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diff --git a/.gitattributes b/.gitattributes new file mode 100644 index 0000000..6833f05 --- /dev/null +++ b/.gitattributes @@ -0,0 +1,3 @@ +* text=auto +*.txt text +*.md text diff --git a/37512-8.txt b/37512-8.txt new file mode 100644 index 0000000..086df85 --- /dev/null +++ b/37512-8.txt @@ -0,0 +1,3259 @@ +The Project Gutenberg EBook of A Revision of Snakes of the Genus Conophis +(Family Colubridae, from Middle America), by John Wellman + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: A Revision of Snakes of the Genus Conophis (Family Colubridae, from Middle America) + +Author: John Wellman + +Release Date: September 23, 2011 [EBook #37512] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK A REVISION OF SNAKES OF THE *** + + + + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +http://www.pgdp.net + + + + + + + +Transcriber's Note + +Typographical corrections are listed at the end of this version. +The list of publications has been compiled after the article's text. + + * * * * * + + + 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. + + 1861. Contributions to the ophiology of Lower California, + México and Central America. Proc. Acad. Nat. Sci. + Philadelphia, 13:292-306. December 28. + + 1867. Fifth contribution to the herpetology of tropical + America. Proc. Acad. Nat. Sci. Philadelphia, 18:317-323. + February 20. + + 1871. Ninth contribution to the herpetology of tropical + America. Proc. Acad. Nat. Sci. Philadelphia, 23(2):200-224. + October 24. + + 1876. On the batrachia and reptilia of Costa Rica. Journ. + Acad. Nat. Sci. Philadelphia, series 2, 8(4):93-154, 6 pls. + + 1895. The classification of the ophidia. Trans. Amer. Philos. + Soc., 18:186-219, 33 pls. April 15. + + 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 + reptiles. Paris, 7(pt. 2):xii + 785. February 25. + Atlas, 24 pp., 108 pls. + + +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. + Essex Inst., 16:1-46. January 9. + + 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. + Soc. London, pt. 2:385-417, pp., pls. 27-33. + + +PETERS, W. + + 1860. Drei neue amerikanisches Schlangen. Monatsb. Akad. Wiss. + Berlin, 1860:517-521, pl., fig. 3. October. + + +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 + +MUSEUM OF NATURAL HISTORY + + +Institutional libraries interested in publications exchange may obtain +this series by addressing the Exchange Librarian, University of Kansas +Library, Lawrence, Kansas. Copies for individuals, persons working in +a particular field of study, may be obtained by addressing instead the +Museum of Natural History, University of Kansas, Lawrence, Kansas. +There is no provision for sale of this series by the University +Library, which meets institutional requests, or by the Museum of +Natural History, which meets the requests of individuals. However, +when individuals request copies from the Museum, 25 cents should be +included, for each separate number that is 100 pages or more in +length, for the purpose of defraying the costs of wrapping and +mailing. + +* An asterisk designates those numbers of which the Museum's supply +(not the Library's supply) is exhausted. Numbers published to date, in +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, + 7 figures in text, 2 tables. October 10, 1951. + + Index. Pp. 651-681. + + *Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, + 41 plates, 31 figures in text. December 27, 1951. + + Vol. 5. Nos. 1-37 and index. Pp. 1-676, 1951-1953. + + *Vol. 6. (Complete) Mammals of Utah, taxonomy and distribution. By + Stephen D. Durrant. Pp. 1-549, 91 figures in text, + 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 + + + + + +End of the Project Gutenberg EBook of A Revision of Snakes of the Genus +Conophis (Family Colubridae, from Middle America), by John Wellman + +*** END OF THIS PROJECT GUTENBERG EBOOK A REVISION OF SNAKES OF THE *** + +***** This file should be named 37512-8.txt or 37512-8.zip ***** +This and all associated files of various formats will be found in: + http://www.gutenberg.org/3/7/5/1/37512/ + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +http://www.pgdp.net + + +Updated editions will replace the previous one--the old editions +will be renamed. + +Creating the works from public domain print editions means that no +one owns a United States copyright in these works, so the Foundation +(and you!) can copy and distribute it in the United States without +permission and without paying copyright royalties. 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You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: A Revision of Snakes of the Genus Conophis (Family Colubridae, from Middle America) + +Author: John Wellman + +Release Date: September 23, 2011 [EBook #37512] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK A REVISION OF SNAKES OF THE *** + + + + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +http://www.pgdp.net + + + + + + +</pre> + + +<div class="book"> +<div class="fig_center" style="width: 255px;"> +<a name="cover" id="cover"></a> +<img src="images/cover.jpg" width="255" height="411" alt="" title="" /> +</div> +<br /> +<br /> + +<div class="trans_notes"> +<div class="caption2">Transcriber's Note</div> +<p>The cover above is simulated. Typographical corrections listed at + the <a href="#typos">end of this version</a> are highlighted in + <ins title="Correction: sample">light green</ins>. The + <a href="#PUBLICATIONS">list of publications</a> has been compiled + after the article's text.</p> +<br /> +</div> +<br /> + +<p><span class="pagenum"><a name="Page_251" id="Page_251">[Pg_251]</a></span></p> +<div class="center"> +<br /> +<br /> +<img src="images/bar_double.png" width="100%" height="15" alt="double bar" /> +<div class="caption2 smcap">University of Kansas Publications</div> +<br /> +<div class="caption2 smcap">Museum of Natural History</div> +<br /> +<img src="images/bar_single.png" width="125" height="15" title="bar" alt="bar" /> +<br /> +<div class="caption2">Volume 15, No. 6, pp. 251-295, 9 figs.</div><br /> +<img src="images/bar_single.png" width="250" height="15" title="bar" alt="bar" /> + <span class="caption2">October 4, 1963</span> +<img src="images/bar_single.png" width="250" height="15" title="bar" alt="bar" /> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<div class="caption1"> +A Revision of Snakes of the Genus Conophis<br /> +(Family Colubridae, from Middle America)<br /> +</div> +<br /> +<br /> +<br /> +<br /> +<div class="caption3">BY</div> +<br /> +<br /> +<div class="caption2">JOHN WELLMAN</div> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<div class="caption2"> +<span class="smcap">University of Kansas</span><br /> +<span class="smcap">Lawrence</span><br /> +1963 +</div> +<br /> +<br /> +<br /> +<br /> +</div> + +<p><span class="pagenum"><a name="Page_252" id="Page_252">[Pg_252]</a></span></p> + +<div class="center"> +<div class="caption3"> +<span class="smcap">University of Kansas Publications, Museum of Natural History</span><br /> +<br /> +Editors: E. Raymond Hall, Chairman, Henry S. Fitch,<br /> +Theodore H. Eaton, Jr.<br /> +<br /> +<br /> +<br /> +<br /> +Volume 15, No. 6, pp. 251-295, 9 figs.<br /> +<br /> +Published October 18, 1963<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<br /> +<span class="smcap">University of Kansas</span><br /> +Lawrence, Kansas<br /> +<br /> +</div> +<br /> +<br /> +<br /> +<br /> +<div class="caption4"> +PRINTED BY<br /> +JEAN M. NEIBARGER, STATE PRINTER<br /> +TOPEKA, KANSAS<br /> +1963<br /> +<img src="images/union_label.png" width="71" height="26" alt="Look for the Union Label" title="Look for the Union Label" /><br /> +29-5936<br /> +</div> +</div> +<br /> +<br /> +<br /> +<br /> + +<p><span class="pagenum"><a name="Page_253" id="Page_253">[Pg_253]</a></span></p> + +<div class="caption1">A Revision of Snakes of the Genus Conophis</div> +<div class="caption1">(Family Colubridae, from Middle America)</div> + +<div class="caption3">BY</div> + +<div class="caption2">JOHN WELLMAN</div> +<br /> +<br /> + +<div class="caption2">CONTENTS</div> + +<table width="100%" summary="Contents"> +<tr> + <td> </td> + <td class="text_rt vtop">PAGE</td> +</tr> +<tr> + <td class="smcap"><a href="#INTRODUCTION">Introduction</a></td> + <td class="text_rt vtop">253</td> +</tr> +<tr> + <td colspan="2"> </td> +</tr> +<tr> + <td class="smcap"><a href="#ACKNOWLEDGMENTS">Acknowledgments</a></td> + <td class="text_rt vtop">254</td> +</tr> +<tr> + <td colspan="2"> </td> +</tr> +<tr> + <td class="smcap"><a href="#MATERIALS_AND_METHODS">Materials and Methods</a></td> + <td class="text_rt vtop">254</td> +</tr> +<tr> + <td colspan="2"> </td> +</tr> +<tr> + <td><a href="#Genus_Conophis_Peters"><span class="smcap">Genus</span> Conophis Peters</a></td> + <td class="text_rt vtop">255</td> +</tr> +<tr> + <td> <a href="#Key_to_the_Species_and_Subspecies">Key to the Species and Subspecies</a></td> + <td class="text_rt vtop">257</td> +</tr> +<tr> + <td> <a href="#Analysis_of_Characters">Analysis of Characters</a></td> + <td class="text_rt vtop">257</td> +</tr> +<tr> + <td> <a href="#Scutellation">Scutellation</a></td> + <td class="text_rt vtop">258</td> +</tr> +<tr> + <td> <a href="#Size_and_Proportions">Size and Proportions</a></td> + <td class="text_rt vtop">258</td> +</tr> +<tr> + <td> <a href="#Color_Pattern">Color Pattern</a></td> + <td class="text_rt vtop">260</td> +</tr> +<tr> + <td> <a href="#Sexual_Dimorphism">Sexual Dimorphism</a></td> + <td class="text_rt vtop">260</td> +</tr> +<tr> + <td> <a href="#Conophis_lineatus"><i>C. lineatus</i></a></td> + <td class="text_rt vtop">262</td> +</tr> +<tr> + <td> <a href="#Conophis_lineatus_dunni"><i>C. lineatus dunni</i></a></td> + <td class="text_rt vtop">262</td> +</tr> +<tr> + <td> <a href="#Conophis_lineatus_lineatus"><i>C. lineatus lineatus</i></a></td> + <td class="text_rt vtop">267</td> +</tr> +<tr> + <td> <a href="#Conophis_lineatus_concolor"><i>C. lineatus concolor</i></a></td> + <td class="text_rt vtop">270</td> +</tr> +<tr> + <td> <a href="#Conophis_nevermanni"><i>C. nevermanni</i></a></td> + <td class="text_rt vtop">272</td> +</tr> +<tr> + <td> <a href="#Conophis_pulcher"><i>C. pulcher</i></a></td> + <td class="text_rt vtop">274</td> +</tr> +<tr> + <td> <a href="#Conophis_vittatus"><i>C. vittatus</i></a></td> + <td class="text_rt vtop">277</td> +</tr> +<tr> + <td> <a href="#Skull">Skull</a></td> + <td class="text_rt vtop">282</td> +</tr> +<tr> + <td> <a href="#Dentition">Dentition</a></td> + <td class="text_rt vtop">288</td> +</tr> +<tr> + <td> <a href="#Vertebrae">Vertebrae</a></td> + <td class="text_rt vtop">288</td> +</tr> +<tr> + <td> <a href="#Hemipenes">Hemipenes</a></td> + <td class="text_rt vtop">289</td> +</tr> +<tr> + <td> <a href="#Food_and_Feeding">Food and Feeding</a></td> + <td class="text_rt vtop">289</td> +</tr> +<tr> + <td> <a href="#Effect_of_Poison">Effect of Poison</a></td> + <td class="text_rt vtop">290</td> +</tr> +<tr> + <td colspan="2"> </td> +</tr> +<tr> + <td><div class="smcap"><a href="#TAXONOMIC_RELATIONSHIPS_AND_EVOLUTION">Taxonomic Relationships and Evolution</a></div></td> + <td class="text_rt vtop">29</td> +</tr> +<tr> + <td colspan="2"> </td> +</tr> +<tr> + <td><div class="smcap"><a href="#SUMMARY">Summary</a></div></td> + <td class="text_rt vtop">292</td> +</tr> +<tr> + <td colspan="2"> </td> +</tr> +<tr> + <td><div class="smcap"><a href="#LITERATURE_CITED">Literature Cited</a></div></td> + <td class="text_rt vtop">293</td> +</tr> +</table> +<br /> + + +<div class="caption2"><a name="INTRODUCTION" id="INTRODUCTION"></a> +INTRODUCTION</div> + +<p>Need for a comprehensive systematic review of the snakes of the +genus <i>Conophis</i> 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.<span class="pagenum"><a name="Page_254" id="Page_254">[Pg_254]</a></span></p> + +<div class="caption2"><a name="ACKNOWLEDGMENTS" id="ACKNOWLEDGMENTS"></a> +ACKNOWLEDGMENTS</div> + +<p>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.</p> + +<p>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.</p> + + +<div class="caption2"><a name="MATERIALS_AND_METHODS" id="MATERIALS_AND_METHODS"></a> +MATERIALS AND METHODS</div> + +<p>Of the 325 specimens of the genus <i>Conophis</i> 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.</p> + +<p>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.</p> + +<p>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 +<span class="pagenum"><a name="Page_255" id="Page_255">[Pg_255]</a></span> +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.</p> + +<p>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.</p> + +<p>Except in three dried skeletons, teeth were counted on dentigerous bones +<i>in situ</i>. Since teeth are often missing, the sockets were counted in order to +obtain an accurate count.</p> + +<p>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).</p> + +<p>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.</p> + +<p>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."</p> + +<p>The abbreviations for the museum collections are:</p> + +<table summary="abbreviations"> +<tr> +<td>AMNH</td> +<td>American Museum of Natural History</td> +</tr> +<tr> +<td>ANSP</td> +<td>Academy of Natural Sciences of Philadelphia</td> +</tr> +<tr> +<td>BMNH</td> +<td>British Museum (Natural History)</td> +</tr> +<tr> +<td>CAS</td> +<td>California Academy of Sciences</td> +</tr> +<tr> +<td>CNHM</td> +<td>Chicago Natural History Museum</td> +</tr> +<tr> +<td>ERA-WTN</td> +<td>E. Ross Allen-Wilfred T. Neill, Ross Allen's Reptile Institute</td> +</tr> +<tr> +<td>KU</td> +<td>University of Kansas Museum of Natural History</td> +</tr> +<tr> +<td>MCZ</td> +<td>Museum of Comparative Zoology, Harvard</td> +</tr> +<tr> +<td>MVZ</td> +<td>Museum of Vertebrate Zoology, University of California</td> +</tr> +<tr> +<td>NMW</td> +<td>Naturhistorisches Museums Wien, Vienna</td> +</tr> +<tr> +<td>SU</td> +<td>Stanford University Natural History Museum</td> +</tr> +<tr> +<td>TCWC</td> +<td>Texas Cooperative Wildlife Collection, Agricultural and + Mechanical College of Texas</td> +</tr> +<tr> +<td>UF</td> +<td>University of Florida Collections</td> +</tr> +<tr> +<td>UIMNH</td> +<td>University of Illinois Museum of Natural History</td> +</tr> +<tr> +<td>UMMZ</td> +<td>University of Michigan Museum of Zoology</td> +</tr> +<tr> +<td>USNM</td> +<td>United States National Museum</td> +</tr> +</table> +<br /> +<br /> + +<div class="caption2">Family <span class="smcap">Colubridae</span></div> + +<div class="caption2">Subfamily Xenodontinae</div> + +<div class="caption2"><a name="Genus_Conophis_Peters" id="Genus_Conophis_Peters"></a> +Genus <b>Conophis</b> Peters</div> + +<div class="species"><i>Tomodon</i> (part) Duméril, Bibron and Duméril, +<ins title='Correction: was "Érpétologie Genérale"'>Erpétologie Générale</ins>, +7(pt.2):936, February 25, 1854 (<i>lineatus</i> and <i>vittatus</i>); Salvin, Proc. +Zool. Soc. London, 28:455, 1860 (<i>pulcher</i>).