July 10, 1939; Stuart, Contr. Lab. Vert. Biol. Univ. Michigan,
65:23, March, 1954; Alvarez del Toro, Reptiles de Chiapas, pp.
153-154, 1960.
_Conophis lineatus_ Cope, Proc. Acad. Nat. Sci. Philadelphia,
16(3):167, 1864 [_nec_ Duméril, Bibron and Duméril,
Érpétologie Genérale, 7(pt. 2):936-938, atlas, pl. 73,
February 25, 1854; specimen from Colima]; Sumichrast, Arch.
Sci. Nat., p. 246, 1873.
_Tomodon vittatus_, Bocourt, Journ. de Zool., p. 407, 1876.
_Conophis sumichrasti sumichrasti_ Cope, Journ. Acad. Nat.
Sci. Philadelphia, ser. 2, 8:137, 1876 (Types.--United
States National Museum, nos. 29123, 30258; type
locality.--Tehuantepec, México); Bull. U. S. Natl. Mus.,
32:77, 1887; Smith and Taylor, Univ. Kansas Sci. Bull., 33(pt.
2):334, March 20, 1950; Maldonado-Koerdell, Inst. Mexicanos
Recursos Nat. Renov., p. 124, 1953.
_Conophis sumichrasti viduus_ Cope, Journ. Acad. Nat. Sci.,
Philadelphia, ser. 2, 8:137, 1876 (Type.--United States
National Museum, no. 30259; type locality.--Tehuantepec,
México); Bull. U. S. Natl. Mus., 32:77, 1887; Cochran,
Bull. U. S. Natl. Mus., 220:167, 1961.
_Conophis sumichrasti_, Cope, Proc. Amer. Philos. Soc.,
18:271, August 11, 1879; Sumichrast, Bull. Soc. Zool. France,
p. 182, 1880; Cope, Trans. Amer. Philos. Soc., 18:194, April
15, 1895; Cochran, Bull. U. S. Natl. Mus., 220:167, 1961.
_Tachymenis lineata_ (in part), Garman, Mem. Mus. Comp. Zool.,
8:60-61, July, 1884.
_Conophis vittatus sumichrasti_, Cope, Ann. Rept. U. S. Natl.
Mus. for 1898, p. 1095, 1900.
_Conophis vittatus videns_ Cope, Ann. Rept. U. S. Natl. Mus.,
for 1898, p. 1095, 1900 (apparent _lapus_ for _viduus_).
_Conophis vittatus vittatus_, Cope, Ann. Rept. U. S. Natl.
Mus. for 1898, p. 1095, 1900; Smith, Journ. Washington Acad.
Sci., 31:119-120, March 15, 1941; Proc. U. S. Natl. Mus.,
92:395, November 5, 1942; Proc. U. S. Natl. Mus., 93:408,
October 29, 1943; Ann. Carnegie Mus., 30:91, November 2, 1944;
Smith and Taylor, Bull. U. S. Natl. Mus., 187:44, October 5,
1945; Smith, Rev. Soc. Mexicanos Hist. Nat., 7:71, December,
1946; Smith and Taylor, Univ. Kansas Sci. Bull., 33(pt.
2):331, March 20, 1950; Davis and Smith, Herpetologica, 8:134,
January 30, 1953; Maldonado-Koerdell, Inst. Mexicanos Recursos
Nat. Renov., p. 130, 1953; Peters, Occas. Papers Mus. Zool.
Univ. Michigan, 554:22, June 23, 1954; Duellman, Occas. Papers
Mus. Zool. Univ. Michigan, 560:15, October 22, 1954; Webb and
Fugler, Herpetologica, 13:35, March 30, 1957; Duellman, Occas.
Papers Mus. Zool. Univ. Michigan, 589:15, March 21, 1958;
Zweifel, Amer. Mus. Novitates, 1949:2, 5, June 17, 1959;
Duellman, Univ. Kansas Publ. Mus. Nat. Hist., 15(1):91-92,
December 20, 1961.
_Conophis vittata_, Gadow, Proc. Zool. Soc. London, 2:196,
1905; Through Southern México, p. 181, 1908.
_Conophis viduus_, Smith, Zool. Ser. Field Mus. Nat. Hist.,
24:31, January 30, 1939; Hartweg and Oliver, Misc. Publ. Mus.
Zool. Univ. Michigan, 47:26-27, July 13, 1940.
_Conophis vittatus viduus_, Smith, Journ. Washington Acad.
Sci., 31:120-121, March 15, 1941; Proc. U. S. Natl. Mus.,
92:395, November 5, 1942; Proc. U. S. Natl. Mus., 93:408,
October 29, 1943; Woodbury and Woodbury, Journ. Washington
Acad. Sci., 34(11):370, 1944; Smith and Taylor, Proc. U. S.
Natl. Mus., 187:44, October 5, 1945; Univ. Kansas Sci. Bull.,
33(pt. 2):340, March 20, 1950; Werler and Smith, Texas Journ.
Sci., 4:565, fig. 16, December 30, 1952; Maldonado-Koerdell,
Inst. Mexicanos Recursos Nat. Renov., p. 130, 1953; Davis and
Dixon, Proc. Biol. Soc. Washington, 72:82-83, July 24, 1959.
_Conophis vittatus vittatus_ × _Conophis vittatus viduus_,
Alvarez del Toro and Smith, Herpetologica, 12:13, March 6,
1956.
_Type._--Zoologisches Museum Berlin. Type locality not given, for the
specimen was purchased from a dealer in Hamburg. The type locality was
first restricted to "Acapulco," Guerrero, by Smith (1941:119), then to
Laguna Coyuca, Guerrero, México, by Smith and Taylor (1950:331).
_Diagnosis._--Three or four dorsal dark stripes, each involving two or
more adjacent scale-rows; never having brown or black on the 1st
scale-row; seven supralabials immaculate white or pale tannish-white.
_Variation._--One hundred seventy-one specimens have 149 to 181 (163.7
± 6.33) ventrals. One hundred fifty-three of these having complete
tails have 55 to 76 (64.8 ± 4.90) subcaudals; the number of ventrals
plus subcaudals varies from 214 to 245 (228.5). In 170 specimens the
reduction from 19 to 17 dorsal scales takes place between ventrals 84
and 118 (102.3 ± 6.60). Sexual dimorphism is evident in the number of
subcaudals; 58 females have 55 to 66 (60.0) and 95 males have 59 to 76
(67.8) subcaudals. The longest specimen (AMNH 68004) is a male from
Escurano, Oaxaca, México, having a body length of 668 mm., a tail
length of 182 mm. and a total length of 850 mm. A juvenile (CNHM
40435) from Tehuantepec, Oaxaca, México, has a body length of 133 mm.,
a tail length of 31 mm. and a total length of 164 mm.
Variation in coloration is of such magnitude that it has been used as
the basis for recognition of subspecies. Unfortunately, until this
time, most specimens reported upon in the literature represented the
two extremes of variation. After examining the coloration of 174
specimens with respect to geographic distribution, I conclude that
only one highly variable species is represented. Specimens from the
northern and western parts of the range (Michoacán, Colima, and
Durango) have the color pattern of _C. vittatus_ as described by
Peters (1860:518-521); these snakes have four narrow black stripes on
a white or pale tan background, and an immaculate white venter. The
lateral dark stripe, which on the head passes through the eye, is
present on the dorsal half of the 3rd and the ventral half of the 4th
scale-rows; the dorsolateral dark stripe, which passes along the
middle of the head and splits on the nape, is present on the middle of
the 8th scale-row. The other extreme in color pattern consists of
three broad stripes; the two dorsolateral stripes are fused. This
pattern is prevalent in specimens from the area around Tehuantepec,
Oaxaca. The lateral stripes include the dorsal half to two-thirds of
the 2nd, all of the 3rd and 4th, and half of the 5th scale-rows; the
fused dorsolateral stripes sometimes cover all of the area dorsal to
and including the dorsal third of the 7th scale-row.
