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+The Project Gutenberg EBook of Systematic Status of the Colubrid Snake,
+Leptodeira discolor Gunther, by William E. Duellman
+
+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: Systematic Status of the Colubrid Snake, Leptodeira discolor Gunther
+
+Author: William E. Duellman
+
+Release Date: January 14, 2011 [EBook #34954]
+
+Language: English
+
+Character set encoding: ISO-8859-1
+
+*** START OF THIS PROJECT GUTENBERG EBOOK SYSTEMATIC STATUS OF THE ***
+
+
+
+
+Produced by Chris Curnow, Monika Krämer, Joseph Cooper and
+the Online Distributed Proofreading Team at
+http://www.pgdp.net
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+
+
+
+ UNIVERSITY OF KANSAS PUBLICATIONS
+ MUSEUM OF NATURAL HISTORY
+
+ Volume 11, No. 1, pp. 1-9, 4 figs.
+ July 14, 1958
+
+
+ Systematic Status of the Colubrid Snake,
+ Leptodeira discolor Günther
+
+ BY
+
+ WILLIAM E. DUELLMAN
+
+
+ UNIVERSITY OF KANSAS
+ LAWRENCE
+ 1958
+
+
+
+ UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY
+
+ Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
+ Robert W. Wilson
+
+
+ Volume 11, No. 1, pp 1-9, 4 figs.
+ Published July 14, 1958
+
+
+ UNIVERSITY OF KANSAS
+ Lawrence, Kansas
+
+
+ PRINTED IN
+ THE STATE PRINTING PLANT
+ TOPEKA, KANSAS
+ 1958
+
+ 27-6708
+
+
+
+
+ Systematic Status of the Colubrid Snake,
+ Leptodeira discolor Günther
+
+ BY
+
+ WILLIAM E. DUELLMAN
+
+
+At the time of completing my study of the genus _Leptodeira_ (1958) I
+had seen no specimens of _Leptodeira discolor_, a species described by
+Günther in 1860 and subsequently referred to the genus _Hypsiglena_ by
+Cope (1887), Boulenger (1894), and Mocquard (1908), and to the genus
+_Pseudoleptodeira_ by Taylor (1938). Günther's description was based on
+two syntypes (British Museum of Natural History numbers 1946.1.23.67
+and 68) collected in Oaxaca, México, by Auguste Sallé. Information
+concerning the scutellation and coloration of the syntypes was provided
+by J. C. Battersby; in my revisionary study (_op. cit._) this
+information was included in a short discussion of the species, which
+was referred to _incerta sedis_ until specimens could be examined and
+the relationships of the species determined.
+
+Through the courtesy of John M. Legler of the Museum of Natural History,
+University of Kansas, I have been able to study a specimen of
+_Leptodeira discolor_ obtained six miles southeast of Tamazulápam,
+Oaxaca, México, by J. R. Alcorn on June 22, 1955. Superficial
+examination of the external characters of this snake shows a striking
+resemblance to _Leptodeira_. The specimen has a vertical pupil, divided
+anal, 21 scale rows, and two apical pits. The enlarged posterior
+maxillary teeth are without a trace of a groove. Examination of the
+hemipenis revealed that the organ was bifurcate and had a forked sulcus;
+these penial characteristics are diagnostic of the subfamily
+Xenodontinae and not the subfamily Colubrinae that includes the genera
+_Hypsiglena_ and _Leptodeira_.
+
+Examination of all available xenodontine genera indicates that this
+snake belongs to a heretofore unnamed genus. In recognition of the
+mental torment customarily suffered by workers attempting to ascertain
+the relationships of the many genera of colubrid snakes, I propose the
+generic name
+
+
+ _Tantalophis_, new genus
+
+ _Leptodeira_ (in part), Günther, Proc. Zool. Soc. London, pp.
+ 317-318, 1860; Garman, Bull. Essex Inst., vol. 16, p. 23, January
+ 9, 1884; Dunn, Proc. Nat. Acad. Sci., vol. 22, pp. 697-698,
+ December, 1936; Duellman, Bull. Amer. Mus. Nat. Hist., vol. 114
+ (1), pp. 95-96, February 24, 1958.
+
+ _Hypsiglena_ (in part), Cope, Bull. U. S. Nat. Mus., no. 32, p. 78,
+ 1887; Günther, Biologia Centrali-Americana, Reptilia, pp. 137-138,
+ pl. 49, fig. A, October, 1894; Boulenger, Catalogue Snakes British
+ Museum, vol. 2, p. 211, September 23, 1894; Mocquard, _in_ Duméril
+ and Bocourt, Mission Scientifique Mexique l'Amerique Centrale, vol.
+ 3, p. 871, 1908; Amaral, Mem. Inst. Butantan, vol. 4, p. 183, May,
+ 1930.
+
+ _Pseudoleptodeira_ (in part) Taylor, Univ. Kansas Sci. Bull., vol.
+ 25, no. 15, p. 343, June 1, 1938.
+
+ _Type Species._--_Leptodeira discolor_ Günther, Proc. Zool. Soc.