</div> + +<p><span class="pagenum"><a name="Page_256" id="Page_256">[Pg_256]</a></span></p> + +<div class="species"><i>Psammophis</i> (part), Günther, Catalogue of Colubrine Snakes in the Collection +of the British Museum, London, 1858:135 (<i>lineatus</i>).</div> + +<div class="species"><i>Conophis</i> Peters, Monatsb. Akad. Wiss. Berlin, 1860:519-520, pl., fig. 3 +(<i>vittatus</i>); Cope, Proc. Acad. Nat. Sci. Philadelphia, 13:300, December +28, 1861 (<i>lineatus concolor</i>); Proc. Acad. Nat. Sci. Philadelphia, +18:318-319, February 20, 1867 (<i>lineatus concolor</i>); Proc. Acad. Nat. Sci. +Philadelphia, ser. 2, 8:137, 1876 (<i>pulcher</i>); Bocourt in Duméril, Bocourt +and Mocquard, Mission Scientifique au Mexique et dans l'Amerique +Centrale, 2:643-644, pl. 38, fig. 5, 1886 (<i>lineatus lineatus</i>); 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 +(<i>nevermanni</i>).</div> + +<div class="species"><i>Tachymenis</i> (in part), Garman, Bull. Essex Inst., 16:33, January 9, 1884 +(<i>vittatus</i> and <i>lineatus</i>).</div> + +<div class="species"><i>Erythrolamprus</i> (in part), Ditmars, Bull. Antivenin Inst. Amer., 2(2):27-29, +June.</div> + +<div class="species"><i>Coniophanes</i> (in part), Wettstein, Sitz. Akad. Wiss. Wien, mathem-naturw. +kl. 143:37-38, 1934 (<i>nevermanni</i>).</div> + +<p><i>Historical summary.</i>—In 1854 Duméril, Bibron and Duméril described and +figured <i>Tomodon lineatum</i> from America. In 1860 Peters described and +figured as a new genus and species, <i>Conophis vittatus</i>, 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 <i>lineatus</i> in +the genus <i>Conophis</i>. Cope (1861) proposed the name <i>Conophis vittatus</i> (<i>nec</i> +Peters, 1860). Later (1900) he changed its name to <i>Conophis lineaticeps</i>. +Early uncertainty of the relationships of the species <i>lineatus</i> caused Günther +(1858) to place it in the genus <i>Psammophis</i>. With the exception of Garman +(1884a and 1884b) who placed <i>lineatus</i> in the genus <i>Tachymenis</i>, and Wettstein +(1934) who reported five specimens of <i>Conophis nevermanni</i> as <i>Coniophanes +i. imperialis</i>, all specimens reported after 1876 were placed in the genus +<i>Conophis</i>.</p> + +<p>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 (<i>nevermanni</i>).</p> + +<p><i>Description.</i>—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 +<span class="pagenum"><a name="Page_257" id="Page_257">[Pg_257]</a></span> +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.</p> + +<p><i>Distribution.</i>—Semi-arid regions of southern México and Central America +as far south as Costa Rica.</p> +<br /> + +<div class="smcap center"><a name="Key_to_the_Species_and_Subspecies" id="Key_to_the_Species_and_Subspecies"></a> +Key to the Species and Subspecies</div> + +<p>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.</p> + +<table width="100%" summary="Species Key"> +<tr> + <td class="vtop">1. </td> + <td>Seven supralabials (3rd and 4th below orbit); 3 to 8 dark + stripes along body</td> +</tr> +<tr> + <td colspan="2" class="text_rt">2</td> +</tr> +<tr> + <td> </td> + <td>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</td> +</tr> +<tr> + <td colspan="2" class="text_rt">3</td> +</tr> +<tr> + <td class="vtop">2. </td> + <td>Dark stripes involving no more than one + longitudinal scale-row</td> +</tr> +<tr> + <td colspan="2" class="text_rt"><a href="#Conophis_lineatus_lineatus"><i>C. lineatus lineatus</i> (part), p. 267</a></td> +</tr> +<tr> + <td> </td> + <td>Dark stripes involving at least + two adjacent scale-rows</td> +</tr> +<tr> + <td colspan="2" class="key_text_rt"><a href="#Conophis_vittatus"><i>C. vittatus</i>, p. 277</a></td> +</tr> +<tr> + <td class="vtop">3. </td> + <td>Supralabials having black borders above; + head and body generally black with 2 or 4 white lines running length + of body</td> +</tr> +<tr> + <td colspan="2" class="key_text_rt"><a href="#Conophis_nevermanni"><i>C. nevermanni</i>, p. 272</a></td> +</tr> +<tr> + <td> </td> + <td>Supralabials immaculate or having + dark borders below; head and body usually pale with dark stripes, or + without stripes</td> +</tr> +<tr> + <td colspan="2" class="key_text_rt">4</td> +</tr> +<tr> + <td class="vtop">4. </td> + <td>Lateral dark stripe through eye involving + upper half of second scale-row; dark stripe on paravertebral row, at + least posteriorly</td> +</tr> +<tr> + <td colspan="2" class="key_text_rt"><a href="#Conophis_pulcher"><i>C. pulcher</i>, p. 274</a></td> +</tr> +<tr> + <td> </td> + <td>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</td> +</tr> +<tr> + <td colspan="2" class="key_text_rt">5</td> +</tr> +<tr> + <td class="vtop">5. </td> + <td>Stripes disappearing posteriorly + (except for small spots of pigment on scale-row 4 or 7); + 1st scale-row unpigmented</td> +</tr> +<tr> + <td colspan="2" class="key_text_rt"><a href="#Conophis_lineatus_concolor"><i>C. lineatus concolor</i>, p. 270</a></td> +</tr> +<tr> + <td> </td> + <td>Stripes present posteriorly; + 1st scale-row pigmented</td> +</tr> +<tr> + <td colspan="2" class="key_text_rt">6</td> +</tr> +<tr> + <td class="vtop">6. </td> + <td>Lateral stripes narrow on nape, restricted + to 4th scale-row on body</td> +</tr> +<tr> + <td colspan="2" class="key_text_rt"><a href="#Conophis_lineatus_lineatus"><i>C. lineatus lineatus</i> (part), p. 267</a></td> +</tr> +<tr> + <td> </td> + <td>Lateral stripes involving 3rd and 4th rows, at least on nape</td> +</tr> +<tr> + <td colspan="2" class="key_text_rt"><a href="#Conophis_lineatus_dunni"><i>C. lineatus dunni</i>, p. 262</a></td> +</tr> +</table> +<br /> +<div class="caption3"><a name="Analysis_of_Characters" id="Analysis_of_Characters"></a> +Analysis of Characters</div> + +<p>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.</p> + +<p><span class="pagenum"><a name="Page_258" id="Page_258">[Pg_258]</a></span></p> +<div class="caption3"><a name="Scutellation" id="Scutellation"></a> +Scutellation</div> + +<p>Labials, dorsals, ventrals, and subcaudals were the most useful scales.</p> + +<p><i>Labials.</i>—All species usually have eight supralabials except <i>C. vittatus</i>, +which has seven. The only other population having a relatively high frequency +of occurrence of seven supralabials is <i>C. l. lineatus</i>. 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.</p> + +<p><i>Dorsals.</i>—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).</p> + +<div class="smcap">Table 1.—Variation in the Place of Dorsal Scale Reduction in +Conophis.</div> + +<table width="100%" style="height:9px; border-collapse: collapse" summary="Sexual Dimorphism"> +<tr> + <td class="brdtp2 brdbt center">Taxon</td> + <td class="brdtp2 brdbt brdlf center">Number of Specimens</td> + <td class="brdtp2 brdbt brdlf center">Range</td> + <td class="brdtp2 brdbt brdlf center">Mean</td> + <td class="brdtp2 brdbt brdlf center">Standard Deviation</td> + <td class="brdtp2 brdbt brdlf center">Standard Error</td> + <td class="brdtp2 brdbt brdlf center">Coefficient of Variation</td> +</tr> +<tr> + <td><i>l. concolor</i></td> + <td class="brdlf center">45</td> + <td class="brdlf center">89-114</td> + <td class="brdlf center">102.5</td> + <td class="brdlf center">5.57</td> + <td class="brdlf center">0.83</td> + <td class="brdlf center">5.43</td> +</tr> +<tr> + <td><i>l. dunni</i></td> + <td class="brdlf center">36</td> + <td class="brdlf center">91-111</td> + <td class="brdlf center">102.1</td> + <td class="brdlf center">4.59</td> + <td class="brdlf center">0.77</td> + <td class="brdlf center">4.50</td> +</tr> +<tr> + <td><i>l. lineatus</i></td> + <td class="brdlf center">26</td> + <td class="brdlf center">91-107</td> + <td class="brdlf center">100.2</td> + <td class="brdlf center">3.59</td> + <td class="brdlf center">0.72</td> + <td class="brdlf center">3.58</td> +</tr> +<tr> + <td><i>nevermanni</i></td> + <td class="brdlf center">6</td> + <td class="brdlf center">84- 97</td> + <td class="brdlf center">93.2</td> + <td class="brdlf center">4.71</td> + <td class="brdlf center">1.92</td> + <td class="brdlf center">5.05</td> +</tr> +<tr> + <td><i>pulcher</i></td> + <td class="brdlf center">26</td> + <td class="brdlf center">94-119</td> + <td class="brdlf center">104.6</td> + <td class="brdlf center">4.90</td> + <td class="brdlf center">0.96</td> + <td class="brdlf center">4.68</td> +</tr> +<tr> + <td class="brdbt"><i>vittatus</i></td> + <td class="brdbt brdlf center">170</td> + <td class="brdbt brdlf center">84-118</td> + <td class="brdbt brdlf center">102.3</td> + <td class="brdbt brdlf center">6.60</td> + <td class="brdbt brdlf center">0.16</td> + <td class="brdbt brdlf center">6.45</td> +</tr> +</table> + +<p><i>Ventrals.</i>—The number of ventral scutes varies from 149-183, and shows +no significant variation in the means (table 2).</p> + +<p><i>Subcaudals.</i>—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.</p> + + +<div class="caption3"><a name="Size_and_Proportions" id="Size_and_Proportions"></a> +Size and Proportions</div> + +<p>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 <i>C. lineatus concolor</i>; all +others are smaller and of about the same size and proportions. The longest +specimen, a male of <i>C. l. concolor</i>, has a body length of 893 mm., a tail length +of 274 mm., and a total length of 1167 mm.</p> + +<p><span class="pagenum"><a name="Page_259" id="Page_259">[Pg_259]</a></span></p> + +<div class="smcap">Table 2.—Variation in the Number of Ventrals in Conophis.</div> + +<table width="100%" style="height:9px; border-collapse: collapse" summary="Sexual Dimorphism"> +<tr> + <td class="brdtp2 brdbt center">Taxon</td> + <td class="brdtp2 brdbt brdlf center">Number of Specimens</td> + <td class="brdtp2 brdbt brdlf center">Range</td> + <td class="brdtp2 brdbt brdlf center">Mean</td> + <td class="brdtp2 brdbt brdlf center">Standard Deviation</td> + <td class="brdtp2 brdbt brdlf center">Standard Error</td> + <td class="brdtp2 brdbt brdlf center">Coefficient of Variation</td> +</tr> +<tr> + <td><i>l. concolor</i></td> + <td class="brdlf center"> 45</td> + <td class="brdlf center">158-170</td> + <td class="brdlf center">163.7</td> + <td class="brdlf center">1.56</td> + <td class="brdlf center">0.23</td> + <td class="brdlf center">0.95</td> +</tr> +<tr> + <td><i>l. dunni</i></td> + <td class="brdlf center">36</td> + <td class="brdlf center">159-178</td> + <td class="brdlf center">167.2</td> + <td class="brdlf center">4.56</td> + <td class="brdlf center">0.76</td> + <td class="brdlf center">2.72</td> +</tr> +<tr> + <td><i>l. lineatus</i></td> + <td class="brdlf center">26</td> + <td class="brdlf center">157-169</td> + <td class="brdlf center">163.5</td> + <td class="brdlf center">3.59</td> + <td class="brdlf center">0.72</td> + <td class="brdlf center">2.20</td> +</tr> +<tr> + <td><i>nevermanni</i> </td> + <td class="brdlf center">6</td> + <td class="brdlf center">173-183</td> + <td class="brdlf center">176.5</td> + <td class="brdlf center">4.00</td> + <td class="brdlf center">1.63</td> + <td class="brdlf center">2.27</td> +</tr> +<tr> + <td><i>pulcher</i></td> + <td class="brdlf center"> 26</td> + <td class="brdlf center">149-180</td> + <td class="brdlf center">169.5</td> + <td class="brdlf center">5.31</td> + <td class="brdlf center">1.04</td> + <td class="brdlf center">3.13</td> +</tr> +<tr> + <td class="brdbt"><i>vittatus</i></td> + <td class="brdbt brdlf center">171</td> + <td class="brdbt brdlf center">149-180</td> + <td class="brdbt brdlf center">163.7</td> + <td class="brdbt brdlf center">6.33</td> + <td class="brdbt brdlf center">0.15</td> + <td class="brdbt brdlf center">3.87</td> +</tr> +</table> +<br /> + +<div class="smcap">Table 3.—Sexual Dimorphism as Indicated by Variation in the Number<br /> +of Subcaudals in Conophis</div> + +<table width="100%" style="height:9px; border-collapse: collapse" summary="Sexual Dimorphism"> +<tr> + <td class="brdtp2 brdbt center">Taxon</td> + <td class="brdtp2 brdbt brdlf center">Sex</td> + <td class="brdtp2 brdbt brdlf center">Number of Specimens</td> + <td class="brdtp2 brdbt brdlf center">Range</td> + <td class="brdtp2 brdbt brdlf center">Mean</td> + <td class="brdtp2 brdbt brdlf center">Standard Deviation</td> + <td class="brdtp2 brdbt brdlf center">Standard Error</td> + <td class="brdtp2 brdbt brdlf center">Coefficient of Variation</td> + <td class="brdtp2 brdbt brdlf center">Coefficient of Difference</td> +</tr> +<tr> + <td><i>lineatus concolor</i></td> + <td class="center brdlf">♂</td> + <td class="center brdlf">22</td> + <td class="center brdlf">68-74</td> + <td class="center brdlf">70.3</td> + <td class="center brdlf">2.14</td> + <td class="center brdlf">0.46</td> + <td class="center brdlf">3.04</td> + <td class="brdlf"> </td> +</tr> +<tr style="height:9px"> + <td> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="center brdlf">1.97</td> +</tr> +<tr style="height:9px"> + <td> </td> + <td class="center brdlf">♀</td> + <td class="center brdlf">16</td> + <td class="center brdlf">56-65</td> + <td class="center brdlf">61.8</td> + <td class="center brdlf">2.18</td> + <td class="center brdlf">0.55</td> + <td class="center brdlf">3.53</td> + <td class="brdlf"> </td> +</tr> +<tr> + <td> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> +</tr> +<tr> + <td><i>lineatus dunni</i></td> + <td class="center brdlf">♂</td> + <td class="center brdlf">14</td> + <td class="center brdlf">67-80</td> + <td class="center brdlf">74.5</td> + <td class="center brdlf">3.86 </td> + <td class="center brdlf">1.03</td> + <td class="center brdlf">5.18</td> + <td class="brdlf"> </td> +</tr> +<tr> + <td> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="center brdlf">0.95</td> +</tr> +<tr> + <td> </td> + <td class="center brdlf">♀</td> + <td class="center brdlf">16</td> + <td class="center brdlf">60-72</td> + <td class="center brdlf">67.1</td> + <td class="center brdlf">3.91</td> + <td class="center brdlf">0.97</td> + <td class="center brdlf">5.82</td> + <td class="brdlf"> </td> +</tr> +<tr> + <td> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> +</tr> +<tr> + <td><i>lineatus lineatus</i></td> + <td class="center brdlf">♂</td> + <td class="center brdlf">11</td> + <td class="center brdlf">67-73</td> + <td class="center brdlf">69.8</td> + <td class="center brdlf">6.17</td> + <td class="center brdlf">1.85</td> + <td class="center brdlf">8.84</td> + <td class="brdlf"> </td> +</tr> +<tr> + <td> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="center brdlf">0.60</td> +</tr> +<tr> + <td> </td> + <td class="center brdlf">♀</td> + <td