Snakes from areas between Tehuantepec and the margins of the
distribution of this species are variously intermediate between the
extremes described above. In some snakes from these areas the lateral
stripes are broad and include either the dorsal half of the 2nd
scale-row or the ventral half of the 5th scale-row, but not both on
the same specimen. Also, the dorsolateral stripes are broad and
include most of the 9th and a part of the 10th scale-rows. Many
specimens from the area around Tehuantepec, where the three-striped
pattern is prevalent, have an intermediate pattern. Some have white on
the center of the 10th scale-row or lateral stripes that are not so
broad as to include the 3rd and 4th and half of each of the 2nd and
5th scale-rows.
The supralabials are immaculate white or pale tan, except that in some
specimens the dorsalmost part of some supralabials are dark brown or
black as they are included in the ventral boundary of the dark stripe
that passes through the eye. There are no dusky markings on the chin
or on any of the ventral scales.
There is no ontogenetic change in color pattern; juveniles have the
same coloration as adults from the same geographic area.
Color in life is not greatly different from that of preserved
specimens. One specimen (UMMZ 114483) from 10.8 miles south of Oaxaca,
had in life black stripes, a pale yellowish tan dorsal ground-color
and a pale off-white venter.
An excellent photograph of this species appears in Schmidt and Inger
(1957:230) under the name _Conophis lineatus_.
_Remarks._--I have been unable to find variation of geographic
importance in scutellation in this species. A wide range of variation
in the characters of scutellation is present in specimens from most
localities; it shows no significant clinal or geographic trends. As I
have stated previously, in the discussion of variation, coloration has
been the feature primarily used by previous workers to distinguish two
"subspecies" for this species; _C. vittatus vittatus_ having four
black stripes and _C. vittatus viduus_ having three black stripes.
Most of the three-striped snakes occur in the vicinity of Tehuantepec,
Oaxaca, whereas the four-striped snakes are found near the margins of
the range of the species in Durango, Colima, Michoacán, Morelos and
Puebla. Specimens that would have to be considered intergrades between
the "subspecies" are found in Michoacán, Guerrero, Oaxaca and Chiapas.
At the time the subspecies were proposed only specimens from
Tehuantepec or from marginal areas were known. Utilizing the large
number of specimens of this species presently available, geographic
variation is found to be clinal, from those with three stripes from
near Tehuantepec, through several intermediate patterns present on
specimens from single localities in Guerrero, Oaxaca and Chiapas, to
those with four dark stripes in areas farthest removed to the north
and west from Tehuantepec. Since only coloration shows geographic
variation, and since this variation represents a continuous cline,
subspecies cannot be recognized for this species.
The presence and position of the three or four dark stripes on the
body and the absence of brown on the 1st scale-row or on the ventral
scales, in combination with the generic characters, distinguish
_Conophis vittatus_ from all other Méxican snakes. The only other
snake that occurs in western México that has been confused with _C.
vittatus_ is _Coniophanes piceivittus taylori_, which has 25, instead
of 19, scale-rows.
_Distribution._--Semi-arid habitats on Pacific slopes from extreme
southern Durango southeastward to Tuxtla Gutierrez, Chiapas, and
inland in the eastern Balsas Basin to Morelos and western Puebla
(fig. 5).
[Illustration: FIG. 5. Selected locality records for _Conophis
vittatus_.]
_Specimens examined._--Total of 174, as follows: MÉXICO: _no specific
locality_, AMNH 66150-52, SU 9465. _Chiapas_: Piedra Parada, USNM
121453. _Pizo de Oro_, UIMNH 40821. Tuxtla Gutierrez, Parque Madero,
UIMNH 37992-93, 38036-37. _Colima: no specific locality_, MCZ 46860,
USNM 31394, 31396-97. 1 mi. SW Colima, AMNH 12783. S of Manzanillo,
AMNH 19641. _Durango_: Hacienda de Gabriel, AMNH 14217. _Guerrero:
Acahuizotla_, TCWC 7419, 9469. _1 mi. W Acahuizotla_, TCWC 7418. 3 mi.
W Acapulco, AMNH 71626. _6 mi. E Acapulco_, TCWC 9476-77. _10 mi. S
Acapulco_, TCWC 8578. _Agua del Obispo_, CNHM 104948, TCWC 11586. near
Chilpancingo, MVZ 45067, UMMZ 85722-23. _1 mi. SW Colotlipa_, TCWC
9471-74. _2 mi. SW Colotlipa_, TCWC 9475. 14 mi. S Ixtapán de la Sal,
KU 67648. _Laguna Coyuca_, CNHM 25881, UMMZ 80942. near La Unión, AMNH
66337. _Magueyes, Laguna Coyuca_, AMNH 66149. _Playa Encantada_, TCWC
9470. 1 mi. S Tierra Colorada, KU 67649. near _Xaltinanguis, km. 405_,
CNHM 104947. _Michoacán_: Coalcomán, UMMZ 104693. _1/2 mi. SE
Coalcomán_, UMMZ 104492. _1 mi. N. Coalcomán_, UMMZ 112543. _1 mi. NE
Coalcomán_, UMMZ 104692. Puerta de la Playa, UMMZ 105155. 12 mi. S
Tzitzio (by road), UMMZ 99153. _Morelos: 12 km. NW Axochiapan_, TCWC
7311, UIMNH 17613, 25924. 7 mi. SE Cuernavaca, MVZ 32258. _Huajintlán,
km. 133_, CNHM 103270. 12 km. S Puente de Ixtla, km. 133, CNHM 104949.
_Oaxaca: Bisiliana_, AMNH 68010. _near Caoba, foot of Cerro Arenal_,
AMNH 68009. _Cerro Arenal_, AMNH 68000-03. _Cerro de Laollaga_, UIMNH
36213. _Cerro de San Pedro_, UIMNH 17616. _Cerro Palma de Oro_, UIMNH
37116. "_C. Madrena, Sto. T. Quieri_," UIMNH 46904. near Chivela, MCZ
25021. Cinco Cerros, UIMNH 37114. _Dami Liesa_, AMNH 66877, UIMNH
6158, 37115. _Escuranos_, AMNH 66873-74, 68004-06. _Finca Santa
Teresa, 2 km. NW Tehuantepec_, UMMZ 82648. _Huilotepec_, AMNH 66878,
UIMNH 40820. _between Huilotepec and Tehuantepec_, AMNH 65106, UMMZ
82644-45. _Las Tejas_, UIMNH 6151-54. _Mixtequilla_, UIMNH 6157,
36211. _between Mixtequilla Mountains and Tehuantepec_, UMMZ 82652.
_between Niltepec and "Carixxal,"_ AMNH 68876. 10.8 mi. SE Oaxaca,
UMMZ 114483. _Quiengola_, UIMNH 17617. _between Quiengola Mountains
and Tehuantepec_, UMMZ 82647. _Rancho Poso Río, 6 km. S Tehuantepec_,
UIMNH 6144-49, 37117-19, UMMZ 82649-51. _Rincón Bamba_, CNHM
105129-30, UIMNH 17615. _Salazar_, AMNH 66875. _vicinity of Salina
Cruz_, UMMZ 82653. _San Gerónimo_, AMNH 4306, CNHM 1457. _San Lucas
Ixtepec_, UIMNH 36206. San Juan Lajarcia, UIMNH 36212. San Mateo del
Mar, AMNH 65914. _San Pablo_, UIMNH 36207. _Santa María (Cerro de
Liesa)_, AMNH 68011. Tapanatepec, MCZ 27806-11. Tehuantepec, AMNH
19644, 65107-09, 65907-13 plus 7, 66871-72, 66879, 68007-08, CNHM
40435-36, 105126-28, MCZ 46403, UIMNH 6150, 17614, 17618, 29692,
36208, 37120-21, UMMZ 82642-43, 82646, USNM 109709-14, _1-2 leagues
SSE Tehuantepec_, UMMZ 82639-41. Tenango, UIMNH 36209-10. between
Tlacolulita and Tequisistlán, CNHM 105125. _Yerba Santa_, UIMNH
6155-56. Puebla: Atencingo, KU 39626.