+ London, pp. 317-318, 1860.
+
+ _Diagnosis._--A xenodontine colubrid snake having a bifurcate
+ hemipenis with a forked sulcus spermaticus, many longitudinal folds
+ on basal portion, and small spines and calyces on distal part; 12
+ or 13 maxillary teeth followed by short diastema and two somewhat
+ enlarged maxillary teeth lacking grooves; small parotid gland;
+ normal colubrid skull; no hypapophyses on posterior vertebrae;
+ elliptical pupils; two apical pits; smooth scales; normal colubrid
+ head shields; divided anal; paired caudals.
+
+ The generic name comes from the Greek Tantalos, a mythological
+ character symbolic of eternal torment, and from the Greek ophis
+ for snake.
+
+
+ _Tantalophis discolor_ (Günther) New comb.
+
+ The synonymy for the species is indicated in the account of the
+ genus. The description below of the species is based on an adult
+ male from 6 miles southeast of Tamazulápam, Oaxaca, México
+ (University of Kansas Museum of Natural History No. 40143).
+
+ _Scutellation._--Head shields normal; upper labials 7-7 (third and
+ fourth entering orbit); lower labials 9-9 (1-4 in contact with
+ anterior chin-shield, 4 and 5 in contact with posterior
+ chin-shield); preoculars 1-1 and not in contact with frontal;
+ postoculars 2-2; temporals 1-2-3, 1-2-3; nasals divided by a
+ distinct groove below nostril and faint groove above; portion of
+ rostral visible from above, one-third length of internasals;
+ internasals pentagonal and one-half as long as prefrontals;
+ parietal suture approximately as long as frontal; ventrals 178;
+ anal divided; caudals 80. Scales in 21 rows at midbody and showing
+ the following reduction:
+
+ 2 + 3 (130) 8 + 9 (162)
+ 21 --------------- 19 -------------- 17 (178)
+ 2 + 3 (130) 8 + 9 (152)
+
+ _Coloration._--Dorsal ground-color light brown and extending onto
+ edges of ventrals; transverse body blotches numbering 50, each
+ 1-1/2 to 3 scales long and separated by light interspaces 1-1/2 to
+ 2 scales long; blotches brownish black and extending onto second
+ scale row; lateral intercalary spots forming dark smudges on rows 1
+ and 2. Top of head black, flecked with tan; nape cream, followed by
+ dark band six scales long; dark nape stripe from posterior edges of
+ parietals to first dark body band. Venter cream-tan; throat and
+ labials cream; posterior margins of all upper labials and of lower
+ labials 1-3 black-edged (Figure 1).
+
+ _Size and Proportions._--Head and body 312 mm. long; tail 118 mm.,
+ amounting to 37.8 per cent of length of head and body.
+
+ _Variation._--Data on the syntypes of _Leptodeira discolor_
+ furnished by J. C. Battersby give some indication of the variation
+ in the species. Both are males, and they have 175 and 180 ventrals,
+ 88 and 89 caudals, 1 preocular, 2 postoculars, 1-2-3 temporals, 7
+ and 8 upper labials, 9 lower labials. They have body lengths of
+ 365 and 402 mm., total lengths of 509 and 555 mm., tail/body ratios
+ of 38.0 and 39.4. They have 51 and 54 dark blotches on the body.
+
+ [Illustration: Fig. 1. Dorsal and lateral views of the head of
+ _Tantalophis discolor_ (Günther). (KU No. 40143). × 7.]
+
+ _Skull._--The skull is typically colubrid and shows no
+ modifications. The quadrate has both a median and a lateral
+ depression, forming a strong lateral flange on the anterior edge;
+ the columellar process is elliptical, and the supra-columellar
+ crest is robust. The posteroinferior vomerine process extends
+ directly posteriorly and then angles sharply posterodorsally,
+ enclosing an elliptical vomerine fenestra. The lateral processes of
+ the premaxillary are slightly pointed; the median spine is
+ relatively thin and high. The pterygoids bear 23 and 21 teeth that
+ decrease in size posteriorly; the transpalatine articulating
+ process of the pterygoid is rounded, not robust; the lateral crest
+ is high and moderately robust; the depression in the ventral
+ surface of the pterygoid is moderate. There are 12 and 13 maxillary
+ teeth that increase in size posteriorly; these are followed by a
+ short diastema and two larger, solid teeth. The prediastemal teeth
+ are slightly curved and slender. The maxillary is laterally
+ compressed; the posterior knob is not robust; there is one foramen
+ in the lateral face of the bone (Figure 2). The 10 palatine teeth
+ are almost uniform in size; the dentary bears 19 teeth that
+ decrease in size posteriorly.
+
+ [Illustration: Fig. 2. Lateral view of the left maxillary of
+ _Tantalophis discolor_ (Günther). (KU No. 40143). × 17.]
+
+ A thin and otherwise small parotid gland or "venom sac" extends
+ posteriorly from beneath the eye to about the angle of the jaw; a
+ minute duct connects with the anteromedian surface and extends to
+ the fleshy part of the mouth at the posterior end of the maxillary
+ (Figure 3).