class="center brdlf">9</td> + <td class="center brdlf">60-66</td> + <td class="center brdlf">62.4</td> + <td class="center brdlf">6.17</td> + <td class="center brdlf">2.06</td> + <td class="center brdlf">9.89</td> + <td class="brdlf"> </td> +</tr> +<tr> + <td> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> +</tr> +<tr> + <td><i>nevermanni</i></td> + <td class="center brdlf">♂</td> + <td class="center brdlf">3</td> + <td class="center brdlf">82-89</td> + <td class="center brdlf">85.3</td> + <td class="center brdlf">……</td> + <td class="center brdlf">……</td> + <td class="center brdlf">……</td> + <td class="brdlf"> </td> +</tr> +<tr> + <td> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="center brdlf">……</td> +</tr> +<tr> + <td> </td> + <td class="center brdlf">♀</td> + <td class="center brdlf">2</td> + <td class="center brdlf">71-76</td> + <td class="center brdlf">73.5</td> + <td class="center brdlf">……</td> + <td class="center brdlf">……</td> + <td class="center brdlf">……</td> + <td class="brdlf"> </td> +</tr> +<tr> + <td> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> +</tr> +<tr> + <td><i>pulcher</i></td> + <td class="center brdlf">♂</td> + <td class="center brdlf">7</td> + <td class="center brdlf">70-79</td> + <td class="center brdlf">74.3</td> + <td class="center brdlf">3.11</td> + <td class="center brdlf">1.17</td> + <td class="center brdlf">4.19</td> + <td class="brdlf"> </td> +</tr> +<tr> + <td> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="center brdlf">0.93</td> +</tr> +<tr> + <td> </td> + <td class="center brdlf">♀</td> + <td class="center brdlf">11</td> + <td class="center brdlf">65-71</td> + <td class="center brdlf">68.2</td> + <td class="center brdlf">3.42</td> + <td class="center brdlf">1.08</td> + <td class="center brdlf">5.01</td> + <td class="brdlf"> </td> +</tr> +<tr> + <td> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> +</tr> +<tr> + <td><i>vittatus</i></td> + <td class="center brdlf">♂</td> + <td class="center brdlf">95</td> + <td class="center brdlf">59-76</td> + <td class="center brdlf">67.8</td> + <td class="center brdlf">3.33</td> + <td class="center brdlf">0.34</td> + <td class="center brdlf">4.91</td> + <td class="brdlf"> </td> +</tr> +<tr> + <td> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="brdlf"> </td> + <td class="center brdlf">1.28</td> +</tr> +<tr> + <td class="brdbt"> </td> + <td class="brdbt brdlf center">♀</td> + <td class="brdbt brdlf center">58</td> + <td class="brdbt brdlf center">55-66</td> + <td class="brdbt brdlf center">60.0</td> + <td class="brdbt brdlf center">2.75</td> + <td class="brdbt brdlf center">0.36</td> + <td class="brdbt brdlf center">4.58</td> + <td class="brdbt brdlf center"> </td> +</tr> +</table> +<br /> + +<p><span class="pagenum"><a name="Page_260" id="Page_260">[Pg_260]</a></span></p> + +<div class="caption3"><a name="Color_Pattern" id="Color_Pattern"></a> +Color Pattern</div> + +<p>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 <i>vittatus</i>, and as many as 13 in <i>l. dunni</i>; except that there is none in <i>C. l. +concolor</i>. 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.</p> + +<p>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.</p> + +<p>In some specimens of <i>Conophis</i> the lateral tips of the ventrals are spotted, +one spot on each end of each ventral. Otherwise, the ventrals are immaculate +white.</p> + +<p>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 <i>C. lineatus dunni</i> and <i>C. pulcher</i> +can be separated on the basis of which scale-rows are darkly pigmented. +<i>C. l. dunni</i> 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.</p> + + +<div class="caption3"><a name="Sexual_Dimorphism" id="Sexual_Dimorphism"></a> +Sexual Dimorphism</div> + +<p>Sexual dimorphism is evident in all species and subspecies of +<i>Conophis</i>. 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.</p> + +<p>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, +<span class="pagenum"><a name="Page_261" id="Page_261">[Pg_261]</a></span> +that seem to be part of a continuous or internal cline (Huxley, +1942) are not used for characterizing subspecies.</p> + +<p><span class="pagenum"><a name="Page_262" id="Page_262">[Pg_262]</a></span></p> +<a name="Fig_1" id="Fig_1"></a> +<div class="fig_center" style="width: 499px;"> +<img src="images/fig_1.png" width="499" height="603" alt="" title="" /> +<div class="fig_caption"> +<span class="smcap">Fig. 1.</span> 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.</div> +</div> +<br /> + + +<div class="caption3"><a name="Conophis_lineatus" id="Conophis_lineatus"></a> +<b>Conophis lineatus</b> (Duméril, Bibron and Duméril)</div> + +<div class="species"> +<p><i>Tomodon lineatum</i> (in part) Duméril, Bibron and Duméril, +<ins title='Correction: was "Érpétologie Genérale"'>Erpétologie Générale</ins>, +7(pt. 2):936-938, February 25, 1854.</p> +</div> + +<p><i>Diagnosis.</i>—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.</p> + +<p><i>Variation.</i>—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 <i>C. l. concolor</i> (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 +<i>C. l. dunni</i> (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.</p> + +<p>The greatest variation is in coloration. Dark color, or lack thereof, has +been used to separate the subspecies of <i>C. lineatus</i>. 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.</p> + +<p>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.</p> + +<p>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) (<a href="#Fig_2">Fig. 2</a>). 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.</p> + + +<div class="caption3"><a name="Conophis_lineatus_dunni" id="Conophis_lineatus_dunni"></a> +<b>Conophis lineatus dunni</b> Smith</div> + +<div class="species"> +<i>Psammophis lineatus</i>, Günther, Catalogue of Colubrine Snakes in the Collection +of the British Museum, p. 135, 1858. +</div> + +<p><span class="pagenum"><a name="Page_263" id="Page_263">[Pg_263]</a></span></p> + +<a name="Fig_2" id="Fig_2"></a> +<div class="fig_center" style="width: 503px;"> +<img src="images/fig_2.png" width="503" height="456" alt="" title="" /> +<div class="fig_caption"> +<span class="smcap">Fig. 2.</span> Selected locality records for the subspecies of <i>Conophis lineatus</i>.</div> +</div> +<br /> + +<div class="species"> +<p><i>Conophis lineatus</i>, 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.</p> + +<p><i>Conophis lineatus similis</i> Smith, Journ. Washington Acad. Sci., 31:123-124, +March 15, 1941 (Type.—United States National Museum, No. 79963; +type locality.—Managua, Nicaragua; <i>nec</i> Bocourt <i>in</i> 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.</p> + +<p><i>Conophis lineatus dunni</i> 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.</p> + +<p><i>Conophis pulcher pulcher</i>, Stuart, Misc. Publ. Mus. Zool. Univ. Michigan, +69:79, June 12, 1948; Contr. Lab. Vert. Biol. Univ. Michigan, 45:24, +<span class="pagenum"><a name="Page_264" id="Page_264">[Pg_264]</a></span> +May, 1950; Contr. Lab. Vert. Biol. Univ. Michigan, 49:14, August, 1951; +Contr. Lab. Vert. Biol. Univ. Michigan, 65:19-20 (part), March, 1954.</p> + +<p><i>Conophis pulcher plagosus</i>, Mertens, Zool. Anz., 148:93, February, 1952; +Abhand. Senken. Naturw. Gesell., 487:61-62, December 1, 1952.</p> + +<p><i>Conophis lineatus nevermanni</i>, Taylor, Univ. Kansas Sci. Bull., 37(pt. 1):563-565, +fig. 16, October 15, 1955.</p> +</div> + +<p><i>Type.</i>—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).</p> + +<p><i>Diagnosis.</i>—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 +<ins title='Correction: was "immaculaate"'>immaculate</ins> or having +dark ventral margins.</p> + +<p><i>Variation.</i>—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.</p> + +<p>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 +(<a href="#Fig_1">fig. 1, C</a>). 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 (<a href="#Fig_1">fig. 1, A</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.</p> + +<p>The primary stripes, <ins title='Correction: was "chacteristic"'>characteristic</ins> +of the species <i>lineatus</i>, 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 +<span class="pagenum"><a name="Page_265" id="Page_265">[Pg_265]</a></span> +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 <i>C. lineatus dunni</i>. 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.</p> + +<p>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.</p> + +<p>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.</p> + +<p>Probably the most common pattern of dorsal coloration consists of eight +or ten dark stripes (<a href="#Fig_1">fig. 1, B</a>). 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.</p> + +<p>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.</p> + +<a name="Fig_3" id="Fig_3"></a> +<table width="80%" class="center" summary="img_frame"> +<tr> + <td><img src="images/fig_3.png" width="247" height="165" alt="" title="" /></td> + <td> </td> + <td class="justify"><span class="smcap">Fig.</span> 3. Patterns of dorsal coloration + at mid-body of juveniles of two sympatric species of Conophis. + A. <i>C. lineatus dunni</i> (MCZ 49794) from Tegucigalpa, Honduras. + B. <i>C. pulcher</i> (MCZ 49791) from Tegucigalpa, Honduras. + Approximately × 1.</td> +</tr> +</table> +<br /> + +<p>In juveniles, there are six or eight black stripes boldly contrasting with a +white or pale tan ground-color (<a href="#Fig_3">fig. 3, A</a>). The first pair of stripes is on the +<span class="pagenum"><a name="Page_266" id="Page_266">[Pg_266]</a></span> +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.</p> + +<div class="larger"> +<p><i>Remarks.</i>—Savage (1949:483-486) stated that his specimen of +<i>C. l. dunni</i> (from Honduras) resembled <i>l. lineatus</i> 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 <i>l. dunni</i>. +The specimen in no way represents an intergrade between <i>C. l. +dunni</i> and <i>l. lineatus</i>.</p> + +<p>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 <i>C. l. dunni</i> 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.</p> + +<p>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 <i>C. l. dunni</i> from all other +snakes in México and Central America. The only sympatric species +of this genus, <i>C. pulcher</i>, differs in that it has paravertebral stripes +(though never a vertebral dark stripe). <i>Conophis pulcher</i> has a +lateral dark stripe that includes the upper half of the second +scale-row on the anterior part of the body; stripes of <i>C. l. dunni</i> +never include more than the 3rd and 4th rows. Even as juveniles +the paravertebral row is not darkly pigmented in <i>C. l. dunni</i> as it +is in <i>C. pulcher</i>.</p> +</div> + +<p><i>Distribution.</i>—Semi-arid habitats from sea level to +<ins title='Correction: was "elevatons"'>elevations</ins> 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 (<a href="#Fig_2">fig. 2</a>).</p> + +<p><i>Specimens examined.</i>—Total of 41 specimens, as follows: +<span class="smcap">British Honduras</span>: +<i>Cayo District</i>: Augustine, ERA-WTN BH-300; <i>Mountain Pine Ridge, +10 mi. E Augustine</i>, ERA-WTN BH-298.</p> + +<p><span class="smcap">Costa Rica</span>: <i>no specific locality</i>, AMNH 17309. "<i>Cartago</i>," BMNH +71.11.22.15. <i>Puntarenas</i>: 32 km. N Barranca, KU 35630; Esparta, USNM +37758. "<i>San José</i>," ANSP 3480, 12232.</p> + +<p><span class="smcap">El Salvador</span>: <i>Morazan</i>: El Divisadero, CNHM 10999. <i>San Miguel: San +Pedro</i>, MCZ 57061.</p> + +<p><span class="smcap">Guatemala</span>: <i>El Petén</i>: Sojio (Toocog), AMNH 69969, 69986. <i>Huehuetenango</i>: +flood plain Río Cuilco, W of Finca Canibal, 18 km. N Tacaná, +UMMZ 98283. <i>Santa Rosa</i>: Santa Rosa, UMMZ 107339.</p> + +<p><span class="pagenum"><a name="Page_267" id="Page_267">[Pg_267]</a></span> +<span class="smcap">Honduras</span>: <i>no specific locality</i>, AMNH 32814, UF 7657. <i>Cortes: Cofradía</i>, +SU 8422; <i>Gracias</i>, CNHM 28560; <i>Hacienda de Santa Ana, W San Pedro +Sula</i>, CNHM 5297; San Pedro Sula, UMMZ 68695(2); <i>near San Pedro Sula</i>, +MCZ 27563. <i>Francisco Morazan: Potrero de Melio, Escuela Agricola Pan-Americana</i>, +MCZ 49987; Tegucigalpa, MCZ 49784, 49786, 49789-90, 49792, 49794.</p> + +<p><span class="smcap">Mexico</span>: <i>no specific locality</i>, BMNH 53.2.4.16.</p> + +<p><span class="smcap">Nicaragua</span>: <i>no specific locality</i>, UMMZ 65633, USNM 25237. +<i>Leon</i>: El Polvón, MCZ 5645, 5696. <i>Managua</i>: Managua, USNM 79963-64; <i>3 mi. SW +Managua</i>, KU 42315; <i>8 mi. WNW Managua</i>, KU 42314; <i>1 mi. N Sabana +Grande</i>, KU 42311-13. <i>Matagalpa</i>: 1.5 mi. N Matagalpa, UMMZ 116537.</p> + + +<div class="caption3"><a name="Conophis_lineatus_lineatus" id="Conophis_lineatus_lineatus"></a> +<b>Conophis lineatus lineatus</b> (<ins title='Correction: was "Dumeril"'>Duméril</ins>, Bibron and +<ins title='Correction: was "Dumeril"'>Duméril</ins>)</div> + +<div class="species"> +<p><i>Tomodon lineatum</i> (in part) Duméril, Bibron and Duméril, +<ins title='Correction: was "Érpétologie Genérale"'>Erpétologie Générale</ins>, +7(pt. 2):936-938, atlas, pl. 73, February 25, 1854; Bocourt, +Journ. de Zool., 5:406-407, 1876.</p> + +<p><i>Tomodon lineatus</i>, 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 Mexico. 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.</p> + +<p><i>Tachymenis lineata</i> (in part), Garman, Bull. Essex Inst., 16: 33, January +9, 1884; Mem. Mus. Comp. Zool., 8:60-61, July, 1884.</p> + +<p><i>Conophis lineatus</i>, Bocourt <i>in</i> 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.</p> + +<p><i>Conophis lineatus lineatus</i>, 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.</p> +</div> + +<p><i>Type.</i>—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 <i>C. l. lineatus</i>. 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 <i>C. l. dunni</i> and <i>C. l. lineatus</i> can be separated.</p> + +<p><i>Diagnosis.</i>—Lateral dark stripe anteriorly passing through eye and posteriorly +involving fourth scale-row only; first scale-row darkly pigmented; no +<span class="pagenum"><a name="Page_268" id="Page_268">[Pg_268]</a></span> +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.</p> + +<p><i>Variation.</i>—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.</p> + +<p>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 (<a href="#Fig_1">fig. 1, D</a>). 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.</p> + +<p>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.</p> + +<p>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.