Skull
In studying the osteology of the genus _Conophis_, I have examined
two complete skeletons (one _C. vittatus_ and one _C. lineatus_); two
additional skulls of _C. vittatus_ and _C. lineatus_; and 24 sets of
dentigerous bones, representing all of the species. Terminology
of the skeletal elements is that of Duellman (1958), Parker (1878),
Radovanovic (1937) and Szunyoghy (1932). The drawing of
the right side of the skull of a specimen of _Tomodon lineatus_ that
appears in Jan and Sordelli (1881:liv. 50, pl. 2, fig. 34) is of little
value due to its small size and lack of detail.
The skull of _Conophis_ is typical of a relatively unspecialized
colubrid snake. Skulls of _Conophis lineatus concolor_ and _C. vittatus_
closely resemble each other. The following description is based
primarily on the skull of _C. lineatus concolor_ (UMMZ S-778).
The elements are discussed in the following order: nasal region,
cranium and associated elements, maxillo-palatal-pterygoid arch,
mandible, dentition, and vertebrae.
[Illustration: FIG. 6. The skull, lacking dentigerous bones, of
_Conophis lineatus concolor_ (UMMZ S-788) showing (A) dorsal,
(B) lateral, and (C) ventral views. × 3.]
_Nasal region._--The premaxillary is relatively heavy and has a
concavity posteroventrally. The lateral processes slope downward, but
remain fairly thick, and do not project far laterally. This shape
(fig. 6) tends to strengthen the nasal region; this anterior
strengthening may be a reflection of the fossorial habits of these
snakes. There are no posterior processes of the premaxillary; thus the
line of fusion with the nasals and septo-maxillaries is broad. The
nasal plate is more than twice as long as wide. The nasals are
relatively flat above, although each curves slightly downward medially
and fuses into the medial nasal septum; laterally each nasal is
narrower and deflected downward, forming a small dorsal shield over
the nasal cavity. The septo-maxillaries are closely associated with
the vomers and form the cavity in which the organ of Jacobson is
situated. The broad medial part of the septo-maxillary forms the roof
and anterior border of the cavity, whereas the anterior part of the
vomer contains the main part of the capsule and forms the posterior
and most of the lateral borders of the cavity. The vomer has a thin
anterior ridge that gradually disappears before it reaches the border
of the premaxillary. The vomer is approximately U-shaped, when viewed
from below. It has no posterior process and does not articulate with
the parasphenoid; there is a sizeable gap between the two bones. The
septo-maxillary has a lateral process that terminally is directed
slightly anteriorly.
_Cranium and associated elements._--The frontal is almost three times
as long as it is wide; it is flat above with an emarginate
dorsolateral margin that forms the upper limit of the optic capsule.
Ventrally the frontal is concave and forms the median limits of the
optic cavity. Farther ventrally the frontal joins with the
parasphenoid, which at this place forms the ventral extent of the
skull, and together with the basisphenoid forms the ventral part of
the posterior three-fourths of the skull. In ventral aspect, the
parasphenoid is a long, thin bone, slightly expanded anteriorly. It
forms the anterior floor of the optic foramen; whereas the frontal
forms the anterior roof of the same opening. The frontal and its
septo-maxillary process surround the olfactory fenestra. The
prefrontal articulates with the anterolateral process of the frontal.
The posterior surface of the prefrontal forms the anterior wall of the
orbit of the eye. The articulating surface upon which the median
process of the maxillary bone rests is situated ventrally. The
anterior dorsal surface of the prefrontal, together with the
anterolateral edge of the frontal, extends slightly over the nasal
cavity, affording some degree of protection for the contained organs
and forming the posterior border of the cavity. A small nasal process
also extends anteriorly from the ventrolateral surface of the
prefrontal. The orbital-nasalis foramen is located in the anterior
surface of the prefrontal. The parietals are fused into one large bone
that forms the roof and sides of the middle part of the cranial
cavity. From its suture with the frontal, the dorsal surface of the
parietal is relatively flat in the area bounded laterally by the
parietal crests, which extend posteromedially from the anterolateral
corners of the bone and converge medially at a point near its
posterior margin. A slight posterior extension of the parietal crests
forms the supratemporal crest, which is present on the posterior part
of the parietal and on the anterior part of the supraoccipital. The
postfrontals are attached to the anterolateral processes of the
parietal. Together the anterior surfaces of these two bones form the
posterior rim of the orbit of the eye. The postfrontal extends
laterally and ventrally and has a terminal extension that projects
anterolaterally. In an articulated skull the trans-palatine articulates
with the ventrolateral articulating surface of the postfrontal.
Anteromedially, the parietal forms the roof and posterior margin of
the optic foramen. The basisphenoid, which is fused with the
parasphenoid, also forms a small part of the posteroventral margin of
the optic foramen. The basisphenoid forms the floor of the middle part
of the cranial cavity and the ventromedial down-pouching that
contains the pituitary body. Posterolateral to the parietal and dorsal
to the posterior part of the basisphenoid is the prootic. Laterally
this bone is deeply emarginate; posteriorly it forms a large part of
the otic notch, through which the columella passes. The columella is a
long, thin bony rod that terminates posteriorly in cartilage. It is
the cartilagenous part of the columella that connects with the
external sound detecting mechanism. There are several foramina on the
lateral surface of the prootic. On the anterolateral surface of the
prootic, branches of the trigeminal nerve pass through three foramina
whereas the facial nerve passes through the single posterior foramen
near the otic notch. The squamosal is attached dorsoventrally to the
posterior part of the parietal and to the lateral part of the prootic.
At this place of attachment there is on the prootic a relatively heavy
crest that forms a rather broad articulating base. The squamosal is
long, flat, and curves slightly in a dorsal direction throughout its
length; it becomes thinner and narrower posteriorly. The posterior
third of the squamosal forms a broad base by means of which the
squamosal articulates with the quadrate. The columella and the
squamosal extend posteriorly beyond the limits of the braincase.
Posteriorly the skull consists of four bones: an unpaired median
dorsal supraoccipital, an unpaired median ventral basioccipital and
two lateral exoccipitals. The basioccipital does not have noticeable
pterygoid processes, but is rather smooth ventrally and only slightly
emarginate on its posterolateral margins. Posteriorly, this bone forms
the ventral part of the occipital condyle. The rest of the condyle, on
each side, is formed by the exoccipitals. The exoccipitals also form
part of the base to which the squamosal is attached. The exoccipitals
extend around the sides of the foramen magnum and meet dorsally. Each
exoccipital also forms the posterior part of the otic notch, which
traverses the exoccipital. The exoccipitals bear moderate occipital
crests that extend posterolaterally across the supraoccipital as
branches from the supratemporal crest. The supraoccipital also has a
medial crest that extends a short distance posteriorly from the
supratemporal crest onto the exoccipitals at their dorsal line of
fusion.
[Illustration: FIG. 7. The maxillo-palatal-pterygoid arch of
_Conophis lineatus concolor_ (UMMZ S-788) showing (A) dorsal,
(B) lateral, and (C) ventral views. × 3. Teeth shown by means
of broken lines were represented only by their sockets.]