+
+ [Illustration: Fig. 3. Lateral view of the head of _Tantalophis
+ discolor_ (Günther), showing the position and relative size of the
+ parotid gland. (KU No. 40143). × 3.]
+
+ _Hemipenis._--In _situ_ the hemipenis extends to the posterior edge
+ of the thirteenth caudal. The unforked part of the organ is
+ bedecked with numerous heavy longitudinal folds alternating with
+ thinner folds. The basal parts of the two heads are covered with
+ moderate sized spines, those closest to the base and the sulcus
+ being the smallest. The distal parts of the heads are covered with
+ calyces. The sulcus bifurcates on the unforked part of the organ at
+ a point about two-thirds of the distance from the base to the
+ division of the organ. The sulcus is a deep groove between heavy
+ folds proximally and is a shallower furrow distally (Figure 4).
+
+ [Illustration: Fig. 4. Hemipenis of _Tantalophis discolor_
+ (Günther). The organ was cut on the ventral surface and opened.
+ (KU No. 40143). × 4.]
+
+_Relationships._--Using Dunn's (1928) arrangement of the American
+colubrid snakes as a guide permits the taxonomist to group _Tantalophis_
+with several genera, some of which occur in South America and others in
+the West Indies. Although the significance of such generic characters as
+scale pits and nature of the hemipenis is not clear, these characters
+must, of necessity, be utilized in attempting to ascertain the
+relationships of _Tantalophis_ to other colubrid snakes. Assuming that
+the primary divisions of the American colubrids into subfamilies based
+on the nature of the sulcus spermaticus and the presence or absence of
+hypapophyses on the posterior vertebrae have some reality, _Tantalophis_
+must be placed in the subfamily Xenodontinae comprising genera chiefly
+South American in their distribution, but with several genera in Middle
+America and a few in North America and the West Indies. In order to
+limit the number of genera to be compared with _Tantalophis_, only those
+xenodontines having apical pits and bifurcate hemipenis are considered.
+These include _Cyclagras_, _Drepanoides_, _Hypsirhynchus_, _Ialtris_,
+_Leimadophis_, _Pseudablabes_, _Siphlophis_, _Tachymenis_, _Tomodon_,
+and _Trypanurgos_. Aside from differences in scutellation,
+_Leimadophis_, _Siphlophis_, and _Trypanurgos_ have the heads of the
+hemipenes terminating in a disc, and _Ialtris_ has a plicate hemipenis.
+_Tomodon_ has basal spines on the hemipenis. The hemipenes of the other
+genera have proximal folds, distal spines, and distal calyces, not
+greatly unlike the condition found in _Tantalophis_. Of these,
+_Cyclagras_, _Hypsirhynchus_, and _Pseudablabes_ have round pupils and
+certain differences in scutellation. _Drepanoides_ and _Tachymenis_ have
+elliptical pupils like those of _Tantalophis_, but _Tachymenis_ has only
+one apical pit, and _Drepanoides_ has one apical pit or none. In the
+above characters no especially close relationship between _Tantalophis_
+and any one of these genera is apparent.
+
+If the characteristics usually employed in distinguishing and relating
+genera are ignored and other more subjective criteria are used, the
+relationships of _Tantalophis_ still remain obscure. Of the xenodontine
+genera _Tantalophis_ approaches _Leimadophis_ in general physiognomy;
+perhaps it represents an early divergent stock of _Leimadophis_ that has
+undergone radical changes in the hemipenis and other characters. On the
+other hand, if the nature of the hemipenis is of no importance in
+defining supergeneric groups of colubrid snakes, _Tantalophis_ may have
+its relationships with _Leptodeira_ and _Hypsiglena_. Although
+herpetologists have been working intensively on American colubrids for
+many decades, the relationships of the majority of the groups are not
+well understood. Until the hemipenes and skulls of all of the forms have
+been studied and compared, and the evolutionary significance has been
+determined for the characters of the hemipenes, dentition, and apical
+pits, our knowledge of the relationships of these snakes will be
+incomplete.
+
+_Remarks._--The individual on which this paper is based is the only
+specimen of the species with definite locality data. It is from a
+locality six miles southeast of Tamazulápam in northwestern Oaxaca. This
+town lies at an elevation of about 6500 feet in the upper reaches of the
+Balsas Basin, an arid interior valley that expands in its upper end to
+form a broad basin of rolling and dissected terrain ranging from about
+4000 to 6800 feet in elevation. The countryside around Tamazulápam is
+arid and supports plants of the genera _Prosopis_, _Acacia_, _Ipomoea_,
+and _Cassia_, and also columnar cacti. Oaks and pines grow on the
+limestone hills rising above the rolling valley. _Tantalophis_ may be
+endemic to the Balsas Basin, as are many other species of reptiles.
+However, if the snake has its relatives to the south in lower Central
+America and South America, such a distribution seems unlikely, even for
+an apparent relict.