</p> + +<p>Usually there are eight supralabials on each side; however, seven of the +27 specimens examined have seven supralabials on each side, and three others +<span class="pagenum"><a name="Page_269" id="Page_269">[Pg_269]</a></span> +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.</p> + +<p>Variation in coloration and in number of supralabials appears to be of no +geographic significance.</p> + +<p>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.</p> + +<p>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.</p> + +<div class="larger"> +<p><i>Remarks.</i>—In his diagnosis of <i>C. l. lineatus</i>, 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.</p> + +<p>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 <i>C. l. lineatus</i> from +all other snakes in México.</p> +</div> + +<p><i>Distribution.</i>—Semi-arid habitats on the coastal plain of Veracruz, México, +from Tecolutla to Lerdo de Tejada and Piedras Negras (<a href="#Fig_2">fig. 2</a>).</p> + +<p><i>Specimens examined.</i>—Total of 27, as follows: +<span class="smcap">México</span>: <i>no specific locality</i>, +AMNH 19614-15, 19621-24, 19642-43, NMW 16827. <i>Veracruz: no specific +locality</i>, AMNH 19618-20, CAS 73640, NMW 16829; <i>4 km. S Alvarado</i>, KU +58124; <i>14 mi. N Alvarado</i>, UIMNH 46978; 6 mi. SE Boca del Río, UIMNH +28023; Etiopa, 2 mi. S Tecolutla, UIMNH 3847; <i>ca.</i> 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; <i>W side Veracruz</i>, AMNH 19616; <i>2 mi. W +Veracruz</i>, AMNH 19617, 19619.</p> + +<p><span class="pagenum"><a name="Page_270" id="Page_270">[Pg_270]</a></span></p> + + +<div class="caption3"><a name="Conophis_lineatus_concolor" id="Conophis_lineatus_concolor"></a> +<b>Conophis lineatus concolor</b> Cope</div> + +<div class="species"><i>Conophis vittatus</i> Cope, Proc. Acad. Nat. Sci. +Philadelphia, 13:300, December 28, 1861 (<i>nec</i> 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.</div> + +<div class="species"><i>Conophis concolor</i> Cope, Proc. Acad. Nat. Sci. Philadelphia, +18:318-319, February 20, 1867; Journ. Acad. Nat. Sci. Philadelphia, +ser. 2, 8:137, 1876; Bocourt <i>in</i> 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.</div> + +<div class="species"><i>Conophis lineatus</i> (in part), Günther, Biologica +Centrali-Americana, p. 165, March, 1895; Gaige <i>in</i> Pearse, <i>et al.</i> +Carnegie Inst. Washington Publ., 457:302, February 5, 1936.</div> + +<div class="species"><i>Conophis lineaticeps</i> Cope, Ann. Rept. U. S. Natl. Mus. for 1898, +pp. 1094-95, 1900 (Substitute name for <i>Conophis vittatus</i> Cope, +1861, <i>nec</i> Peters, 1860).</div> + +<div class="species"><i>Conophis lineatus concolor</i>, 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.</div> + +<p><i>Types.</i>—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).</p> + +<p><i>Diagnosis.</i>—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.</p> + +<p><i>Variation.</i>—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.</p> + +<p>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.</p> + +<p><span class="pagenum"><a name="Page_271" id="Page_271">[Pg_271]</a></span> +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.</p> + +<div class="larger"> +<p><i>Remarks.</i>—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 "<i>Conophis vittatus</i>, 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 <i>C. +lineaticeps</i> by Cope who thought the specimen differed significantly +from <i>C. concolor</i> (Cope, 1867:318-319). This specimen has the +coloration normal for <i>C. l. concolor</i> as far posteriorly as mid-body; +beyond mid-body the dark lines, typical of <i>C. l. lineatus</i> or of <i>C. l. +dunni</i>, are present. It is likely that this specimen is an intergrade +between <i>C. l. concolor</i> and <i>C. l. dunni</i>, the other subspecies present +in Guatemala.</p> + +<p>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 <i>C. l. +concolor</i> in northern Honduras. This individual may be merely a +genetically aberrant specimen from an area where normal specimens +are <i>C. l. dunni</i>. Neill and Allen (1961:44-45) suggested that +the specimen from Patuca implies widely overlapping distributions +for <i>C. l. dunni</i> and <i>C. concolor</i>. The occurrence of <i>C. l. concolor</i> 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 <i>C. l. dunni</i> and <i>C. l. concolor</i> +could occur through a relatively broad area of El Petén and British +Honduras.</p> + +<p>Neill and Allen (1961:44-45) further suggest that the present +range of <i>C. l. dunni</i> extends "presumably still farther northward +toward the Méxican state of Veracruz where <i>C. l. lineatus</i> exists." +Actually the presence of the subspecies <i>C. l. dunni</i> and <i>C. l. lineatus</i> +as presently disjunct populations implies merely that they were +presumably a continuous population at some time in the past.</p> + +<p>The characteristics of the genus in combination with the reduction +<span class="pagenum"><a name="Page_272" id="Page_272">[Pg_272]</a></span> +of dark coloration posterior to the head distinguish this snake +from all other snakes in México and Central America.</p> +</div> + +<p><i>Distribution.</i>—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 (<a href="#Fig_2">fig. 2</a>).</p> + +<p><i>Specimens examined.</i>—Total of 48, as follows: +<span class="smcap">British Honduras</span>: <i>Belize +District</i>: 13.0 mi. W, 1.5 mi. S Belize, ERA-WTN BH-1562.</p> + +<p><span class="smcap">Guatemala</span>: <i>El Petén, no specific locality</i>, USNM 4941.</p> + +<p><span class="smcap">Honduras</span>: <i>Colón</i>: Patuca, USNM 20271.</p> + +<p><span class="smcap">México</span>: <i>Campeche</i>: Champotón, UMMZ 73063-66; Encarnación, CNHM +106462. <i>Yucatán: no specific locality</i>, 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; <i>Libré Union</i>, +CNHM 36298, 36302, 36310-11, 36314; Mayapán, CNHM 40720; Mérida, +CNHM 19411, 19413, NMW 16828; Progreso, CNHM 40721; Tekom, CNHM +49374; Yokdzonot, CNHM 36315.</p> + + +<div class="caption3"><a name="Conophis_nevermanni" id="Conophis_nevermanni"></a> +<b>Conophis nevermanni</b> Dunn</div> + +<div class="species"><i>Coniophanes imperialis imperialis</i>, Wettstein, Sitz. +Akad. Wiss. Wien. mathem-naturw. Kl., 143:37-38, 1934.</div> + +<div class="species"><i>Conophis nevermanni</i> 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.</div> + +<p><i>Type.</i>—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.</p> + +<p><i>Diagnosis.</i>—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.</p> + +<p><i>Variation.</i>—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.</p> + +<p>The dorsal coloration (<a href="#Fig_1">fig. 1, E</a>) 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.</p> + +<p><span class="pagenum"><a name="Page_273" id="Page_273">[Pg_273]</a></span> +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.</p> + +<p>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.</p> + +<div class="fig_center" style="width: 498px;"> +<a name="Fig_4" id="Fig_4"></a> +<img src="images/fig_4.png" width="498" height="405" alt="" title="" /> +<div class="fig_caption"> +<span class="smcap">Fig. 4.</span> Selected locality records for <i>Conophis pulcher</i> +and <i>Conophis nevermanni</i>.</div> +</div> +<br /> + +<div class="larger"> +<p><i>Remarks.</i>—Taylor (1955:563-565) hesitantly referred a specimen +(KU 35630) from 32 kilometers north of Barranca, Puntarenas +<span class="pagenum"><a name="Page_274" id="Page_274">[Pg_274]</a></span> +Province, Costa Rica, to <i>Conophis lineatus nevermanni</i>. 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 <i>C. nevermanni</i>. Furthermore, the ventral margins +of the supralabials are brown, and the pale dorsal stripes are +tan and too wide for <i>C. nevermanni</i> (compare <a href="#Fig_1">figs. 1, C and E</a>). +The specimen definitely is <i>C. lineatus dunni</i>, and corresponds well +with another specimen from Costa Rica (ANSP 12232).</p> + +<p>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 <i>Conophis nevermanni</i> from other Central American +snakes.</p></div> + +<p><i>Distribution.</i>—Pacific coastal plain of northwestern Costa Rica and the +Meseta Central of central Costa Rica (<a href="#Fig_4">fig. 4</a>).</p> + +<p><i>Specimens examined.</i>—Total of six, as follows: +<span class="smcap">Costa Rica</span>: <i>Guanacaste</i>: +Bebedero, Río Tenorio, NMW 16838(5). "<i>San José</i>," ANSP 22424.</p> + + +<div class="caption3"><a name="Conophis_pulcher" id="Conophis_pulcher"></a> +<b>Conophis pulcher</b> Cope</div> + +<div class="species"><i>Tomodon lineatus</i> (in part), Salvin, Proc. Zool. +Soc. London, 28:455, 1860.</div> + +<div class="species"><i>Conophis pulcher</i> Cope, Proc. Acad. Nat. Sci. Philadelphia, 20(5):308, +1869; Journ. Acad. Nat. Sci. Philadelphia, ser. 2, 8:137, 1876; Bocourt +<i>in</i> 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.</div> + +<div class="species"><i>Tomodon pulcher</i>, Bocourt, Journ. de Zool., p. 408, 1876.</div> + +<div class="species"><i>Conophis pulcher</i> var. <i>similis</i> Bocourt <i>in</i> 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)].</div> + +<div class="species"><i>Conophis lineatus</i>, 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.</div> + +<div class="species"><i>Conophis pulcher pulcher</i>, 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.</div> + +<div class="species"><i>Conophis pulcher plagosus</i> 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.</div> + +<div class="species"><i>Conophis pulcher similis</i>, 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. +<span class="pagenum"><a name="Page_275" id="Page_275">[Pg_275]</a></span> +Kansas Sci. Bull., 33(pt. 2):43-44, March 20, 1950; Maldonado-Koerdell, +Inst. Mexicanos Recursos Nat. Renov. pp. 132-133, 1953.</div> + +<p><i>Types.</i>—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 <i>Conophis +pulcher</i> 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.</p> + +<p><i>Diagnosis.</i>—Paravertebral stripes present at least posteriorly +(<a href="#Fig_1">fig. 1, F</a>); 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.</p> + +<p><i>Variation.</i>—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.</p> + +<p>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.</p> + +<p>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.</p> + +<p>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 +<span class="pagenum"><a name="Page_276" id="Page_276">[Pg_276]</a></span> +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 (<a href="#Fig_1">fig. 1, F</a>). The variation +in continuity and width described above is found in all of the dorsal dark +stripes.</p> + +<p>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.</p> + +<p>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.</p> + +<p>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 (<a href="#Fig_3">fig. 3, B</a>). +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.</p> + +<p>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.</p> + +<p>Three excellent photographs of this species have been published under +the name <i>Conophis lineatus</i> (Ditmars, 1931:pls. 26 and 27).</p> + +<div class="larger"> +<p><i>Remarks.</i>—Smith (1941:121-122) described <i>C. pulcher plagosus</i> +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 <i>p. pulcher</i> or any member of the <i>lineatus</i> 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, +<span class="pagenum"><a name="Page_277" id="Page_277">[Pg_277]</a></span> +the spotting varies from a few inconspicuous spots to many large +spots. Paravertebral rows were continuous posteriorly in alimens +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.</p> + +<p>Mertens (1952a:93, and 1952b:61-62) designated three specimens +from El Salvador as <i>C. pulcher plagosus</i>. In the latter paper, Mertens, +on the basis of a description of a specimen of "<i>C. lineatus</i>" from +Divisadero, El Salvador, given by Schmidt (1928:200), referred that +specimen also to <i>C. pulcher plagosus</i>. I have examined this specimen +and refer it to <i>C. lineatus dunni</i>. 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 <i>C. lineatus dunni</i>.</p> + +<p>The presence of paravertebral stripes in combination with the +characteristics of the genus distinguish <i>Conophis pulcher</i> from all +other snakes in southern México and Central America. The only +sympatric species of this genus, <i>C. lineatus dunni</i>, differs in that it +lacks paravertebral stripes, although it may have a single vertebral +stripe. <i>Conophis lineatus dunni</i> has lateral dark stripes that are +present on the 3rd and 4th scale-rows, never on the anterior third +of the body as in <i>C. pulcher</i>. Even in juveniles the third pair of +dark stripes includes the lower part of the 9th scale-row in <i>C. pulcher</i>, +whereas the dorsal most dark stripe of <i>C. lineatus dunni</i> never +includes more than the lower part of the 8th scale-row.</p> +</div> + +<p><i>Distribution.</i>—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 (<a href="#Fig_4">fig. 4</a>).</p> + +<p><i>Specimens examined.</i>—Total of 27, as follows: +<span class="smcap">Guatemala</span>: <i>no specific +locality</i>, CNHM 22912, NMW 16830. <i>Jutiapa</i>: Hacienda Mongoy, UMMZ +106725. <i>El Progreso</i>: El Progreso, CAS 67000; <i>El Rancho</i>, UMMZ 106724; +<i>San Antonio</i>, CAS 66999. "Peten," USNM 6751(2), 6803. <i>Sacatepequez</i>: +Dueñas, BMNH 64.1.26.17, 64.1.26.126-127. <i>Zacapa</i>: Pepesca, AMNH +72555-56.</p> + +<p><span class="smcap">Honduras</span>: <i>no specific locality</i>, AMNH 58364. <i>Cortes</i>: San Pedro Sula, +CNHM 5295-96. <i>Francisco Morazan: El Zamarano</i>, AMNH 70189; Tegucigalpa, +MCZ 49785, 49787-88, 49791, 49793, 49795.</p> + +<p><span class="smcap">México</span>: <i>Chiapas</i>: <i>Soconusco</i>, UIMNH 33646-47; Tonalá, USNM 109707.</p> + + +<div class="caption3"><a name="Conophis_vittatus" id="Conophis_vittatus"></a> +<b>Conophis vittatus</b> Peters</div> +<br /> + +<div class="species"><i>Tomodon lineatum</i> (in part), Duméril, Bibron and +<ins title='Correction: was "Duméil"'>Duméril</ins>, +<ins title='Correction: was "Érpétologie Genérale"'>Erpétologie Générale</ins>, +7(pt. 2):936-938, February 25, 1854.</div> + +<div class="species"><i>Conophis vittatus</i> Peters, Monatsb. Akad. Wiss. Berlin, pp. 519-520, pl., +<span class="pagenum"><a name="Page_278" id="Page_278">[Pg_278]</a></span> +fig. 3, October, 1860; Cope, Proc. Amer. Philos. Soc., 11:162, 1870; Bocourt +<i>in</i> 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.</div> + +<div class="species"><i>Conophis lineatus</i> Cope, Proc. Acad. Nat. Sci. Philadelphia, 16(3):167, 1864 +[<i>nec</i> Duméril, Bibron and Duméril, +<ins title='Correction: was "Érpétologie Genérale"'>Erpétologie Générale</ins>, +7(pt. 2):936-938, atlas, pl. 73, February 25, 1854; specimen from Colima]; +Sumichrast, Arch. Sci. Nat., p. 246, 1873.