_Maxillo-palatal-pterygoid arch._--In an articulated skull, the
anterior edge of the maxillary is immediately posterior to the lateral
tip of the premaxillary (fig. 7). The maxillary is curved moderately
laterally and is not robust at its tip, but it becomes heavier about
one-third of its length posteriorly. A dorsomedian process begins at
about one-third of its distance from the anterior end; the prefrontal
articulates with this process. The process is broad and almost flat,
except that at its medial end, an elongate, rounded knob extends
ventrally. The dorsomedian process of the maxillary extends toward,
but does not meet, a lateral process from the palatine. The maxillary
teeth are set in sockets on the ventral surface of the bone. Just
posterior to the level of the last prediastemal tooth is the median
trans-palatine process that articulates with the anteromedian part of
the trans-palatine. Immediately posterior to this process, the
maxillary narrows slightly; then it broadens to form an obliquely
oriented knob. The posteroventral surface of the posterior knob of the
maxillary bears one or two enlarged maxillary teeth. (These teeth are
discussed further in the section on Dentition.) The anterolateral part
of the trans-palatine articulates with the dorsal surface of the
posterior knob of the maxillary. Toward the middle of its length, the
trans-palatine narrows considerably; then it broadens again and
articulates with the pterygoid. The palatine is slightly rounded at
its anterior end, which extends anteriorly to the posterior margin of
the vacuity containing Jacobson's organ. The palatine extends
posteriorly to the trans-palatine process of the maxilla, where the
palatine articulates with the pterygoid. A posterior pterygoid process
from the palatine projects posteromedially from the end of the
palatine and overlaps the anterior end of the pterygoid. Just less
than one-half the distance from the anterior end of the palatine,
there is a lateral process that curves ventrolaterally forming a blunt
tip posteriorly. Slightly more posteriorly and on the medial side of
the palatine, is a medial sphenoid process, which is thin, rather
broad, and curves ventromedially; ultimately it comes to lie near the
anterior part of the parasphenoid. The palatine teeth are set in
shallow sockets on the ventral edge of the bone. Of the bones of the
maxillo-palatal-pterygoid arch, those on the pterygoid extend farthest
posteriorly. The pterygoid is broad medially and posteriorly, although
pointed at its posterior tip. The trans-palatine articulates in a
broad line at about one-third of the distance along the lateral margin
of the pterygoid. Immediately posterior to this articulation, there is
a median ridge on the pterygoid; lateral to the pterygoid ridge is an
abrupt hollow, the pterygoid groove. Posteromedially, this groove
becomes gradually more shallow and disappears. The dorsal surface of
the pterygoid is rounded anteriorly and somewhat flattened
posteriorly, whereas the ventral surface is gently rounded along its
length, except that there is a high median crest. The pterygoid teeth
are situated in shallow sockets along this crest. The teeth diminish
in size posteriorly.
[Illustration: FIG. 8. The left mandible and associated
quadrate of _Conophis lineatus concolor_ (UMMZ S-788) showing
(A) lateral and (B) medial views. × 3. Teeth shown by means of
broken lines were represented only by their sockets.]
_Mandible._--The dentary (fig. 8) is compressed laterally and rounded
below. The teeth, which are longest about one-third of the way from
the anterior end of the dentary, are set in sockets on the medial side
of the bone. The posterior half of the dentary overlies the fused
surangular-prearticular part of the articular. Ventrally, the
posterior part of the dentary underlies the splenial, which is set in
a median trench within the dentary. Near the common suture of the
dentary and the splenial is the large inferior alveolar foramen;
completely within the splenial and ventral to the inferior alveolar
foramen is the anterior mylohyoid foramen. Posterior to the splenial
and also forming a part of the ventral surface of the mandible is the
wedge-shaped angular, which lies directly beneath the fused
surangular-prearticular. As has been implied, the articular, the
surangular, and the prearticular are fused. The prearticular part of
this bone forms a part of Meckel's canal. In the surangular part,
immediately posterior to the end of the dentary, is the large
surangular foramen. Lying in a longitudinal axis along the medial
surface of the articular is a high crest, dorsal to which is a deep
hollow. The lateral wall of the articular above this hollow is thin
and rounded dorsally; the ventral surface is uniformly round and
slightly curved dorsally, except that it ends with a short tympanic
crest, which projects beyond the articulation with the quadrate. Where
the quadrate articulates with the dorsolateral surface of the
posterior portion of the squamosal, the former is broad and has a high
mid-lateral crest, which extends about one-third of the distance down
the quadrate before disappearing. The columellar process (the place of
fusion of the columella) is about two-thirds of the way down the
medial surface of the quadrate. Ventrally the quadrate has a narrow
neck dorsal to its articulation with the articular. The articulation
is formed by two lateral flanges of the quadrate that fit over a
medial ridge formed by the articular.
Dentition
Teeth on the maxillary and pterygoid decrease in size posteriorly,
whereas those of the dentary do likewise except for the first one or
two that are usually slightly smaller than those immediately
posterior. The palatine teeth are subequal in size. The enlarged,
grooved teeth on the maxillary are in shallow sockets on the
posteroventral surface of the posterior knob of the maxillary. These
teeth point posteriorly. The grooves are deep and are situated
anterolaterally. One or two enlarged grooved teeth are present on a
given maxillary. There seems to be a correlation between the type of
preservation, the age of the snake, and the number of grooved teeth.
Old (large) individuals always have only one grooved tooth that is
rooted and functional, whereas some of the younger animals have two in
place. Usually replacement teeth are present in alcoholic specimens,
but these unrooted teeth are lost in the preparation of dried
skeletons. Thus, it seems that in _Conophis_ only one pair of grooved
teeth is functional at any one time, although usually replacement
teeth are present behind and beside the functional one. Some specimens
have one tooth in the medial socket on one side and one in the lateral
socket on the other. Replacement teeth on the maxillary and dentary
are present in the buccal tissue on the medial side of the bones,
whereas on the palatines and pterygoids, the replacement teeth are
present laterally. Apparently there are no significant differences in
dentition among the members of the genus _Conophis_.
Vertebrae
The fiftieth vertebra of _Conophis vittatus_ (UMMZ 82642) can be
described as follows: The neural spine is elongate, thin and low; the
posterior edge is sharply emarginate, and the anterior edge is only
slightly emarginate. The zygosphene is thin dorsoventrally; in a
ventral or dorsal view the zygosphene has a slightly concave anterior
edge, the flat surface of which is oriented ventrolaterally. The
centrum is elongate and triangular from below; it is widest at the
paradiapophyses and narrowest at the short condylar neck. The condylus
is directed posteriorly. The centrum, when viewed laterally, is
slightly concave and has prominent subcentral ridges that extend from
the median side of the paradiapophysial articular surfaces posteriorly
to the neck of the condylus. The paradiapophysial articular surfaces
are well developed and have two facets. The diapophysial surface is
larger and more spherical than the parapophysial one. The
parapophysial process projects beyond the parapophysial articular
surface and is nearly even with the lip of the cotyle, which is
slightly oval. The neural arch is slightly depressed; its width is
somewhat less than the width of the cotyle. The articular surfaces of
the postzygapophyses are oval and are directed posterolaterally. There
is a strongly developed concave interzygapophysial ridge. A
well-developed accessory spine extends laterally beyond the oval
articular facets of the pre-zygapophysis and forms a slightly
flattened, blunt spine. Excellent drawings of the middle thoracic
vertebra of _Conophis lineatus dunni_ from Honduras were published by
Auffenberg (1958:6).