+
+_Acknowledgments._--For permission to study and report on this specimen
+I am indebted to Dr. E. Raymond Hall and Mr. John M. Legler. I am
+grateful to Dr. Laurence C. Stuart for many helpful suggestions and to
+Dr. Norman E. Hartweg for placing at my disposal the facilities of the
+Museum of Zoology at the University of Michigan.
+
+
+LITERATURE CITED
+
+ AMARAL, A. DO
+
+ 1930 Estudos sobre ophidios neotropicos XVIII--Lista remissiva dos
+ ophidios da região neotropica. Mem. Inst. Butantan, 4:129-275.
+
+ BOULENGER, G. A.
+
+ 1894 Catalogue of the snakes in the British Museum (Natural
+ History). London, 2:xi + 382, pls. 1-20, figs. 1-25.
+
+ COPE, E. D.
+
+ 1887 Catalogue of the batrachians and reptiles of Central America
+ and Mexico. Bull. U. S. Nat. Mus., 32:1-98.
+
+ DUELLMAN, W. E.
+
+ 1958 A monographic study of the colubrid snake genus _Leptodeira_.
+ Bull. Amer. Mus. Nat. Hist., 114:1-152, pls. 1-31, figs. 1-25,
+ maps 1-25.
+
+ DUMÉRIL, A. M., and BOCOURT, F.
+
+ 1870-1909 Études sur les reptiles. Mission scientifique au Mexique
+ et dans l'Amerique Centrale, Recherches zoologiques. Paris,
+ Pt. 3, 1:xiv + 1012, pls. 1-77.
+
+ DUNN, E. R.
+
+ 1928 A tentative key and arrangement of the American genera of
+ Colubridae. Bull. Antivenin Inst. Amer., 2 (1):18-24.
+
+ 1936 Notes on North American _Leptodeira_. Proc. Nat. Acad. Sci.,
+ 22 (12):689-698.
+
+ GARMAN, S.
+
+ 1884 The North American reptiles and batrachians. Bull. Essex
+ Inst., 16:1-46, 3 figs.
+
+ GÜNTHER, A. C. L. G.
+
+ 1860 On new reptiles and fishes from Mexico. Proc. Zool. Soc.
+ London, pp. 316-319.
+
+ 1885-1902 Biologia Centrali-Americana. Reptilia and Batrachia.
+ London, pp. xx + 1-326, pls. 1-76.
+
+ TAYLOR, E. H.
+
+ 1938 Notes on Mexican snakes of the genus _Leptodeira_, with a
+ proposal of a new snake genus, _Pseudoleptodeira_. Univ.
+ Kansas Sci. Bull., 25:315-355, pls. 30-34, figs. 1-7.
+
+_Transmitted March 11, 1958. Contribution No. 11 from the Department of
+Biology, Wayne State University, Detroit 2, Michigan._
+
+
+
+
+
+End of the Project Gutenberg EBook of Systematic Status of the Colubrid
+Snake, Leptodeira discolor Gunther, by William E. Duellman
+
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+ The Project Gutenberg eBook of Systematic Status of the Colubrid Snake,
+ Leptodeira discolor G&uuml;nther, by William E. Duellman.
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+
+The Project Gutenberg EBook of Systematic Status of the Colubrid Snake,
+Leptodeira discolor Gunther, by William E. Duellman
+
+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: Systematic Status of the Colubrid Snake, Leptodeira discolor Gunther
+
+Author: William E. Duellman
+
+Release Date: January 14, 2011 [EBook #34954]
+
+Language: English
+
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+*** START OF THIS PROJECT GUTENBERG EBOOK SYSTEMATIC STATUS OF THE ***
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+
+
+
+
+<p><span class="pagenum"><a name="Front_cover" id="Front_cover">[Pg 1]</a></span></p>
+
+<div class="titlepage"><span class="smcap">University of Kansas Publications</span><br />
+<span class="smcap">Museum of Natural History</span>
+
+<br /><br />
+
+Volume 11, No. 1, pp. 1-9, 4 figs.<br />
+July 14, 1958</div>
+
+
+<div class="titlepage" style="font-size:200%;">Systematic Status of the Colubrid Snake,<br />
+Leptodeira discolor G&uuml;nther</div>
+
+<div class="titlepage">BY</div>
+
+<div class="titlepage" style="font-size:150%;">WILLIAM E. DUELLMAN</div>
+
+
+<div class="titlepage"><span class="smcap">University of Kansas</span><br />
+<span class="smcap">Lawrence</span><br />
+
+1958
+</div>
+
+<hr style="width:65%;" />
+
+<p><span class="pagenum"><a name="Inside_front_cover" id="Inside_front_cover">[Pg 2]</a></span></p>
+
+
+<div class="titlepage">
+<span class="smcap">University of Kansas Publications, Museum of Natural History</span><br />
+</div>
+
+<div class="titlepage">Editors: E. Raymond Hall, Chairman, Henry S. Fitch,<br />
+Robert W. Wilson<br /></div>
+
+
+<div class="titlepage">Volume 11, No. 1, pp 1-9, 4 figs.<br />
+Published July 14, 1958<br /><br /></div>
+
+
+<div class="titlepage">
+<span class="smcap">University of Kansas</span><br />
+Lawrence, Kansas<br /></div>
+
+
+<div style="text-align:center; margin-top:2em;">
+PRINTED IN<br />
+THE STATE PRINTING PLANT<br />
+TOPEKA, KANSAS<br />
+1958</div>
+
+<div style="text-align:center; margin-top:1em;">27-6708</div>
+
+<hr style="width:65%;" />
+<p><span class='pagenum'><a name="Page_3" id="Page_3">[Pg 3]</a></span></p>
+
+
+
+<h1>Systematic Status of the Colubrid Snake,<br />
+Leptodeira discolor G&uuml;nther</h1>
+
+<p class="center">BY</p>
+
+<p class="center">WILLIAM E. DUELLMAN</p>
+
+
+<p>At the time of completing my study of the genus <i>Leptodeira</i>
+(1958) I had seen no specimens of <i>Leptodeira discolor</i>, a species
+described by G&uuml;nther in 1860 and subsequently referred to the
+genus <i>Hypsiglena</i> by Cope (1887), Boulenger (1894), and Mocquard
+(1908), and to the genus <i>Pseudoleptodeira</i> by Taylor (1938).