</div> + +<div class="species"><i>Tomodon vittatus</i>, Bocourt, Journ. de Zool., p. 407, 1876.</div> + +<div class="species"><i>Conophis sumichrasti sumichrasti</i> 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.</div> + +<div class="species"><i>Conophis sumichrasti viduus</i> 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.</div> + +<div class="species"><i>Conophis sumichrasti</i>, 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.</div> + +<div class="species"><i>Tachymenis lineata</i> (in part), Garman, Mem. Mus. Comp. Zool., 8:60-61, +July, 1884.</div> + +<div class="species"><i>Conophis vittatus sumichrasti</i>, Cope, Ann. Rept. U. S. Natl. Mus. for 1898, +p. 1095, 1900.</div> + +<div class="species"><i>Conophis vittatus videns</i> Cope, Ann. Rept. U. S. Natl. Mus., for 1898, p. +1095, 1900 (apparent <i>lapus</i> for <i>viduus</i>).</div> + +<div class="species"><i>Conophis vittatus vittatus</i>, 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.</div> + +<div class="species"><i>Conophis vittata</i>, Gadow, Proc. Zool. Soc. London, 2:196, 1905; Through +Southern Mexico, p. 181, 1908.</div> + +<div class="species"><i>Conophis viduus</i>, 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.</div> + +<p><span class="pagenum"><a name="Page_279" id="Page_279">[Pg_279]</a></span></p> + +<div class="species"><i>Conophis vittatus viduus</i>, 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.</div> + +<div class="species"><i>Conophis vittatus vittatus</i> × <i>Conophis vittatus viduus</i>, Alvarez del Toro +and Smith, Herpetologica, 12:13, March 6, 1956.</div> + +<p><i>Type.</i>—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).</p> + +<p><i>Diagnosis.</i>—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.</p> + +<p><a name="Variation" id="Variation"></a><i>Variation.</i>—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 <ins title='Correction: was "840"'>850</ins> mm. A juvenile (CNHM 40435) from +<ins title='Correction: was "Tehauntepec"'>Tehuantepec</ins>, +Oaxaca, México, has a body length of 133 mm., a tail length of 31 mm. and +a total length of 164 mm.</p> + +<p>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 <i>C. +vittatus</i> 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.</p> + +<p>Snakes from areas between Tehuantepec and the margins of the distribution +<span class="pagenum"><a name="Page_280" id="Page_280">[Pg_280]</a></span> +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.</p> + +<p>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.</p> + +<p>There is no ontogenetic change in color pattern; juveniles have the same +coloration as adults from the same geographic area.</p> + +<p>Color in life is not greatly different from that of preserved specimens. One +specimen (UMMZ 114483) from 10.8 miles south of +<ins title='Correction: "Oaxaca," repeated'>Oaxaca,</ins> had in +life black stripes, a pale yellowish tan dorsal ground-color and a pale off-white +venter.</p> + +<p>An excellent photograph of this species appears in Schmidt and Inger +(1957:230) under the name <i>Conophis lineatus</i>.</p> + +<div class="larger"> +<p><i>Remarks.</i>—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; <i>C. vittatus vittatus</i> +having four black stripes and <i>C. vittatus viduus</i> 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.</p> + +<p><span class="pagenum"><a name="Page_281" id="Page_281">[Pg_281]</a></span> +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 +<i>Conophis vittatus</i> from all other Méxican snakes. The +only other snake that occurs in western México that has been +confused with <i>C. vittatus</i> is <i>Coniophanes piceivittus taylori</i>, which +has 25, instead of 19, scale-rows.</p></div> + +<p><i>Distribution.</i>—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 (<a href="#Fig_5">fig. 5</a>).</p> + +<div class="fig_center" style="width: 510px;"> +<a name="Fig_5" id="Fig_5"></a> +<img src="images/fig_5.png" width="510" height="357" alt="" title="" /> +<div class="fig_caption"> +<span class="smcap">Fig. 5.</span> Selected locality records for +<i>Conophis vittatus</i>.</div> +</div> +<br /> + +<p><i>Specimens examined.</i>—Total of 174, as follows: <span class="smcap">México</span>: <i>no specific locality</i>, +AMNH 66150-52, SU 9465. <i>Chiapas</i>: Piedra Parada, USNM 121453. +<i>Pizo de Oro</i>, UIMNH 40821. Tuxtla Gutierrez, Parque Madero, UIMNH +37992-93, 38036-37. <i>Colima: no specific locality</i>, MCZ 46860, USNM 31394, +31396-97. 1 mi. SW Colima, AMNH 12783. S of Manzanillo, AMNH 19641. +<i>Durango</i>: Hacienda de Gabriel, AMNH 14217. <i>Guerrero: Acahuizotla</i>, TCWC +7419, 9469. <i>1 mi. W Acahuizotla</i>, TCWC 7418. 3 mi. W Acapulco, AMNH +71626. <i>6 mi. E Acapulco</i>, TCWC 9476-77. <i>10 mi. S Acapulco</i>, TCWC 8578. +<i>Agua del Obispo</i>, CNHM 104948, TCWC 11586. near Chilpancingo, MVZ +45067, UMMZ 85722-23. <i>1 mi. SW Colotlipa</i>, TCWC 9471-74. <i>2 mi. SW +Colotlipa</i>, TCWC 9475. 14 mi. S Ixtapán de la Sal, KU 67648. <i>Laguna +Coyuca</i>, CNHM 25881, UMMZ 80942. near La Unión, AMNH 66337. +<i>Magueyes, Laguna Coyuca</i>, AMNH 66149. <i>Playa Encantada</i>, TCWC 9470. +1 mi. S Tierra Colorada, KU 67649. near <i>Xaltinanguis, km. 405</i>, CNHM +104947. <i>Michoacán</i>: Coalcomán, UMMZ 104693. <i>1/2 mi. SE Coalcomán</i>, +UMMZ 104492. <i>1 mi. N. Coalcomán</i>, UMMZ 112543. <i>1 mi. NE Coalcomán</i>, +UMMZ 104692. Puerta de la Playa, UMMZ 105155. 12 mi. S +<span class="pagenum"><a name="Page_282" id="Page_282">[Pg_282]</a></span> +Tzitzio (by road), UMMZ 99153. <i>Morelos: 12 km. NW Axochiapan</i>, TCWC +7311, UIMNH 17613, 25924. 7 mi. SE Cuernavaca, MVZ 32258. <i>Huajintlán, +km. 133</i>, CNHM 103270. 12 km. S Puente de Ixtla, km. 133, CNHM 104949. +<i>Oaxaca: Bisiliana</i>, AMNH 68010. <i>near Caoba, foot of Cerro Arenal</i>, AMNH +68009. <i>Cerro Arenal</i>, AMNH 68000-03. <i>Cerro de Laollaga</i>, UIMNH 36213. +<i>Cerro de San Pedro</i>, UIMNH 17616. <i>Cerro Palma de Oro</i>, UIMNH 37116. +"<i>C. Madrena, Sto. T. Quieri</i>," UIMNH 46904. near Chivela, MCZ 25021. +Cinco Cerros, UIMNH 37114. <i>Dami Liesa</i>, AMNH 66877, UIMNH 6158, +37115. <i>Escuranos</i>, AMNH 66873-74, 68004-06. <i>Finca Santa Teresa, 2 km. +NW Tehuantepec</i>, UMMZ 82648. <i>Huilotepec</i>, AMNH 66878, UIMNH +40820. <i>between Huilotepec and Tehuantepec</i>, AMNH 65106, UMMZ 82644-45. +<i>Las Tejas</i>, UIMNH 6151-54. <i>Mixtequilla</i>, UIMNH 6157, 36211. <i>between +Mixtequilla Mountains and Tehuantepec</i>, UMMZ 82652. <i>between Niltepec +and "Carixxal,"</i> AMNH 68876. 10.8 mi. SE Oaxaca, UMMZ 114483. <i>Quiengola</i>, +UIMNH 17617. <i>between Quiengola Mountains and Tehuantepec</i>, UMMZ +82647. <i>Rancho Poso Río, 6 km. S Tehuantepec</i>, UIMNH 6144-49, 37117-19, +UMMZ 82649-51. <i>Rincón Bamba</i>, CNHM 105129-30, UIMNH 17615. <i>Salazar</i>, +AMNH 66875. <i>vicinity of Salina Cruz</i>, UMMZ 82653. <i>San Gerónimo</i>, +AMNH 4306, CNHM 1457. <i>San Lucas Ixtepec</i>, UIMNH 36206. San Juan +Lajarcia, UIMNH 36212. San Mateo del Mar, AMNH 65914. <i>San Pablo</i>, +UIMNH 36207. <i>Santa María (Cerro de Liesa)</i>, 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, <i>1-2 leagues SSE Tehuantepec</i>, UMMZ 82639-41. Tenango, +UIMNH 36209-10. <i>between Tlacolulita and Tequisistlán</i>, CNHM 105125. +<i>Yerba Santa</i>, UIMNH 6155-56. Puebla: Atencingo, KU 39626.</p> + + +<div class="caption3"><a name="Skull" id="Skull"></a> +Skull</div> + +<p>In studying the osteology of the genus <i>Conophis</i>, I have examined +two complete skeletons (one <i>C. vittatus</i> and one <i>C. lineatus</i>); two +additional skulls of <i>C. vittatus</i> and <i>C. lineatus</i>; 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 <i>Tomodon lineatus</i> 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.</p> + +<p>The skull of <i>Conophis</i> is typical of a relatively unspecialized +colubrid snake. Skulls of <i>Conophis lineatus concolor</i> and <i>C. vittatus</i> +closely resemble each other. The following description is based +primarily on the skull of <i>C. lineatus concolor</i> (UMMZ S-778).</p> + +<p>The elements are discussed in the following order: nasal region, +cranium and associated elements, maxillo-palatal-pterygoid arch, +mandible, dentition, and vertebrae.</p> + +<p><i>Nasal region.</i>—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 (<a href="#Fig_6">fig. 6</a>) 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 +<span class="pagenum"><a name="Page_283" id="Page_283">[Pg_283]</a></span> +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.</p> + +<div class="fig_center" style="width: 474px;"> +<a name="Fig_6" id="Fig_6"></a> +<img src="images/fig_6.png" width="474" height="625" alt="" title="" /> +<div class="fig_caption"> +<span class="smcap">Fig. 6.</span> The skull, lacking dentigerous bones, +of <i>Conophis lineatus concolor</i> (UMMZ S-788) showing (A) dorsal, +(B) lateral, and (C) ventral views. × 3.</div> +</div> +<br /> + +<p><span class="pagenum"><a name="Page_284" id="Page_284">[Pg_284]</a></span> +<i>Cranium and associated elements.</i>—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 +<span class="pagenum"><a name="Page_285" id="Page_285">[Pg_285]</a></span> +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 +<span class="pagenum"><a name="Page_286" id="Page_286">[Pg_286]</a></span> +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.</p> + +<div class="fig_center" style="width: 420px;"> +<a name="Fig_7" id="Fig_7"></a> +<img src="images/fig_7.png" width="420" height="466" alt="" title="" /> +<div class="fig_caption"> +<span class="smcap">Fig. 7.</span> The maxillo-palatal-pterygoid arch of +<i>Conophis lineatus concolor</i> (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.</div> +</div> +<br /> + +<p><i>Maxillo-palatal-pterygoid arch.</i>—In an articulated skull, the anterior edge +of the maxillary is immediately posterior to the lateral tip of the premaxillary +(<a href="#Fig_7">fig. 7</a>). 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.</p> + +<div class="fig_center" style="width: 429px;"> +<a name="Fig_8" id="Fig_8"></a> +<img src="images/fig_8.png" width="429" height="418" alt="" title="" /> +<div class="fig_caption"> +<span class="smcap">Fig. 8.</span> The left mandible and associated quadrate +of <i>Conophis lineatus concolor</i> (UMMZ S-788) showing (A) lateral and +(B) medial views. × 3. Teeth shown by means of broken lines +were represented only by their sockets.</div> +</div> +<br /> + +<p><i>Mandible.</i>—The dentary (<a href="#Fig_8">fig. 8</a>) 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. +<span class="pagenum"><a name="Page_287" id="Page_287">[Pg_287]</a></span> +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 +<span class="pagenum"><a name="Page_288" id="Page_288">[Pg_288]</a></span> +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.</p> + + +<div class="caption3"><a name="Dentition" id="Dentition"></a> +Dentition</div> + +<p>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 <i>Conophis</i> 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 <i>Conophis</i>.</p> + + +<div class="caption3"><a name="Vertebrae" id="Vertebrae"></a> +Vertebrae</div> + +<p>The fiftieth vertebra of <i>Conophis vittatus</i> (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 +<span class="pagenum"><a name="Page_289" id="Page_289">[Pg_289]</a></span> +of the middle thoracic vertebra of <i>Conophis lineatus dunni</i> from Honduras +were published by Auffenberg (1958:6).</p> + + +<div class="caption3"><a name="Hemipenes" id="Hemipenes"></a> +Hemipenes</div> + +<p>The hemipenes of <i>Conophis</i> are moderately caliculate, having spines +covering the surface from the base to near the apex (<a href="#Fig_9">fig. 9</a>). 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 <i>C. vittatus</i> and one lamina in <i>C. +lineatus concolor</i>. 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 <i>C. vittatus</i>) or a deeply bilobed appearance in others +(especially <i>C. lineatus concolor</i>).</p> + +<div class="fig_center" style="width: 242px;"> +<a name="Fig_9" id="Fig_9"></a> +<img src="images/fig_9.png" width="242" height="389" alt="" title="" /> +<div class="fig_caption"> +<span class="smcap">Fig. 9.</span> The everted left hemipenis of +Conophis vittatus (UMMZ 82650). × 5.</div> +</div> +<br /> + +<p>The everted hemipenis reaches +posteriorly to the eighth subcaudal +scale. The sulcus bifurcates at the +third subcaudal scale. The situation +is similar <i>in situ</i> (Cope, 1895:pl. 28, +fig. 2).</p> + +<p>There are no apparent hemipenial +differences among the species of the +genus <i>Conophis</i>. As can be seen in +the above description, the hemipenis +of <i>C. vittatus</i> is less bilobed and has a +more pronounced disc at the apex than +the others. The hemipenis of <i>C. lineatus concolor</i> is most bilobed, but has +the smallest apical disc. The other species and subspecies vary widely within +these extremes.</p> + + +<div class="caption3"><a name="Food_and_Feeding" id="Food_and_Feeding"></a> +Food and Feeding</div> + +<p><i>Conophis</i> eats mostly small lizards, especially <i>Cnemidophorus</i>. +In México <i>Conophis</i> occurs in semi-arid habitat where <i>Cnemidophorus</i> +is common. A specimen each of <i>Conophis vittatus</i> and +<i>C. lineatus lineatus</i> were obtained while I was collecting <i>Cnemidophorus</i>. +The only record of <i>Conophis</i> having fed on a warm-blooded +vertebrate was obtained in the course of this study, when +I recovered from the stomach of a <i>Conophis lineatus concolor</i> +(CNHM 36299) from Chichén Itzá, Yucatán, a heteromyid rodent +(<i>Heteromys gaumeri</i>).</p> + +<p><span class="pagenum"><a name="Page_290" id="Page_290">[Pg_290]</a></span> +Ralph Axtell (personal communication) observed <i>Conophis</i> actively +searching for food at dusk. His observations were made near +Tehuantepec, Oaxaca, and the snakes were seen to chase lizards +of the genus <i>Cnemidophorus</i>. Near Alvarado, Veracruz, in the late +afternoon, I watched a <i>Conophis lineatus lineatus</i> follow a lizard +into a hole.</p> + +<p>Mittleman (1944:122) presents the only discussion of the mode +of feeding of a captive specimen of <i>Conophis lineatus</i> ssp. When +presented with a <i>Thamnophis</i> slightly smaller than itself, the +<i>Conophis</i> struck, and within eight minutes immobilized the +<i>Thamnophis</i>. Within one-half hour the <i>Thamnophis</i> was swallowed. +Three days later the <i>Conophis</i> ate another <i>Thamnophis</i>, +though still distended from its first meal; nine days later it ate a +<i>Storeria</i>. In the course of several months, the <i>Conophis</i> ate various +toads and hylids and two more <i>Storeria</i>. Apparently members of +the genus <i>Conophis</i> do not constrict their prey, but rely upon a +combination of loss of blood and action of the venom to completely +immobilize their prey.