Hemipenes
The hemipenes of _Conophis_ are moderately caliculate, having spines
covering the surface from the base to near the apex (fig. 9). These
spines are largest near the base and are reduced to small papillate
projections near the apex. The apex terminates in a small disc having
three to five laminae in _C. vittatus_ and one lamina in _C. lineatus
concolor_. The sulcus is bifurcate; the fork is near the base and
almost gives the appearance of two sulci on some specimens. Distally
the apices are widely separated, and the intervening space gives the
hemipenis a slightly bilobed appearance in some species (especially
_C. vittatus_) or a deeply bilobed appearance in others (especially
_C. lineatus concolor_).
[Illustration: FIG. 9. The everted left hemipenis of _Conophis
vittatus_ (UMMZ 82650). × 5.]
The everted hemipenis reaches posteriorly to the eighth subcaudal
scale. The sulcus bifurcates at the third subcaudal scale. The
situation is similar _in situ_ (Cope, 1895:pl. 28, fig. 2).
There are no apparent hemipenial differences among the species of the
genus _Conophis_. As can be seen in the above description, the
hemipenis of _C. vittatus_ is less bilobed and has a more pronounced
disc at the apex than the others. The hemipenis of _C. lineatus
concolor_ is most bilobed, but has the smallest apical disc. The other
species and subspecies vary widely within these extremes.
Food and Feeding
_Conophis_ eats mostly small lizards, especially _Cnemidophorus_. In
México _Conophis_ occurs in semi-arid habitat where _Cnemidophorus_ is
common. A specimen each of _Conophis vittatus_ and _C. lineatus
lineatus_ were obtained while I was collecting _Cnemidophorus_. The
only record of _Conophis_ having fed on a warm-blooded vertebrate was
obtained in the course of this study, when I recovered from the
stomach of a _Conophis lineatus concolor_ (CNHM 36299) from Chichén
Itzá, Yucatán, a heteromyid rodent (_Heteromys gaumeri_).
Ralph Axtell (personal communication) observed _Conophis_ actively
searching for food at dusk. His observations were made near
Tehuantepec, Oaxaca, and the snakes were seen to chase lizards of the
genus _Cnemidophorus_. Near Alvarado, Veracruz, in the late afternoon,
I watched a _Conophis lineatus lineatus_ follow a lizard into a hole.
Mittleman (1944:122) presents the only discussion of the mode of
feeding of a captive specimen of _Conophis lineatus_ ssp. When
presented with a _Thamnophis_ slightly smaller than itself, the
_Conophis_ struck, and within eight minutes immobilized the
_Thamnophis_. Within one-half hour the _Thamnophis_ was swallowed.
Three days later the _Conophis_ ate another _Thamnophis_, though still
distended from its first meal; nine days later it ate a _Storeria_. In
the course of several months, the _Conophis_ ate various toads and
hylids and two more _Storeria_. Apparently members of the genus
_Conophis_ do not constrict their prey, but rely upon a combination of
loss of blood and action of the venom to completely immobilize their
prey.
Ditmars (1931:pls. 26-27) showed three photographs of "_Conophis
lineatus_" (actually _Conophis pulcher_) ingesting another snake,
identified by him as a young _Ophis (= Xenodon) colubrinus_.
Effect of Poison
The rear fangs of these snakes are large for the size of the snake.
Various collectors have been bitten, and several reports of the effect
of the poison have been published. The snakes are aggressive and bite
constantly while being handled. A field companion, Dale L. Hoyt, was
bitten on the forefinger by a specimen of _C. l. lineatus_ and
immediately felt a burning sensation. The finger swelled, much as it
would if stung by a wasp, but it returned to normal size in about
twenty-four hours. Ditmars (1931:legend pl. 27) reported immediate
burning pain and a localized swelling, an inch in diameter and half an
inch high, which lasted for several hours. Mertens (1952b:83) reported
merely that the hand of the gardener at the Instituto Tropical in San
Salvador bled strongly for a full hour. Edward H. Taylor was bitten by
a specimen of _Conophis vittatus_ (Taylor and Smith, 1939:252); pain
and swelling lasted for some time. Taylor (personal communication) is
still troubled by damage incurred by that bite, which apparently
resulted in mechanical damage to the second joint of the middle
finger, for the joint swells when the finger is used or exercised.
William E. Duellman (personal communication) was bitten on the hand in
July, 1956. There was immediate pain and localized swelling, both of
which disappeared several hours later.
TAXONOMIC RELATIONSHIPS AND EVOLUTION
The genus _Conophis_ is known only from the Recent. Except that
_Conophis_ belongs to the subfamily xenodontinae and probably is of
New World origin, little is known about the relationships of the
genus. Auffenberg (1958) described a new genus and species of fossil
colubrid snake from the Miocene of Montana as _Dryinoides oxyrhachis_
and compared it with several recent genera. This specimen, of which
there is a relatively complete skull and a series of vertebrae, seems
most closely to resemble a specimen of _Conophis lineatus dunni_ (UF
7657) from Honduras, with which it was compared in basic osteology.
The two genera could be related, for the progenitors of _Conophis_
possibly inhabited much of North America in the Miocene.
Another possibility is that the main stock of the xenodontines reached
South America in earliest Tertiary times, and that the formation of
the Panamanian and Colombian seaways that separated South America and
Central America from the Late Paleocene to the middle of the Pliocene
left the _Conophis_ stock isolated in Middle America where members of
the genus dispersed through semi-arid habitats.
Turning our attention now to the species within the genus, instead of
the genus as a whole, _Conophis vittatus_ is readily set apart from
other members of the genus on the basis of the universal presence of
seven supralabials. In basic coloration it also differs, having no
stripe on the 1st scale-row, or spots on the venter, and a maximum of
four broad stripes on the body. The other species appear to be more
closely related; these make up the _C. lineatus_-group. _Conophis
nevermanni_ differs so much from the other species that it might be
placed in a separate group. Nevertheless, the basic striped pattern,
which is masked by the increased melanism of many specimens, indicates
that _nevermanni_ is more closely related to the _lineatus_-group than
to _vittatus_. The _lineatus_-group, thus, consists of _pulcher_,
_nevermanni_ and the three subspecies of _lineatus_. In this group the
color pattern is characterized by the high frequency of ventral
spotting, darkening of part of the supralabials, dark pigmentation on
the 1st scale-row, and more than four dark stripes on the body of
adults. _Conophis lineatus concolor_, on which the dark pigmentation
on the body apparently is secondarily lost, is an exception.
If differences in color pattern be used as an indication of the
relationships between the species and subspecies of the genus
_Conophis_, I would consider _C. vittatus_ the most divergent unit.
The subspecies of _lineatus_ closely resemble one another and, as a
unit, resemble _pulcher_ from which they differ primarily in the
position of the dorsalmost stripes. _Conophis nevermanni_ is more
divergent than is _pulcher_ from the species _lineatus_, but probably
is not so far removed from _lineatus_ as is _vittatus_.
In the light of what has been pointed out immediately above with
respect to resemblances of, and differences between, the species, an
hypothesis to account for their formation and for their presence in
the areas where they are today is the following: Concurrent with
climatic fluctuations in the Late Pliocene and Pleistocene, the
northernmost population differentiated into the species _vittatus_,
and has subsequently spread north and west from the region of
Tehuantepec, México. During the same period _nevermanni_ became
isolated in northern Costa Rica.
The species _pulcher_ probably differentiated from the remaining
_lineatus_ stock during the Early Pleistocene orogenic upheaval in
Guatemala. The _pulcher_ stock was isolated on the Pacific Coastal
slopes of Guatemala, while _lineatus_ moved through the subhumid
corridor of northern Middle America into México and southward toward
Costa Rica (Stuart, 1954a). In the Late Pleistocene and Recent,
_pulcher_ moved back across the central Guatemalan highlands occupying
its present range in northern Middle America. Primarily because of the
formation of unsuitable habitat (wet forest) that presently separates
the geographic ranges of populations of _lineatus_, this species
differentiated into three subspecies.