+G&uuml;nther's description was based on two syntypes (British Museum
+of Natural History numbers 1946.1.23.67 and 68) collected in
+Oaxaca, M&eacute;xico, by Auguste Sall&eacute;. Information concerning the
+scutellation and coloration of the syntypes was provided by J. C.
+Battersby; in my revisionary study (<i>op. cit.</i>) this information was
+included in a short discussion of the species, which was referred to
+<i>incerta sedis</i> until specimens could be examined and the relationships
+of the species determined.</p>
+
+<p>Through the courtesy of John M. Legler of the Museum of Natural
+History, University of Kansas, I have been able to study a
+specimen of <i>Leptodeira discolor</i> obtained six miles southeast of
+Tamazul&aacute;pam, Oaxaca, M&eacute;xico, by J. R. Alcorn on June 22, 1955.
+Superficial examination of the external characters of this snake
+shows a striking resemblance to <i>Leptodeira</i>. The specimen has a
+vertical pupil, divided anal, 21 scale rows, and two apical pits. The
+enlarged posterior maxillary teeth are without a trace of a groove.
+Examination of the hemipenis revealed that the organ was bifurcate
+and had a forked sulcus; these penial characteristics are diagnostic
+of the subfamily Xenodontinae and not the subfamily Colubrinae
+that includes the genera <i>Hypsiglena</i> and <i>Leptodeira</i>.</p>
+
+<p>Examination of all available xenodontine genera indicates that
+this snake belongs to a heretofore unnamed genus. In recognition
+of the mental torment customarily suffered by workers attempting
+to ascertain the relationships of the many genera of colubrid snakes,
+I propose the generic name</p>
+
+
+<p class="center"><b>Tantalophis</b>, new genus</p>
+
+<div class="blockquot"><p><i>Leptodeira</i> (in part), G&uuml;nther, Proc. Zool. Soc. London, pp. 317-318, 1860;
+Garman, Bull. Essex Inst., vol. 16, p. 23, January 9, 1884; Dunn, Proc. Nat.
+Acad. Sci., vol. 22, pp. 697-698, December, 1936; Duellman, Bull. Amer. Mus.
+Nat. Hist., vol. 114 (1), pp. 95-96, February 24, 1958.</p>
+
+<p><i>Hypsiglena</i> (in part), Cope, Bull. U. S. Nat. Mus., no. 32, p. 78, 1887;
+G&uuml;nther, Biologia Centrali-Americana, Reptilia, pp. 137-138, pl. 49, fig. A,<span class='pagenum'><a name="Page_4" id="Page_4">[Pg 4]</a></span>
+October, 1894; Boulenger, Catalogue Snakes British Museum, vol. 2, p. 211,
+September 23, 1894; Mocquard, <i>in</i> Dum&eacute;ril and Bocourt, Mission Scientifique
+Mexique l'Amerique Centrale, vol. 3, p. 871, 1908; Amaral, Mem. Inst. Butantan,
+vol. 4, p. 183, May, 1930.</p>
+
+<p><i>Pseudoleptodeira</i> (in part) Taylor, Univ. Kansas Sci. Bull., vol. 25, no. 15,
+p. 343, June 1, 1938.</p>
+
+<p><i>Type Species.</i>&mdash;<i>Leptodeira discolor</i> G&uuml;nther, Proc. Zool. Soc. London, pp.