</p> + +<p>Ditmars (1931:pls. 26-27) showed three photographs of "<i>Conophis +lineatus</i>" (actually <i>Conophis pulcher</i>) ingesting another snake, +identified by him as a young <i>Ophis (= Xenodon) colubrinus</i>.</p> + + +<div class="caption3"><a name="Effect_of_Poison" id="Effect_of_Poison"></a> +Effect of Poison</div> + +<p>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 <i>C. l. lineatus</i> +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 <i>Conophis +vittatus</i> (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, +<span class="pagenum"><a name="Page_291" id="Page_291">[Pg_291]</a></span> +1956. There was immediate pain and localized swelling, both of +which disappeared several hours later.</p> + + +<div class="caption2"> +<a name="TAXONOMIC_RELATIONSHIPS_AND_EVOLUTION" id="TAXONOMIC_RELATIONSHIPS_AND_EVOLUTION"></a> +TAXONOMIC RELATIONSHIPS AND EVOLUTION</div> + +<p>The genus <i>Conophis</i> is known only from the Recent. Except +that <i>Conophis</i> 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 +<i>Dryinoides oxyrhachis</i> 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 +<i>Conophis lineatus dunni</i> (UF 7657) from Honduras, with which it +was compared in basic osteology. The two genera could be related, +for the progenitors of <i>Conophis</i> possibly inhabited much of North +America in the Miocene.</p> + +<p>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 <i>Conophis</i> stock isolated in Middle +America where members of the genus dispersed through semi-arid +habitats.</p> + +<p>Turning our attention now to the species within the genus, +instead of the genus as a whole, <i>Conophis vittatus</i> 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 <i>C. lineatus</i>-group. +<i>Conophis nevermanni</i> 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 <i>nevermanni</i> is more closely related +to the <i>lineatus</i>-group than to <i>vittatus</i>. The <i>lineatus</i>-group, thus, +consists of <i>pulcher</i>, <i>nevermanni</i> and the three subspecies of <i>lineatus</i>. +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. <i>Conophis lineatus concolor</i>, on +which the dark pigmentation on the body apparently is secondarily +lost, is an exception.</p> + +<p><span class="pagenum"><a name="Page_292" id="Page_292">[Pg_292]</a></span> +If differences in color pattern be used as an indication of the +relationships between the species and subspecies of the genus +<i>Conophis</i>, I would consider <i>C. vittatus</i> the most divergent unit. The +subspecies of <i>lineatus</i> closely resemble one another and, as a unit, +resemble <i>pulcher</i> from which they differ +<ins title='Correction: was "primarly"'>primarily</ins> in the position +of the dorsalmost stripes. <i>Conophis nevermanni</i> is more divergent +than is <i>pulcher</i> from the species <i>lineatus</i>, but probably is not so far +removed from <i>lineatus</i> as is <i>vittatus</i>.</p> + +<p>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 <i>vittatus</i>, +and has subsequently spread north and west from the region of +Tehuantepec, México. During the same period <i>nevermanni</i> became +isolated in northern Costa Rica.</p> + +<p>The species <i>pulcher</i> probably differentiated from the remaining +<i>lineatus</i> stock during the Early Pleistocene orogenic upheaval in +Guatemala. The <i>pulcher</i> stock was isolated on the Pacific Coastal +slopes of Guatemala, while <i>lineatus</i> moved through the subhumid +corridor of northern Middle America into México and southward +toward Costa Rica (Stuart, 1954a). In the Late Pleistocene and +Recent, <i>pulcher</i> 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 <i>lineatus</i>, this species differentiated into three subspecies.</p> +<br /> + +<div class="caption2"><a name="SUMMARY" id="SUMMARY"></a> +SUMMARY</div> + +<p>The genus <i>Conophis</i> Peters, 1860, contains four species. Three +are monotypic and the fourth has three subspecies, making a total +of six taxa.</p> + +<p>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.</p> + +<p>The six taxa are separated primarily on the basis of color pattern, +but characters of scutellation, including numbers of dorsals, ventrals, +<span class="pagenum"><a name="Page_293" id="Page_293">[Pg_293]</a></span> +caudals, and places of reduction of the number of dorsal +scale-rows, were analyzed.</p> + +<p>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 <i>Cnemidophorus</i>; in addition +they are known to eat small rodents and other snakes.</p> + +<p><i>Conophis</i> is a member of the subfamily Xenodontinae and, as +presently understood, has no known living close relatives. A single +specimen of <i>Dryinoides</i> from the Miocene of Montana has been +compared with this genus. The genus <i>Conophis</i> 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 <i>Conophis</i> presently lives.</p> +<br /> +<br /> + + +<div class="caption2"><a name="LITERATURE_CITED" id="LITERATURE_CITED"></a> +LITERATURE CITED</div> +<br /> + +<div class="smcap">Auffenberg, W.</div> + +<div class="reference">1958. A new genus of colubrid snake from the Upper Miocene of North +America. Amer. Mus. Novitates, 1874:1-16. February 27.</div> +<br /> + +<div class="smcap">Cope, E. D.</div> + +<div class="reference">1861. Contributions to the ophiology of Lower California, México and +Central America. Proc. Acad. Nat. Sci. Philadelphia, 13:292-306. +December 28.</div> + +<div class="reference">1867. Fifth contribution to the herpetology of tropical America. Proc. +Acad. Nat. Sci. Philadelphia, 18:317-323. February 20.</div> + +<div class="reference">1871. Ninth contribution to the herpetology of tropical America. Proc. +Acad. Nat. Sci. Philadelphia, 23(2):200-224. October 24.</div> + +<div class="reference">1876. On the batrachia and reptilia of Costa Rica. Journ. Acad. Nat. +Sci. Philadelphia, series 2, 8(4):93-154, 6 pls.</div> + +<div class="reference">1895. The classification of the ophidia. Trans. Amer. Philos. Soc., +18:186-219, 33 pls. April 15.</div> + +<div class="reference">1900. The crocodilians, lizards, and snakes of North America. Ann. Rept. +U. S. Natl. Mus. for 1898, pp. 153-1270, 36 pls.</div> +<br /> + +<div class="smcap">Ditmars, R. L.</div> + +<div class="reference">1931. Snakes of the World. New York, The MacMillan Company, 1931. +xi + 207 pp., 84 pls.</div> +<br /> + +<div class="smcap">Dowling, H. G.</div> + +<div class="reference">1951. A proposed standard system of counting ventrals in snakes. British +Journ. Herpetology, 1(5):97-99, fig. 1.</div> +<br /> + +<div class="smcap">Duellman, W. E.</div> + +<div class="reference">1958. A preliminary analysis of the herpetofauna of Colima, Mexico. +Occas. Papers Mus. Zool. Univ. Michigan, 589:1-22, March 21.</div> +<br /> + +<div class="smcap">Duméril, A. M. C., Bibron, G., and Duméril, A. H. A.</div> + +<div class="reference">1854. Érpétologie genérale, ou histoire naturelle des reptiles. Paris, 7(pt. +2):xii + 785. February 25. Atlas, 24 pp., 108 pls.</div> +<br /> + +<div class="smcap">Duméril, A. H. A., Bocourt, M., and Mocquard, F.</div> + +<div class="reference">1870-1909. Mission Scientifique au Mexique et dans l'Amerique Centrale +… Etudes sur les Reptiles. Paris, vol. 2:xiv + 1012 pp., +77 pls.</div> +<br /> + +<div class="smcap">Garman, S.</div> + +<div class="reference">1884a. The North American reptiles and batrachians. Bull. Essex Inst., +16:1-46. January 9.</div> + +<p><span class="pagenum"><a name="Page_294" id="Page_294">[Pg_294]</a></span></p> + +<div class="reference">1884b. The reptiles and batrachians of North America. Mem. Mus. +Comp. Zool., 8(3):xxxi + 185 pp., 9 pls. July.</div> +<br /> + +<div class="smcap">Günther, A. C. L. G.</div> + +<div class="reference">1858. Catalogue of colubrine snakes in the collection of the British +Museum. London. xiv + 281 pp.</div> +<br /> + +<div class="smcap">Huxley, J.</div> + +<div class="reference">1942. Evolution. The Modern Synthesis. London. 645 pp.</div> +<br /> + +<div class="smcap">Jan, G. and Sordelli, F.</div> + +<div class="reference">1866. Iconographie Generale des Ophidiens. Milano. livr. 19, pls. 1-6. +December.</div> + +<div class="reference">1881. Iconographie Generale des Ophidiens. Milano. livr. 50, pls. 1-7. +November.</div> +<br /> + +<div class="smcap">Mayr, E.</div> + +<div class="reference">1942. Systematics and the Origin of Species. New York, x + 334 pp., +29 figs.</div> +<br /> + +<div class="smcap">Mayr, E., Linsley, E. G., and Usinger, R. L.</div> + +<div class="reference">1953. Methods and Principles of Systematic Zoology. New York. +ix + 328 pp., 45 figs.</div> +<br /> + +<div class="smcap">Mertens, R.</div> + +<div class="reference">1952a. Neues uber die Reptilienfauna von El Salvador. Zool. Anz., +148:87-93. February.</div> + +<div class="reference">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.</div> +<br /> + +<div class="smcap">Mittleman, M. B.</div> + +<div class="reference">1944. Feeding habits of a Central American opisthoglyph snake. Copeia, +no. 2:122. June 30.</div> +<br /> + +<div class="smcap">Neill, W. T. and Allen, R.</div> + +<div class="reference">1961. Further studies on the herpetology of British Honduras. Herpetologica, +17(1):37-52. April 15.</div> +<br /> + +<div class="smcap">Parker, W. K.</div> + +<div class="reference">1878. On the structure and development of the skull in the common +snake (<i>Tropidonotus natrix</i>). Phil. Trans. Roy. Soc. London, pt. 2:385-417, +pp., pls. 27-33.</div> +<br /> + +<div class="smcap">Peters, W.</div> + +<div class="reference">1860. Drei neue amerikanisches Schlangen. Monatsb. Akad. Wiss. Berlin, +1860:517-521, pl., fig. 3. October.</div> +<br /> + +<div class="smcap">Radovanovic, M.</div> + +<div class="reference">1937. Osteologie des Schlangenkopfs. Jenaische Zeitschr. Naturw., +71(2):179-312.</div> +<br /> + +<div class="smcap">Savage, J. M.</div> + +<div class="reference">1949. Notes on the Central American snake, <i>Conophis lineatus dunni</i> +Smith, with a record from Honduras. Trans. Kansas Acad. Sci., +50:483-486. December 31.</div> +<br /> + +<div class="smcap">Schmidt, K. P.</div> + +<div class="reference">1928. Reptiles collected in Salvador for the California Institute of Technology. +Zool. Ser. Field Mus. Nat. Hist., 12(16):193-201. November 21.</div> +<br /> + +<div class="smcap">Schmidt, K. P. and Inger, R. F.</div> + +<div class="reference">1957. Living Reptiles of the World. Garden City, New York, Hanover +House. 287 pp.</div> +<br /> + +<div class="smcap">Smith, H. M.</div> + +<div class="reference">1941. Notes on snakes of the genus <i>Conophis</i>. Journ. Washington Acad. +Sci., 31(3):117-124. March 15.</div> +<br /> + +<div class="smcap">Smith, H. M. and Taylor, E. H.</div> + +<div class="reference">1950. Type localities of Méxican reptiles and amphibians. Univ. Kansas +Sci. Bull., 33:313-380. March 20.</div> +<br /> + +<p><span class="pagenum"><a name="Page_295" id="Page_295">[Pg_295]</a></span></p> + +<div class="smcap">Stuart, L. C.</div> + +<div class="reference">1948. The amphibians and reptiles of Alta Verapaz, Guatemala. Misc. +Publ. Mus. Zool. Univ. Michigan, 69:1-109. June 12.</div> + +<div class="reference">1954a. A description of a subhumid corridor across northern Central +America, with comments on its <ins title='Correction: was "hertetofaunal"'>herpetofaunal</ins> indicators. Contr. +Lab. Vert. Biol. Univ. Michigan, 65:1-26 pp., 6 pls. March.</div> + +<div class="reference">1954b. Herpetofauna of the southeast highlands of Guatemala. Contr. +Lab. Vert. Biol. Univ. Michigan, 68:1-65 pp., 3 pls. November.</div> +<br /> + +<div class="smcap">Szunyoghy, J.</div> + +<div class="reference">1932. Beitrage zur vergleichenden Formenlehre des Colubridenschadels, +nebst einer Kraniologischen Synopsis der fossilen Schlangen Ungarns. +Acta Zool., 13:1-56.</div> +<br /> + +<div class="smcap">Taylor, E. H.</div> + +<div class="reference">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.</div> +<br /> + +<div class="smcap">Taylor, E. H. and Smith, H. M.</div> + +<div class="reference">1939. Miscellaneous notes on Mexican snakes. Univ. Kansas Sci. Bull., +25:239-258. July 10.</div> +<br /> + +<div class="smcap">Wettstein, O.</div> + +<div class="reference">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.</div> +<br /> + +<p><i>Transmitted November 30, 1962.</i></p> +<br /> +<br /> +<br /> +<br /> + +<div class="center"> +<img src="images/square.png" width="16" height="17" alt="" title="" /><br /> +29-5936 +</div> + +<br /> +<br /> +<br /> + +<p><span class="pagenum"><a name="Page_i" id="Page_i">[Pg_i]</a></span></p> + +<div class="caption2"><a name="PUBLICATIONS" id="PUBLICATIONS"></a> +UNIVERSITY OF KANSAS PUBLICATIONS</div> + +<div class="caption2">MUSEUM OF NATURAL HISTORY</div> + +<p>Institutional libraries interested in publications exchange may obtain this +series by addressing the Exchange Librarian, University of Kansas Library, +Lawrence, Kansas. Copies for individuals, persons working in a particular +field of study, may be obtained by addressing instead the Museum of Natural +History, University of Kansas, Lawrence, Kansas. There is no provision for +sale of this series by the University Library, which meets institutional requests, +or by the Museum of Natural History, which meets the requests of individuals. +However, when individuals request copies from the Museum, 25 cents should +be included, for each separate number that is 100 pages or more in length, for +the purpose of defraying the costs of wrapping and mailing.</p> + +<p>* An asterisk designates those numbers of which the Museum's supply (not the Library's +supply) is exhausted. Numbers published to date, in this series, are as follows:</p> + +<table width="100%" summary="publications"> +<tr> + <td class="text_rt vtop">Vol. 1.</td> + <td colspan="2">Nos. 1-26 and index. Pp. 1-638, 1946-1950.</td> +</tr> +<tr> + <td class="text_rt vtop">*Vol. 2.</td> + <td colspan="2">(Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140 + figures in text. April 9, 1948.</td> +</tr> +<tr> + <td class="text_rt vtop">Vol. 3.</td> + <td class="text_rt vtop">*1.</td> + <td>The avifauna of Micronesia, its origin, evolution, and distribution. By Rollin + H. Baker. Pp. 1-359, 16 figures in text. June 12, 1951.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">*2.</td> + <td>A quantitative study of the nocturnal migration of birds. By George H. + Lowery, Jr. Pp. 361-472, 47 figures in text. June 29, 1951.