SUMMARY
The genus _Conophis_ Peters, 1860, contains four species. Three are
monotypic and the fourth has three subspecies, making a total of six
taxa.
The genus is characterized by maxillary teeth of equal size followed
by a diastema and two enlarged grooved fangs. The scales are smooth,
in 19 rows at mid-body, and 17 nearer the tail. The anal is divided,
apical pits are lacking, the head shields are normal for a colubrid,
and the hemipenis is bilobed having many large basal spines.
The six taxa are separated primarily on the basis of color pattern,
but characters of scutellation, including numbers of dorsals,
ventrals, caudals, and places of reduction of the number of dorsal
scale-rows, were analyzed.
Snakes of this genus are distributed throughout semi-arid environments
from southern México southward into Costa Rica. They feed upon
lizards, primarily of the genus _Cnemidophorus_; in addition they are
known to eat small rodents and other snakes.
_Conophis_ is a member of the subfamily Xenodontinae and, as presently
understood, has no known living close relatives. A single specimen of
_Dryinoides_ from the Miocene of Montana has been compared with this
genus. The genus _Conophis_ is thought to have evolved in Middle
America. The present distribution and differentiation probably are
primarily the result of climatic fluctuations in Middle America, which
produced the areas of subhumid environment where _Conophis_ presently
lives.
LITERATURE CITED
AUFFENBERG, W.
1958. A new genus of colubrid snake from the Upper Miocene of
North America. Amer. Mus. Novitates, 1874:1-16. February 27.
COPE, E. D.
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México and Central America. Proc. Acad. Nat. Sci.
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1867. Fifth contribution to the herpetology of tropical
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February 20.
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October 24.
1876. On the batrachia and reptilia of Costa Rica. Journ.
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1895. The classification of the ophidia. Trans. Amer. Philos.
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1900. The crocodilians, lizards, and snakes of North America.
Ann. Rept. U. S. Natl. Mus. for 1898, pp. 153-1270, 36 pls.
DITMARS, R. L.
1931. Snakes of the World. New York, The MacMillan Company,
1931. xi + 207 pp., 84 pls.
DOWLING, H. G.
1951. A proposed standard system of counting ventrals in
snakes. British Journ. Herpetology, 1(5):97-99, fig. 1.
DUELLMAN, W. E.
1958. A preliminary analysis of the herpetofauna of Colima,
Mexico. Occas. Papers Mus. Zool. Univ. Michigan, 589:1-22,
March 21.
DUMÉRIL, A. M. C., BIBRON, G., AND DUMÉRIL, A. H. A.
1854. Érpétologie genérale, ou histoire naturelle des
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DUMÉRIL, A. H. A., BOCOURT, M., AND MOCQUARD, F.
1870-1909. Mission Scientifique au Mexique et dans l'Amerique
Centrale ... Etudes sur les Reptiles. Paris, vol. 2:xiv +
1012 pp., 77 pls.
GARMAN, S.
1884a. The North American reptiles and batrachians. Bull.
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1884b. The reptiles and batrachians of North America. Mem.
Mus. Comp. Zool., 8(3):xxxi + 185 pp., 9 pls. July.
GÜNTHER, A. C. L. G.
1858. Catalogue of colubrine snakes in the collection of the
British Museum. London. xiv + 281 pp.
HUXLEY, J.
1942. Evolution. The Modern Synthesis. London. 645 pp.
JAN, G. AND SORDELLI, F.
1866. Iconographie Generale des Ophidiens. Milano. livr. 19,
pls. 1-6. December.
1881. Iconographie Generale des Ophidiens. Milano. livr. 50,
pls. 1-7. November.
MAYR, E.
1942. Systematics and the Origin of Species. New York, x +
334 pp., 29 figs.
MAYR, E., LINSLEY, E. G., AND USINGER, R. L.
1953. Methods and Principles of Systematic Zoology. New York.
ix + 328 pp., 45 figs.
MERTENS, R.
1952a. Neues uber die Reptilienfauna von El Salvador. Zool.
Anz., 148:87-93. February.
1952b. Die Amphibien und Reptilien von El Salvador auf grund
der reisen von R. Mertens und A. Zilch. Abhand. Senken.
Naturw. Gesell., 487:83, 1 Kart., 16 taf. December 1.
MITTLEMAN, M. B.
1944. Feeding habits of a Central American opisthoglyph snake.
Copeia, no. 2:122. June 30.
NEILL, W. T. AND ALLEN, R.
1961. Further studies on the herpetology of British Honduras.
Herpetologica, 17(1):37-52. April 15.
PARKER, W. K.
1878. On the structure and development of the skull in the
common snake (_Tropidonotus natrix_). Phil. Trans. Roy.
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PETERS, W.
1860. Drei neue amerikanisches Schlangen. Monatsb. Akad. Wiss.
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RADOVANOVIC, M.
1937. Osteologie des Schlangenkopfs. Jenaische Zeitschr.
Naturw., 71(2):179-312.
SAVAGE, J. M.
1949. Notes on the Central American snake, _Conophis lineatus
dunni_ Smith, with a record from Honduras. Trans. Kansas
Acad. Sci., 50:483-486. December 31.
SCHMIDT, K. P.
1928. Reptiles collected in Salvador for the California
Institute of Technology. Zool. Ser. Field Mus. Nat. Hist.,
12(16):193-201. November 21.
SCHMIDT, K. P. AND INGER, R. F.
1957. Living Reptiles of the World. Garden City, New York,
Hanover House. 287 pp.
SMITH, H. M.
1941. Notes on snakes of the genus _Conophis_. Journ.
Washington Acad. Sci., 31(3):117-124. March 15.
SMITH, H. M. AND TAYLOR, E. H.
1950. Type localities of Méxican reptiles and amphibians.
Univ. Kansas Sci. Bull., 33:313-380. March 20.
STUART, L. C.
1948. The amphibians and reptiles of Alta Verapaz, Guatemala.
Misc. Publ. Mus. Zool. Univ. Michigan, 69:1-109. June 12.
1954a. A description of a subhumid corridor across northern
Central America, with comments on its herpetofaunal
indicators. Contr. Lab. Vert. Biol. Univ. Michigan,
65:1-26 pp., 6 pls. March.
1954b. Herpetofauna of the southeast highlands of Guatemala.
Contr. Lab. Vert. Biol. Univ. Michigan, 68:1-65 pp.,
3 pls. November.
SZUNYOGHY, J.
1932. Beitrage zur vergleichenden Formenlehre des
Colubridenschadels, nebst einer Kraniologischen Synopsis
der fossilen Schlangen Ungarns. Acta Zool., 13:1-56.
TAYLOR, E. H.
1955. Additions to the known herpetological fauna of Costa
Rica with comments on other species. No. II. Univ. Kansas
Sci. Bull., 37:299-575. October 15.
TAYLOR, E. H. AND SMITH, H. M.
1939. Miscellaneous notes on Mexican snakes. Univ. Kansas Sci.
Bull., 25:239-258. July 10.
WETTSTEIN, O.
1934. Ergibnisse der osterreichischen biologischen Costa
Rica--Expedition 1930. Die Amphibia und Reptilien. Stiz.
Akad. Wiss. Wien, mathem-naturw. kl., Abt. 1, bd. 143:1-39.