+317-318, 1860.</p>
+
+<p><i>Diagnosis.</i>&mdash;A xenodontine colubrid snake having a bifurcate hemipenis with
+a forked sulcus spermaticus, many longitudinal folds on basal portion, and small
+spines and calyces on distal part; 12 or 13 maxillary teeth followed by short
+diastema and two somewhat enlarged maxillary teeth lacking grooves; small
+parotid gland; normal colubrid skull; no hypapophyses on posterior vertebrae;
+elliptical pupils; two apical pits; smooth scales; normal colubrid head shields;
+divided anal; paired caudals.</p>
+
+<p>The generic name comes from the Greek &#932;&#945;&#957;&#964;&#945;&#955;&#959;&#963;, a mythological character
+symbolic of eternal torment, and from the Greek &#959;&#966;&#953;&#963; for snake.</p></div>
+
+
+<p class="center"><b>Tantalophis discolor</b> (G&uuml;nther) New comb.</p>
+
+<div class="blockquot"><p>The synonymy for the species is indicated in the account of the genus. The
+description below of the species is based on an adult male from 6 miles southeast
+of Tamazul&aacute;pam, Oaxaca, M&eacute;xico (University of Kansas Museum of Natural
+History No. 40143).</p>
+
+<p><i>Scutellation.</i>&mdash;Head shields normal; upper labials 7-7 (third and fourth
+entering orbit); lower labials 9-9 (1-4 in contact with anterior chin-shield,
+4 and 5 in contact with posterior chin-shield); preoculars 1-1 and not in contact
+with frontal; postoculars 2-2; temporals 1-2-3, 1-2-3; nasals divided by a
+distinct groove below nostril and faint groove above; portion of rostral visible
+from above, one-third length of internasals; internasals pentagonal and one-half
+as long as prefrontals; parietal suture approximately as long as frontal; ventrals
+178; anal divided; caudals 80. Scales in 21 rows at midbody and showing the
+following reduction:</p>
+
+<p class="center">
+2 + 3 (130)&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;8 + 9 (162)<br />
+&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;21 &mdash;&mdash;&mdash;&mdash;&mdash;&nbsp; 19&nbsp;&mdash;&mdash;&mdash;&mdash;&mdash;&nbsp; 17 (178)<br />
+2 + 3 (130)&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;8 + 9 (152)<br />
+</p>
+
+<p><i>Coloration.</i>&mdash;Dorsal ground-color light brown and extending onto edges of
+ventrals; transverse body blotches numbering 50, each 1&frac12; to 3 scales long and
+separated by light interspaces 1&frac12; to 2 scales long; blotches brownish black and
+extending onto second scale row; lateral intercalary spots forming dark smudges
+on rows 1 and 2. Top of head black, flecked with tan; nape cream, followed
+by dark band six scales long; dark nape stripe from posterior edges of parietals
+to first dark body band. Venter cream-tan; throat and labials cream; posterior
+margins of all upper labials and of lower labials 1-3 black-edged (Figure 1).</p>
+
+<p><i>Size and Proportions.</i>&mdash;Head and body 312 mm. long; tail 118 mm., amounting
+to 37.8 per cent of length of head and body.</p>
+
+<span class='pagenum'><a name="Page_5" id="Page_5">[Pg 5]</a></span>
+
+<div class="figcenter" style="width: 450px;">
+<img src="images/bild1.jpg" width="450" height="655" alt="Fig. 1." title="Fig. 1." />
+<span class="caption">Fig. 1. Dorsal and lateral views of the head of <i>Tantalophis discolor</i> (G&uuml;nther).
+(KU No. 40143). &times; 7.</span>
+</div>
+
+<p><i>Variation.</i>&mdash;Data on the syntypes of <i>Leptodeira discolor</i> furnished by J. C.
+Battersby give some indication of the variation in the species. Both are males,
+and they have 175 and 180 ventrals, 88 and 89 caudals, 1 preocular, 2 postoculars,
+1-2-3 temporals, 7 and 8 upper labials, 9 lower labials. They have<span class='pagenum'><a name="Page_6" id="Page_6">[Pg 6]</a></span>
+body lengths of 365 and 402 mm., total lengths of 509 and 555 mm., tail/body
+ratios of 38.0 and 39.4. They have 51 and 54 dark blotches on the body.</p>
+
+<p><i>Skull.</i>&mdash;The skull is typically colubrid and shows no modifications. The
+quadrate has both a median and a lateral depression, forming a strong lateral
+flange on the anterior edge; the columellar process is elliptical, and the supracolumellar
+crest is robust. The posteroinferior vomerine process extends directly
+posteriorly and then angles sharply posterodorsally, enclosing an elliptical
+vomerine fenestra. The lateral processes of the premaxillary are slightly
+pointed; the median spine is relatively thin and high. The pterygoids bear
+23 and 21 teeth that decrease in size posteriorly; the transpalatine articulating
+process of the pterygoid is rounded, not robust; the lateral crest is high and
+moderately robust; the depression in the ventral surface of the pterygoid is
+moderate. There are 12 and 13 maxillary teeth that increase in size posteriorly;
+these are followed by a short diastema and two larger, solid teeth. The
+prediastemal teeth are slightly curved and slender. The maxillary is laterally
+compressed; the posterior knob is not robust; there is one foramen in the
+lateral face of the bone (Figure 2). The 10 palatine teeth are almost uniform
+in size; the dentary bears 19 teeth that decrease in size posteriorly.</p>
+
+<div class="figcenter" style="width: 700px;">
+<img src="images/bild2.png" width="700" height="177" alt="Fig. 2." title="Fig. 2." />
+<span class="caption">Fig. 2. Lateral view of the left maxillary of <i>Tantalophis discolor</i> (G&uuml;nther).