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">3.</td> + <td>Phylogeny of the waxwings and allied birds. By M. Dale Arvey. Pp. 473-530, + 530, 49 figures in text, 13 tables. October 10. 1951.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">*4.</td> + <td>Birds from the state of Veracruz, Mexico. By George H. Lowery, Jr., and + Walter W. Dalquest. Pp. 531-649, 7 figures in text, 2 tables. October 10, + 1951.</td> +</tr> +<tr> + <td> </td> + <td colspan="2">Index. Pp. 651-681.</td> +</tr> +<tr> + <td class="text_rt vtop">*Vol. 4.</td> + <td colspan="2">(Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, 31 + figures in text. December 27, 1951.</td> +</tr> +<tr> + <td class="text_rt vtop">Vol. 5.</td> + <td colspan="2">Nos. 1-37 and index. Pp. 1-676, 1951-1953.</td> +</tr> +<tr> + <td class="text_rt vtop">*Vol. 6.</td> + <td colspan="2">(Complete) Mammals of Utah, taxonomy and distribution. By Stephen D. + Durrant. Pp. 1-549, 91 figures in text, 30 tables. August 10, 1952.</td> +</tr> +<tr> + <td class="text_rt vtop">Vol. 7.</td> + <td colspan="2">Nos. 1-15 and index. Pp. 1-651, 1952-1955.</td> +</tr> +<tr> + <td class="text_rt vtop">Vol. 8.</td> + <td colspan="2">Nos. 1-10 and index. Pp. 1-675, 1954-1956.</td> +</tr> +<tr> + <td class="text_rt vtop">Vol. 9.</td> + <td class="text_rt vtop">*1.</td> + <td>Speciation of the wandering shrew. By James S. Findley. Pp. 1-68, 18 + figures in text. December 10, 1955.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">2.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">3.</td> + <td>A new long-eared myotis (Myotis evotis) from northeastern Mexico. By Rollin + H. Baker and Howard J. Stains. Pp. 81-84. December 10, 1955.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">4.</td> + <td>Subspeciation in the meadow mouse, Microtus pennsylvanicus, in Wyoming. + By Sydney Anderson. Pp. 85-104, 2 figures in text. May 10, 1956.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">5.</td> + <td>The condylarth genus Ellipsodon. By Robert W. Wilson. Pp. 105-116, 6 + figures in text. May 19, 1956.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">6.</td> + <td>Additional remains of the multituberculate genus Eucosmodon. By Robert + W. Wilson. Pp. 117-123, 10 figures in text. May 19, 1956.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">7.</td> + <td>Mammals of Coahulia, Mexico. By Rollin H. Baker. Pp. 125-335, 75 figures + in text. June 15, 1956.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">8.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">9.</td> + <td>Extensions of known ranges of Mexican bats. By Sydney Anderson. Pp. + 347-351. August 15, 1956.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">10.</td> + <td>A new bat (Genus Leptonycteris) from Coahulia. By Howard J. Stains. + Pp. 353-356. January 21, 1957.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">11.</td> + <td>A new species of pocket gopher (Genus Pappogeomys) from Jalisco, Mexico. + By Robert J. Russell. Pp. 357-361. January 21, 1957.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">12.</td> + <td>Geographic variation in the pocket gopher, Thomomys bottae, in Colorado. + By Phillip M. Youngman. Pp. 363-<ins title='Correction: was "387"'>384</ins>, + 7 figures in text. February 21, 1958.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">13.</td> + <td>New bog lemming (genus Synaptomys) from Nebraska. By J. Knox Jones, + Jr. Pp. 385-388. May 12, 1958.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">14.</td> + <td>Pleistocene bats from San Josecito Cave, Nuevo León, México. By J. Knox + Jones, Jr. Pp. 389-396. December 19, 1958.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">15.</td> + <td>New subspecies of the rodent Baiomys from Central America. By Robert + L. Packard. Pp. 397-404. December 19, 1958.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">16.</td> + <td>Mammals of the Grand Mesa, Colorado. By Sydney Anderson. Pp. 405-414, + 1 figure in text, May 20, 1959.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">17.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">18.</td> + <td>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. + <span class="pagenum"><a name="Page_ii" id="Page_ii">[Pg_ii]</a></span></td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">19.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">20.</td> + <td>Small carnivores from San Josecito Cave (Pleistocene), Nuevo León, México. + By E. Raymond Hall. Pp. 531-538, 1 figure in text. January 14, 1960.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">21.</td> + <td>Pleistocene pocket gophers from San Josecito Cave, Nuevo León, México. + By Robert J. Russell. Pp. 539-548, 1 figure in text. January 14, 1960.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">22.</td> + <td>Review of the insectivores of Korea. By J. Knox Jones, Jr., and David H. + Johnson. Pp. 549-578. February 23, 1960.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">23.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td colspan="2">Index. Pp. 671-690</td> +</tr> +<tr> + <td class="text_rt vtop">Vol. 10.</td> + <td class="text_rt vtop">1.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">2.</td> + <td>Comparative breeding behavior of Ammospiza caudacuta and A. maritima. + By Glen E. Woolfenden. Pp. 45-75, 6 plates, 1 figure. December 20, 1956.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">3.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">4.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">5.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">*6.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">7.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">8.</td> + <td>Natural history of the salamander, Aneides hardyi. By Richard F. Johnston + and Gerhard A. Schad. Pp. 573-585. October 8, 1959.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">9.</td> + <td>A new subspecies of lizard, Cnemidophorus sacki, from Michoacán, México. + By William E. Duellman. Pp. 587-598, 2 figures in text. May 2, 1960.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">10.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td colspan="2">Index. Pp. 611-626.</td> +</tr> +<tr> + <td class="text_rt vtop">Vol. 11.</td> + <td colspan="2">Nos. 1-10 and index. Pp. 1-703, 1958-1960.</td> +</tr> +<tr> + <td class="text_rt vtop">Vol. 12.</td> + <td class="text_rt vtop">1.</td> + <td>Functional morphology of three bats: Sumops, Myotis, Macrotus. By Terry + A. Vaughan. Pp. 1-153, 4 plates, 24 figures in text. July 8, 1959.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">*2.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">3.</td> + <td>The baculum in microtine rodents. By Sydney Anderson. Pp. 181-216, 49 + figures in text. February 19, 1960.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">*4.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">5.</td> + <td>Natural history of the bell vireo. By Jon C. Barlow. Pp. 241-296, 6 figures + in text. March 7, 1962.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">6.</td> + <td>Two new pelycosaurs from the lower Permian of Oklahoma. By Richard C. + Fox. Pp. 297-307, 6 figures in text. May 21, 1962.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">7.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">8.</td> + <td>Teeth of Edestid sharks. By Theodore H. Eaton, Jr. Pp. 347-362, 10 figures + in text. October 1, 1962.</td> +</tr> +<tr> + <td> </td> + <td colspan="2">More numbers will appear in volume 12.</td> +</tr> +<tr> + <td class="text_rt vtop">Vol. 13.</td> + <td class="text_rt vtop">1.</td> + <td>Five natural hybrid combinations in minnows (Cyprinidae). By Frank B. + Cross and W. L. Minckley. Pp. 1-18. June 1, 1960.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">2.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">3.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">4.</td> + <td>Autecology of the copperhead. By Henry S. Fitch. Pp. 85-288, pls. 13-20, + 26 figures in text. November 30, 1960.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">5.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">6.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">7.</td> + <td>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. + <span class="pagenum"><a name="Page_iii" id="Page_iii">[Pg_iii]</a></span></td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">8.</td> + <td><ins title='Correction: was "Decriptions"'>Descriptions</ins> + 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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">9.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">10.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td colspan="2">Index. Pp. 613-624.</td> +</tr> +<tr> + <td class="text_rt vtop">Vol. 14.</td> + <td class="text_rt vtop">1.</td> + <td>Neotropical bats from western México. By Sydney Anderson. Pp. 1-8. + October 24, 1960.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">2.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">3.</td> + <td>Mammals of Mesa Verde National Park, Colorado. By Sydney Anderson. + Pp. 29-67, pls. 1 and 2, 3 figures in text. July 24, 1961.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">4.</td> + <td>A new subspecies of the black myotis (bat) from eastern Mexico. By E. + Raymond Hall <ins title='Correction: was "anad"'>and</ins> + Ticul Alvarez. Pp. 69-72, 1 figure in text. December 29, 1961.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">5.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">6.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">7.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">8.</td> + <td>A new subspecies of ground squirrel (Spermophilus spilosoma) from Tamaulipas, + Mexico. By Ticul Alvarez. Pp. 121-124. March 7, 1962.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">9.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">10.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">11.</td> + <td>A new subspecies of wood rat (Neotoma) from northeastern Mexico. By + Ticul Alvarez. Pp. 139-143. April 30, 1962.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">12.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">13.</td> + <td>A new bat (Myotis) from Mexico. By E. Raymond Hall. Pp. 161-164, + 1 figure in text. May 21, 1962.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">14.</td> + <td>The mammals of Veracruz. By E. Raymond Hall + <ins title='Correction: was "anad"'>and</ins> Walter W. Dalquest. + Pp. 165-362, 2 figures. May 20, 1963.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">15.</td> + <td>The recent mammals of Tamaulipas, México. By Ticul Alvarez. Pp. 363-473, + 5 figures in text. May 20, 1963.</td> +</tr> +<tr> + <td> </td> + <td colspan="2">More numbers will appear in volume 14.</td> +</tr> +<tr> + <td class="text_rt vtop">Vol. 15.</td> + <td class="text_rt vtop">1.</td> + <td>The amphibians and reptiles of Michoacán, México. By William E. Duellman. + Pp. 1-148, pls. 1-6, 11 figures in text. December 20, 1961.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">2.</td> + <td>Some reptiles and amphibians from Korea. By Robert G. Webb, J. Knox + Jones, Jr., and George W. Byers. Pp. 149-173. January 31, 1962.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">3.</td> + <td>A new species of frog (Genus Tomodactylus) from western México. By + Robert G. Webb. Pp. 175-181, 1 figure in text. March 7, 1962.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">4.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">5.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td class="text_rt vtop">6.</td> + <td>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.</td> +</tr> +<tr> + <td> </td> + <td colspan="2">More numbers will appear in volume 15.</td> +</tr> +</table> +<br /> +<br /> + +<div class="trans_notes"> +<a name="typos" id="typos"></a><div class="caption2">Transcriber's Notes</div> + +<p>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. <a href="#Page_255">255</a>, +<a href="#Page_262">262</a>, <a href="#Page_267">267</a>, +<a href="#Page_277">277</a>, and <a href="#Page_278">278</a>).</p> + +<p>On page 279 under <a href="#Variation"><i>Variation</i></a> there appears to be a miscalculation:</p> + <div class="center">668 mm. + 182 mm. = 850 mm. <b>not</b> + 840 as in original text.</div> +<br /> +<br /> + +<div class="caption2">Typographical Corrections</div> +<br /> + +<table summary="Corrections"> +<tr> + <td class="brdbt">Page</td> + <td> </td> + <td class="brdbt">Correction</td> +</tr> +<tr> + <td><a href="#Page_264">264</a></td> + <td> </td> + <td>immaculaate ⇒ immaculate</td> +</tr> +<tr> + <td><a href="#Page_264">264</a></td> + <td> </td> + <td>chacteristic ⇒ characteristic</td> +</tr> +<tr> + <td><a href="#Page_266">266</a></td> + <td> </td> + <td>elevatons ⇒ elevations</td> +</tr> +<tr> + <td><a href="#Page_267">267</a></td> + <td> </td> + <td>Dumeril ⇒ Duméril</td> +</tr> +<tr> + <td><a href="#Page_277">277</a></td> + <td> </td> + <td>Duméil ⇒ Duméril</td> +</tr> +<tr> + <td><a href="#Page_279">279</a></td> + <td> </td> + <td>Tehauntepec ⇒ Tehuantepec</td> +</tr> +<tr> + <td><a href="#Page_280">280</a></td> + <td> </td> + <td>Deleted repeated "Oaxaca,"</td> +</tr> +<tr> + <td><a href="#Page_292">292</a></td> + <td> </td> + <td>primarly ⇒ primarily</td> +</tr> +<tr> + <td><a href="#Page_295">295</a></td> + <td> </td> + <td>hertetofaunal ⇒ herpetofaunal</td> +</tr> +<tr> + <td><a href="#Page_i">i</a></td> + <td> </td> + <td> V. 9 No. 12 - Pp. 363-387 ⇒ Pp. 363-384</td> +</tr> +<tr> + <td><a href="#Page_iii">iii</a></td> + <td> </td> + <td>V. 13 No. 8 - Decriptions ⇒ Descriptions</td> +</tr> +<tr> + <td><a href="#Page_iii">iii</a></td> + <td> </td> + <td>V. 14 No. 8 - anad ⇒ and</td> +</tr> +<tr> + <td><a href="#Page_iii">iii</a></td> + <td> </td> + <td>V. 14 No. 14 - anad ⇒ and</td> +</tr> +</table> +<br /> +<br /> +</div> +<br /> +<br /> +</div> + + + + + + + + +<pre> + + + + + +End of the Project Gutenberg EBook of A Revision of Snakes of the Genus +Conophis (Family Colubridae, from Middle America), by John Wellman + +*** END OF THIS PROJECT GUTENBERG EBOOK A REVISION OF SNAKES OF THE *** + +***** This file should be named 37512-h.htm or 37512-h.zip ***** +This and all associated files of various formats will be found in: + http://www.gutenberg.org/3/7/5/1/37512/ + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +http://www.pgdp.net + + +Updated editions will replace the previous one--the old editions +will be renamed. + +Creating the works from public domain print editions means that no +one owns a United States copyright in these works, so the Foundation +(and you!) can copy and distribute it in the United States without +permission and without paying copyright royalties. 