_Transmitted November 30, 1962._
29-5936
[]
UNIVERSITY OF KANSAS PUBLICATIONS
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this series, are as follows:
Vol. 1. Nos. 1-26 and index. Pp. 1-638, 1946-1950.
*Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest.
Pp. 1-444, 140 figures in text. April 9, 1948.
Vol. 3. *1. The avifauna of Micronesia, its origin, evolution,
and distribution. By Rollin H. Baker. Pp. 1-359,
16 figures in text. June 12, 1951.
*2. A quantitative study of the nocturnal migration of
birds. By George H. Lowery, Jr. Pp. 361-472,
47 figures in text. June 29, 1951.
3. Phylogeny of the waxwings and allied birds. By
M. Dale Arvey. Pp. 473-530, 530, 49 figures in text,
13 tables. October 10. 1951.
*4. Birds from the state of Veracruz, Mexico. By George H.
Lowery, Jr., and Walter W. Dalquest. Pp. 531-649,
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Index. Pp. 651-681.
*Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466,
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*Vol. 6. (Complete) Mammals of Utah, taxonomy and distribution. By
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30 tables. August 10, 1952.
Vol. 7. Nos. 1-15 and index. Pp. 1-651, 1952-1955.
Vol. 8. Nos. 1-10 and index. Pp. 1-675, 1954-1956.
Vol. 9. *1. Speciation of the wandering shrew. By James S. Findley.
Pp. 1-68, 18 figures in text. December 10, 1955.
2. Additional records and extension of ranges of mammals
from Utah. By Stephen D. Durrant, M. Raymond Lee, and
Richard M. Hansen. Pp. 69-80. December 10, 1955.
3. A new long-eared myotis (Myotis evotis) from northeastern
Mexico. By Rollin H. Baker and Howard J. Stains. Pp. 81-84.
December 10, 1955.
4. Subspeciation in the meadow mouse, Microtus pennsylvanicus,
in Wyoming. By Sydney Anderson. Pp. 85-104, 2 figures in
text. May 10, 1956.
5. The condylarth genus Ellipsodon. By Robert W. Wilson.
Pp. 105-116, 6 figures in text. May 19, 1956.
6. Additional remains of the multituberculate genus
Eucosmodon. By Robert W. Wilson. Pp. 117-123,
10 figures in text. May 19, 1956.
7. Mammals of Coahulia, Mexico. By Rollin H. Baker. Pp.
125-335, 75 figures in text. June 15, 1956.
8. Comments on the taxonomic status of Apodemus peninsulae,
with description of a new subspecies from North China.
By J. Knox Jones, Jr. Pp. 337-346, 1 figure in text,
1 table. August 15, 1956.
9. Extensions of known ranges of Mexican bats. By Sydney
Anderson. Pp. 347-351. August 15, 1956.
10. A new bat (Genus Leptonycteris) from Coahulia. By
Howard J. Stains. Pp. 353-356. January 21, 1957.
11. A new species of pocket gopher (Genus Pappogeomys) from
Jalisco, Mexico. By Robert J. Russell. Pp. 357-361.
January 21, 1957.
12. Geographic variation in the pocket gopher, Thomomys
bottae, in Colorado. By Phillip M. Youngman. Pp.
363-384, 7 figures in text. February 21, 1958.
13. New bog lemming (genus Synaptomys) from Nebraska.
By J. Knox Jones, Jr. Pp. 385-388. May 12, 1958.
14. Pleistocene bats from San Josecito Cave, Nuevo León,
México. By J. Knox Jones, Jr. Pp. 389-396.
December 19, 1958.
15. New subspecies of the rodent Baiomys from Central America.
By Robert L. Packard. Pp. 397-404. December 19, 1958.
16. Mammals of the Grand Mesa, Colorado. By Sydney Anderson.
Pp. 405-414, 1 figure in text, May 20, 1959.
17. Distribution, variation, and relationships of the montane
vole, Microtus montanus. By Sydney Anderson. Pp. 415-511,
12 figures in text, 2 tables. August 1, 1959.
18. Conspecificity of two pocket mice, Perognathus goldmani
and P. artus. By E. Raymond Hall and Marilyn Bailey
Ogilvie. Pp. 513-518, 1 map, January 14, 1960.
19. Records of harvest mice, Reithrodontomys, from Central
America, with description of a new subspecies from
Nicaragua. By Sydney Anderson and J. Knox Jones, Jr.
Pp. 519-529. January 14, 1960.
20. Small carnivores from San Josecito Cave (Pleistocene),
Nuevo León, México. By E. Raymond Hall. Pp. 531-538,
1 figure in text. January 14, 1960.
21. Pleistocene pocket gophers from San Josecito Cave, Nuevo
León, México. By Robert J. Russell. Pp. 539-548, 1 figure
in text. January 14, 1960.
22. Review of the insectivores of Korea. By J. Knox Jones,
Jr., and David H. Johnson. Pp. 549-578. February 23, 1960.
23. Speciation and evolution of the pygmy mice, genus
Baimoys. By Robert L. Packard. Pp. 579-670, 4 plates,
12 figures in text. June 16, 1960.
Index. Pp. 671-690
Vol. 10. 1. Studies of birds killed in nocturnal migration.
By Harrison B. Tordoff and Robert M. Mengel. Pp. 1-44,
6 figures in text, 2 tables. September 12, 1956.
2. Comparative breeding behavior of Ammospiza caudacuta
and A. maritima. By Glen E. Woolfenden. Pp. 45-75,
6 plates, 1 figure. December 20, 1956.
3. The forest habitat of the University of Kansas Natural
History Reservation. By Henry S. Fitch and Ronald R.
McGregor. Pp. 77-127, 2 plates, 7 figures in text,
4 tables. December 31, 1956.
4. Aspects of reproduction and development in the prairie
vole (Microtus ochrogaster). By Henry S. Fitch. Pp.
129-161, 8 figures in text, 4 tables. December 19, 1957.
5. Birds found on the Arctic slope of northern Alaska.
By James W. Bee. Pp. 163-211, plates 9-10, 1 figure
in text. March 12, 1958.
*6. The wood rats of Colorado: distribution and ecology.
By Robert B. Finley, Jr. Pp. 213-552, 34 plates, 8 figures
in text, 35 tables. November 7, 1958.
7. Home ranges and movements of the eastern cottontail in
Kansas. By Donald W. Janes. Pp. 553-572, 4 plates,
3 figures in text. May 4, 1959.
8. Natural history of the salamander, Aneides hardyi.
By Richard F. Johnston and Gerhard A. Schad. Pp. 573-585.
October 8, 1959.
9. A new subspecies of lizard, Cnemidophorus sacki, from
Michoacán, México. By William E. Duellman. Pp. 587-598,
2 figures in text. May 2, 1960.
10. A taxonomic study of the middle American snake, Pituophis
deppei. By William E. Duellman. Pp. 599-610. 1 plate,
1 figure in text. May 2, 1960.
Index. Pp. 611-626.
Vol. 11. Nos. 1-10 and index. Pp. 1-703, 1958-1960.
Vol. 12. 1. Functional morphology of three bats: Sumops, Myotis,
Macrotus. By Terry A. Vaughan. Pp. 1-153, 4 plates,
24 figures in text. July 8, 1959.
*2. The ancestry of modern Amphibia: a review of the evidence.
By Theodore H. Eaton, Jr. Pp. 155-180, 10 figures in text.
July 10, 1959.
3. The baculum in microtine rodents. By Sydney Anderson.
Pp. 181-216, 49 figures in text. February 19, 1960.
*4. A new order of fishlike Amphibia from the Pennsylvanian
of Kansas. By Theodore H. Eaton, Jr., and Peggy Lou
Stewart. Pp. 217-240, 12 figures in text. May 2, 1960.