+(KU No. 40143). &times; 17.</span>
+</div>
+
+<p>A thin and otherwise small parotid gland or "venom sac" extends posteriorly
+from beneath the eye to about the angle of the jaw; a minute duct
+connects with the anteromedian surface and extends to the fleshy part of the
+mouth at the posterior end of the maxillary (Figure 3).</p>
+
+
+<div class="figcenter" style="width: 600px;">
+<img src="images/bild3.png" width="600" height="207" alt="Fig. 3." title="Fig. 3." />
+<span class="caption">Fig. 3. Lateral view of the head of <i>Tantalophis
+discolor</i> (G&uuml;nther), showing the position and relative size of the parotid
+gland. (KU No. 40143). &times; 17.</span>
+</div>
+
+<p><i>Hemipenis.</i>&mdash;In <i>situ</i> the hemipenis extends to the posterior edge of the
+thirteenth caudal. The unforked part of the organ is bedecked with numerous
+heavy longitudinal folds alternating with thinner folds. The basal parts of
+the two heads are covered with moderate sized spines, those closest to the base
+and the sulcus being the smallest. The distal parts of the heads are covered<span class='pagenum'><a name="Page_7" id="Page_7">[Pg 7]</a></span>
+with calyces. The sulcus bifurcates on the unforked part of the organ at a
+point about two-thirds of the distance from the base to the division of the
+organ. The sulcus is a deep groove between heavy folds proximally and is a
+shallower furrow distally (Figure 4).</p>
+</div>
+
+<div class="figcenter" style="width: 480px;">
+<img src="images/bild4.png" width="480" height="615" alt="Fig. 4." title="Fig. 4." />
+<span class="caption">Fig. 4. Hemipenis of <i>Tantalophis discolor</i>
+(G&uuml;nther). The organ was cut on the ventral surface and opened.
+(KU No. 40143). &times; 4.</span>
+</div>
+
+<p><i>Relationships.</i>&mdash;Using Dunn's (1928) arrangement of the American
+colubrid snakes as a guide permits the taxonomist to group
+<i>Tantalophis</i> with several genera, some of which occur in South
+America and others in the West Indies. Although the significance
+of such generic characters as scale pits and nature of the hemipenis
+is not clear, these characters must, of necessity, be utilized in attempting
+to ascertain the relationships of <i>Tantalophis</i> to other
+colubrid snakes. Assuming that the primary divisions of the
+American colubrids into subfamilies based on the nature of the
+sulcus spermaticus and the presence or absence of hypapophyses on
+the posterior vertebrae have some reality, <i>Tantalophis</i> must be
+placed in the subfamily Xenodontinae comprising genera chiefly
+South American in their distribution, but with several genera in
+Middle America and a few in North America and the West Indies.<span class='pagenum'><a name="Page_8" id="Page_8">[Pg 8]</a></span>
+In order to limit the number of genera to be compared with <i>Tantalophis</i>,
+only those xenodontines having apical pits and bifurcate
+hemipenis are considered. These include <i>Cyclagras</i>, <i>Drepanoides</i>,
+<i>Hypsirhynchus</i>, <i>Ialtris</i>, <i>Leimadophis</i>, <i>Pseudablabes</i>, <i>Siphlophis</i>,
+<i>Tachymenis</i>, <i>Tomodon</i>, and <i>Trypanurgos</i>. Aside from differences in
+scutellation, <i>Leimadophis</i>, <i>Siphlophis</i>, and <i>Trypanurgos</i> have the
+heads of the hemipenes terminating in a disc, and <i>Ialtris</i> has a plicate
+hemipenis. <i>Tomodon</i> has basal spines on the hemipenis. The
+hemipenes of the other genera have proximal folds, distal spines, and
+distal calyces, not greatly unlike the condition found in <i>Tantalophis</i>.