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You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: A Revision of Snakes of the Genus Conophis (Family Colubridae, from Middle America) + +Author: John Wellman + +Release Date: September 23, 2011 [EBook #37512] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK A REVISION OF SNAKES OF THE *** + + + + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +http://www.pgdp.net + + + + + + + +Transcriber's Note + +Typographical corrections are listed at the end of this version. +The list of publications has been compiled after the article's text. + + * * * * * + + + 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) Dumeril, Bibron and Dumeril, Erpetologie Generale, + 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), Guenther, 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 + Dumeril, 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 Dumeril, Bibron and Dumeril 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 +Guenther (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 Mexico 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 Rio Blanco, + 20 km. WNW Piedras Negras, Veracruz, Mexico. E. _C. nevermanni_ + (ANSP 22424) "San Jose," Costa Rica. F. _C. pulcher_ (UIMNH + 33646) from Soconusco, Chiapas, Mexico. G. _C. vittatus_ (KU + 39626) from Atencingo, Puebla, Mexico. H. _C. vittatus_ (TCWC + 9473) from 1 mi. S Colotlipa, Guerrero, Mexico. I. _C. + vittatus_ (UMMZ 82653) from "vicinity of" Salina Cruz, Oaxaca, + Mexico. Approximately x 3/4.] + + +=Conophis lineatus= (Dumeril, Bibron and Dumeril) + + _Tomodon lineatum_ (in part) Dumeril, Bibron and Dumeril, + Erpetologie Generale, 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 +Chichen Itza, Yucatan, Mexico, 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 Yucatan Peninsula. The other two have +stripes on the dorsum and vary clinally in coloration from the north +(Veracruz, Mexico) 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_, Guenther, 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; Guenther, + 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_ Dumeril, 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 x 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 "Mexico." Since this +specimen is of _C. l. dunni_ and this subspecies occurs only south of +Mexico, 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 Mexico 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 Jose_," ANSP 3480, 12232. + +EL SALVADOR: _Morazan_: El Divisadero, CNHM 10999. _San Miguel: San +Pedro_, MCZ 57061. + +GUATEMALA: _El Peten_: Sojio (Toocog), AMNH 69969, 69986. +_Huehuetenango_: flood plain Rio Cuilco, W of Finca Canibal, 18 km. N +Tacana, UMMZ 98283. _Santa Rosa_: Santa Rosa, UMMZ 107339. + +HONDURAS: _no specific locality_, AMNH 32814, UF 7657. _Cortes: +Cofradia_, 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. + +MEXICO: _no specific locality_, BMNH 53.2.4.16. + +NICARAGUA: _no specific locality_, UMMZ 65633, USNM 25237. _Leon_: El +Polvon, 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= (Dumeril, Bibron and Dumeril) + + _Tomodon lineatum_ (in part) Dumeril, Bibron and Dumeril, + Erpetologie Generale, 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 Mexico. 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_ Dumeril, 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.--"Mexico," restricted to Veracruz, Veracruz, Mexico, 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 Dumeril, 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 Dumeril, Bibron and Dumeril (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 "Mexico," 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, Mexico, 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, Mexico (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 Mexico. + +_Distribution._--Semi-arid habitats on the coastal plain of Veracruz, +Mexico, from Tecolutla to Lerdo de Tejada and Piedras Negras (fig. 2). + +_Specimens examined._--Total of 27, as follows: MEXICO: _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 Rio, 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; Rio 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--"Peten," + 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_ Dumeril, + Bocourt and Mocquard, Mission Scientifique au Mexique et dans + l'Amerique Centrale, 2:648, 1886; Mueller, 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), Guenther, 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: "Yucatan," restricted to Chichen Itza, +Yucatan, Mexico 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 Chichen +Itza, Yucatan, 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 +Chichen Itza, Yucatan, 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 "Peten" (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 +Peten, Guatemala, due to the presence in the bottle of a piece of +paper inscribed "_Conophis vittatus_, Peten, 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 Yucatan 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 Peten 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 +Mexican 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 Mexico and Central America. + +_Distribution._--The Yucatan Peninsula: eastern Campeche, all of +Yucatan, 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 Peten, no specific locality_, USNM 4941. + +HONDURAS: _Colon_: Patuca, USNM 20271. + +MEXICO: _Campeche_: Champoton, UMMZ 73063-66; Encarnacion, CNHM +106462. _Yucatan: no specific locality_, BMNH 80.7.13.30; Chichen +Itza, 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; _Libre +Union_, CNHM 36298, 36302, 36310-11, 36314; Mayapan, CNHM 40720; +Merida, 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: +Rio Poas de Aserri (a few miles south of San Jose), 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 Jose, 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, Rio Tenorio, NMW 16838(5). "_San Jose_," 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_ Dumeril, 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_ Dumeril, + 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 Tonala, Chiapas, by + Smith and Taylor (1950:326)]. + + _Conophis lineatus_, Guenther, 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: Tonala, 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: "Peten," +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 Cahabon, Negro, or Salama +river basins--all places near the sugar plantation that Hague managed +at San Jeronimo, 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 Tonala, 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 +Tonala, 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 Mexico 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, Mexico, +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_: Duenas, 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. + +MEXICO: _Chiapas_: _Soconusco_, UIMNH 33646-47; Tonala, USNM 109707. + + +=Conophis vittatus= Peters + + _Tomodon lineatum_ (in part), Dumeril, Bibron and Dumeril, + Erpetologie Generale, 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_ Dumeril, Bocourt and + Mocquard, Mission Scientifique au Mexique et dans l'Amerique + Centrale, 2:644-646, pl. 38, fig. 7, 1886; Guenther, 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_ Dumeril, Bibron and Dumeril, + Erpetologie Generale, 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, Mexico); 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, + Mexico); 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 Mexico, 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_ x _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, Mexico, 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, Mexico, 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, Mexico, 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 (Michoacan, 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, Michoacan, Morelos and +Puebla. Specimens that would have to be considered intergrades between +the "subspecies" are found in Michoacan, 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 Mexican snakes. The only other +snake that occurs in western Mexico 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: MEXICO: _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 Ixtapan de la Sal, +KU 67648. _Laguna Coyuca_, CNHM 25881, UMMZ 80942. near La Union, AMNH +66337. _Magueyes, Laguna Coyuca_, AMNH 66149. _Playa Encantada_, TCWC +9470. 1 mi. S Tierra Colorada, KU 67649. near _Xaltinanguis, km. 405_, +CNHM 104947. _Michoacan_: Coalcoman, UMMZ 104693. _1/2 mi. SE +Coalcoman_, UMMZ 104492. _1 mi. N. Coalcoman_, UMMZ 112543. _1 mi. NE +Coalcoman_, 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. _Huajintlan, +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 Rio, 6 km. S Tehuantepec_, +UIMNH 6144-49, 37117-19, UMMZ 82649-51. _Rincon Bamba_, CNHM +105129-30, UIMNH 17615. _Salazar_, AMNH 66875. _vicinity of Salina +Cruz_, UMMZ 82653. _San Geronimo_, 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 Maria (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 Tequisistlan, 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. x 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. x 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. x 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). x 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 +Mexico _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 Chichen +Itza, Yucatan, 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, Mexico. 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 Mexico 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 Mexico 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. + + 1861. Contributions to the ophiology of Lower California, + Mexico and Central America. Proc. Acad. Nat. Sci. + Philadelphia, 13:292-306. December 28. + + 1867. Fifth contribution to the herpetology of tropical + America. Proc. Acad. Nat. Sci. Philadelphia, 18:317-323. + February 20. + + 1871. Ninth contribution to the herpetology of tropical + America. Proc. Acad. Nat. Sci. Philadelphia, 23(2):200-224. + October 24. + + 1876. On the batrachia and reptilia of Costa Rica. Journ. + Acad. Nat. Sci. Philadelphia, series 2, 8(4):93-154, 6 pls. + + 1895. The classification of the ophidia. Trans. Amer. Philos. + Soc., 18:186-219, 33 pls. April 15. + + 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. + + +DUMERIL, A. M. C., BIBRON, G., AND DUMERIL, A. H. A. + + 1854. Erpetologie generale, ou histoire naturelle des + reptiles. Paris, 7(pt. 2):xii + 785. February 25. + Atlas, 24 pp., 108 pls. + + +DUMERIL, 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. + Essex Inst., 16:1-46. January 9. + + 1884b. The reptiles and batrachians of North America. Mem. + Mus. Comp. Zool., 8(3):xxxi + 185 pp., 9 pls. July. + + +GUeNTHER, 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. + Soc. London, pt. 2:385-417, pp., pls. 27-33. + + +PETERS, W. + + 1860. Drei neue amerikanisches Schlangen. Monatsb. Akad. Wiss. + Berlin, 1860:517-521, pl., fig. 3. October. + + +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 Mexican 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 + +MUSEUM OF NATURAL HISTORY + + +Institutional libraries interested in publications exchange may obtain +this series by addressing the Exchange Librarian, University of Kansas +Library, Lawrence, Kansas. Copies for individuals, persons working in +a particular field of study, may be obtained by addressing instead the +Museum of Natural History, University of Kansas, Lawrence, Kansas. +There is no provision for sale of this series by the University +Library, which meets institutional requests, or by the Museum of +Natural History, which meets the requests of individuals. However, +when individuals request copies from the Museum, 25 cents should be +included, for each separate number that is 100 pages or more in +length, for the purpose of defraying the costs of wrapping and +mailing. + +* An asterisk designates those numbers of which the Museum's supply +(not the Library's supply) is exhausted. Numbers published to date, in +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, + 7 figures in text, 2 tables. October 10, 1951. + + Index. Pp. 651-681. + + *Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, + 41 plates, 31 figures in text. December 27, 1951. + + Vol. 5. Nos. 1-37 and index. Pp. 1-676, 1951-1953. + + *Vol. 6. (Complete) Mammals of Utah, taxonomy and distribution. By + Stephen D. Durrant. Pp. 1-549, 91 figures in text, + 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 Leon, + Mexico. 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 Leon, Mexico. By E. Raymond Hall. Pp. 531-538, + 1 figure in text. January 14, 1960. + + 21. Pleistocene pocket gophers from San Josecito Cave, Nuevo + Leon, Mexico. 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 + Michoacan, Mexico. 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, Mexico. + 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, Mexico. 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, Mexico. 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 Mexico. 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, Mexico. 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 Michoacan, Mexico. 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 + Mexico. 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 + Peten, 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 (Mexico) 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 Erpetologie Generale were +correcteded to Erpetologie Generale (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 => Dumeril + 277 Dumeil => Dumeril + 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 + + + + + +End of the Project Gutenberg EBook of A Revision of Snakes of the Genus +Conophis (Family Colubridae, from Middle America), by John Wellman + +*** END OF THIS PROJECT GUTENBERG EBOOK A REVISION OF SNAKES OF THE *** + +***** This file should be named 37512.txt or 37512.zip ***** +This and all associated files of various formats will be found in: + http://www.gutenberg.org/3/7/5/1/37512/ + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +http://www.pgdp.net + + +Updated editions will replace the previous one--the old editions +will be renamed. + +Creating the works from public domain print editions means that no +one owns a United States copyright in these works, so the Foundation +(and you!) can copy and distribute it in the United States without +permission and without paying copyright royalties. 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