5. Natural history of the bell vireo. By Jon C. Barlow.
Pp. 241-296, 6 figures in text. March 7, 1962.
6. Two new pelycosaurs from the lower Permian of Oklahoma.
By Richard C. Fox. Pp. 297-307, 6 figures in text.
May 21, 1962.
7. Vertebrates from the barrier island of Tamaulipas, México.
By Robert K. Selander, Richard F. Johnston, B. J. Wilks,
and Gerald G. Raun. Pp. 309-345, pls. 5-8. June 18, 1962.
8. Teeth of Edestid sharks. By Theodore H. Eaton, Jr.
Pp. 347-362, 10 figures
in text. October 1, 1962.
More numbers will appear in volume 12.
Vol. 13. 1. Five natural hybrid combinations in minnows (Cyprinidae).
By Frank B. Cross and W. L. Minckley. Pp. 1-18.
June 1, 1960.
2. A distributional study of the amphibians of the Isthmus
of Tehuantepec, México. By William E. Duellman. Pp. 19-72,
pls. 1-8, 3 figures in text. August 16, 1960.
3. A new subspecies of the slider turtle (Pseudemys
scripta) from Coahulia, México. By John M. Legler.
Pp. 73-84, pls. 9-12, 3 figures in text. August
16, 1960.
4. Autecology of the copperhead. By Henry S. Fitch.
Pp. 85-288, pls. 13-20, 26 figures in text.
November 30, 1960.
5. Occurrence of the garter snake, Thamnophis sirtalis, in
the Great Plains and Rocky Mountains. By Henry S. Fitch
and T. Paul Maslin. Pp. 289-308, 4 figures in text.
February 10, 1961.
6. Fishes of the Wakarusa river in Kansas. By James E. Deacon
and Artie L. Metcalf. Pp. 309-322, 1 figure in text.
February 10, 1961.
7. Geographic variation in the North American cyprinid fish,
Hybopsis gracilis. By Leonard J. Olund and Frank B. Cross.
Pp. 323-348, pls. 21-24, 2 figures in text.
February 10, 1961.
8. Descriptions of two species of frogs, genus Ptychohyla;
studies of American hylid frogs, V. By William E. Duellman.
Pp. 349-357, pl. 25, 2 figures in text. April 27, 1961.
9. Fish populations, following a drought, in the Neosho and
Marais des Cygnes rivers of Kansas. By James Everett Deacon.
Pp. 359-427, pls. 26-30, 3 figs. August 11, 1961.
10. Recent soft-shelled turtles of North America (family
Trionychidae). By Robert G. Webb. Pp. 429-611, pls. 31-54,
24 figures in text. February 16, 1962.
Index. Pp. 613-624.
Vol. 14. 1. Neotropical bats from western México. By Sydney Anderson.
Pp. 1-8. October 24, 1960.
2. Geographic variation in the harvest mouse.
Reithrodontomys megalotis, on the central Great Plains
and in adjacent regions. By J. Knox Jones, Jr., and
B. Mursaloglu. Pp. 9-27, 1 figure in text. July 24, 1961.
3. Mammals of Mesa Verde National Park, Colorado. By Sydney
Anderson. Pp. 29-67, pls. 1 and 2, 3 figures in text.
July 24, 1961.
4. A new subspecies of the black myotis (bat) from eastern
Mexico. By E. Raymond Hall and Ticul Alvarez. Pp. 69-72,
1 figure in text. December 29, 1961.
5. North American yellow bats, "Dasypterus," and a list of
the named kinds of the genus Lasiurus Gray. By E. Raymond
Hall and J. Knox Jones, Jr. Pp. 73-98, 4 figures in text.
December 29, 1961.
6. Natural history of the brush mouse (Peromyscus boylii) in
Kansas with description of a new subspecies. By Charles A.
Long. Pp. 99-111, 1 figure in text. December 29, 1961.
7. Taxonomic status of some mice of the Peromyscus boylii
group in eastern Mexico, with description of a new
subspecies. By Ticul Alvarez. Pp. 113-120, 1 figure in
text. December 29, 1961.
8. A new subspecies of ground squirrel (Spermophilus
spilosoma) from Tamaulipas, Mexico. By Ticul Alvarez.
Pp. 121-124. March 7, 1962.
9. Taxonomic status of the free-tailed bat, Tadarida
yucatanica Miller. By J. Knox Jones, Jr., and Ticul
Alvarez. Pp. 125-133, 1 figure in text. March 7, 1962.
10. A new doglike carnivore, genus Cynaretus, from the
Clarendonian Pliocene, of Texas. By E. Raymond Hall and
Walter W. Dalquest. Pp. 135-138, 2 figures in text.
April 30, 1962.
11. A new subspecies of wood rat (Neotoma) from northeastern
Mexico. By Ticul Alvarez. Pp. 139-143. April 30, 1962.
12. Noteworthy mammals from Sinaloa, Mexico. By J. Knox Jones,
Jr., Ticul Alvarez, and M. Raymond Lee. Pp. 145-159,
1 figure in text. May 18, 1962.
13. A new bat (Myotis) from Mexico. By E. Raymond Hall.
Pp. 161-164, 1 figure in text. May 21, 1962.
14. The mammals of Veracruz. By E. Raymond Hall and Walter W.
Dalquest. Pp. 165-362, 2 figures. May 20, 1963.
15. The recent mammals of Tamaulipas, México. By Ticul Alvarez.
Pp. 363-473, 5 figures in text. May 20, 1963.
More numbers will appear in volume 14.
Vol. 15. 1. The amphibians and reptiles of Michoacán, México. By
William E. Duellman. Pp. 1-148, pls. 1-6, 11 figures in
text. December 20, 1961.
2. Some reptiles and amphibians from Korea. By Robert G. Webb,
J. Knox Jones, Jr., and George W. Byers. Pp. 149-173.
January 31, 1962.
3. A new species of frog (Genus Tomodactylus) from western
México. By Robert G. Webb. Pp. 175-181, 1 figure in text.
March 7, 1962.
4. Type specimens of amphibians and reptiles in the Museum
of Natural History, the University of Kansas. By William
E. Duellman and Barbara Berg. Pp. 183-204. October 26, 1962.
5. Amphibians and Reptiles of the Rainforests of Southern El
Petén, Guatemala. By William E. Duellman. Pp. 205-249, pls.
7-10, 6 figures in text. October 4, 1963.
6. A revision of snakes of the genus Conophis (Family
Colubridae, from Middle America). By John Wellman.
Pp. 251-295, 9 figures in text. October 4, 1963.
More numbers will appear in volume 15.
Transcriber's Notes
For consistancy, a number of word which had alternate spellings were
altered to match the most prevalent version used. For example, where
the word Mexico was used in the body of the article, the more frequent
spelling (México) was substituted. However, in the reference sections,
the spelling was not altered as that may have been the spelling used
by the article's author. All occurrances of Érpétologie Genérale were
correcteded to Erpétologie Générale (Pp. 255, 262, 267, 277, and 278).
On page 279 under _Variation_ there appears to be a miscalculation:
668 mm. + 182 mm. = 850 mm. not 840 as in original text.
Typographical Corrections
Page Correction
===== ===========================================
264 immaculaate => immaculate
264 chacteristic => characteristic
266 elevatons => elevations
267 Dumeril => Duméril
277 Duméil => Duméril
279 Tehauntepec => Tehuantepec
280 Deleted repeated "Oaxaca,"
292 primarly => primarily
295 hertetofaunal => herpetofaunal
i V. 9 No. 12: Pp. 363-387 => Pp. 363-384
iii V. 13 No. 8: Decriptions => Descriptions
iii V. 14 No. 8: anad => and
iii V. 14 No. 14: anad => and
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