+Of these, <i>Cyclagras</i>, <i>Hypsirhynchus</i>, and <i>Pseudablabes</i> have round
+pupils and certain differences in scutellation. <i>Drepanoides</i> and
+<i>Tachymenis</i> have elliptical pupils like those of <i>Tantalophis</i>, but
+<i>Tachymenis</i> has only one apical pit, and <i>Drepanoides</i> has one apical
+pit or none. In the above characters no especially close relationship
+between <i>Tantalophis</i> and any one of these genera is apparent.</p>
+
+<p>If the characteristics usually employed in distinguishing and relating
+genera are ignored and other more subjective criteria are
+used, the relationships of <i>Tantalophis</i> still remain obscure. Of the
+xenodontine genera <i>Tantalophis</i> approaches <i>Leimadophis</i> in general
+physiognomy; perhaps it represents an early divergent stock of
+<i>Leimadophis</i> that has undergone radical changes in the hemipenis
+and other characters. On the other hand, if the nature of the hemipenis
+is of no importance in defining supergeneric groups of colubrid
+snakes, <i>Tantalophis</i> may have its relationships with <i>Leptodeira</i>
+and <i>Hypsiglena</i>. Although herpetologists have been working intensively
+on American colubrids for many decades, the relationships
+of the majority of the groups are not well understood. Until
+the hemipenes and skulls of all of the forms have been studied and
+compared, and the evolutionary significance has been determined
+for the characters of the hemipenes, dentition, and apical pits, our
+knowledge of the relationships of these snakes will be incomplete.</p>
+
+<p><i>Remarks.</i>&mdash;The individual on which this paper is based is the
+only specimen of the species with definite locality data. It is from
+a locality six miles southeast of Tamazul&aacute;pam in northwestern
+Oaxaca. This town lies at an elevation of about 6500 feet in the
+upper reaches of the Balsas Basin, an arid interior valley that expands
+in its upper end to form a broad basin of rolling and dissected
+terrain ranging from about 4000 to 6800 feet in elevation. The
+countryside around Tamazul&aacute;pam is arid and supports plants of the
+genera <i>Prosopis</i>, <i>Acacia</i>, <i>Ipomoea</i>, and <i>Cassia</i>, and also columnar
+cacti. Oaks and pines grow on the limestone hills rising above the<span class='pagenum'><a name="Page_9" id="Page_9">[Pg 9]</a></span>
+rolling valley. <i>Tantalophis</i> may be endemic to the Balsas Basin, as
+are many other species of reptiles. However, if the snake has its
+relatives to the south in lower Central America and South America,
+such a distribution seems unlikely, even for an apparent relict.</p>
+
+<p><i>Acknowledgments.</i>&mdash;For permission to study and report on this
+specimen I am indebted to Dr. E. Raymond Hall and Mr. John M.
+Legler. I am grateful to Dr. Laurence C. Stuart for many helpful
+suggestions and to Dr. Norman E. Hartweg for placing at my disposal
+the facilities of the Museum of Zoology at the University of
+Michigan.</p>
+
+
+<h3>LITERATURE CITED</h3>
+
+<p><span class="smcap">Amaral, A. do</span></p>
+
+<div class="blockquot"><p>1930 Estudos sobre ophidios neotropicos XVIII&mdash;Lista remissiva dos
+ophidios da regi&atilde;o neotropica. Mem. Inst. Butantan, 4:129-275.</p></div>
+
+<p><span class="smcap">Boulenger, G. A.</span></p>
+
+<div class="blockquot"><p>1894 Catalogue of the snakes in the British Museum (Natural History).
+London, 2:xi + 382, pls. 1-20, figs. 1-25.</p></div>
+
+<p><span class="smcap">Cope, E. D.</span></p>
+
+<div class="blockquot"><p>1887 Catalogue of the batrachians and reptiles of Central America and
+Mexico. Bull. U. S. Nat. Mus., 32:1-98.</p></div>
+
+<p><span class="smcap">Duellman, W. E.</span></p>
+
+<div class="blockquot"><p>1958 A monographic study of the colubrid snake genus <i>Leptodeira</i>.
+Bull. Amer. Mus. Nat. Hist., 114:1-152, pls. 1-31, figs. 1-25, maps
+1-25.</p></div>
+
+<p><span class="smcap">Dum&eacute;ril, A. M.</span>, and <span class="smcap">Bocourt, F.</span></p>
+
+<div class="blockquot"><p>1870-1909 &Eacute;tudes sur les reptiles. Mission scientifique au Mexique et
+dans l'Amerique Centrale, Recherches zoologiques. Paris, Pt. 3,
+1:xiv + 1012, pls. 1-77.</p></div>
+
+<p><span class="smcap">Dunn, E. R.</span></p>
+
+<div class="blockquot"><p>1928 A tentative key and arrangement of the American genera of Colubridae.
+Bull. Antivenin Inst. Amer., 2 (1):18-24.</p>
+
+<p>1936 Notes on North American <i>Leptodeira</i>. Proc. Nat. Acad. Sci., 22
+(12):689-698.</p></div>
+
+<p><span class="smcap">Garman, S.</span></p>
+
+<div class="blockquot"><p>1884 The North American reptiles and batrachians. Bull. Essex Inst.,
+16:1-46, 3 figs.</p></div>
+
+<p><span class="smcap">G&uuml;nther, A. C. L. G.</span></p>
+
+<div class="blockquot"><p>1860 On new reptiles and fishes from Mexico. Proc. Zool. Soc. London,
+pp. 316-319.</p>
+
+<p>1885-1902 Biologia Centrali-Americana. Reptilia and Batrachia. London,
+pp. xx + 1-326, pls. 1-76.</p></div>
+
+<p><span class="smcap">Taylor, E. H.</span></p>
+
+<div class="blockquot"><p>1938 Notes on Mexican snakes of the genus <i>Leptodeira</i>, with a proposal
+of a new snake genus, <i>Pseudoleptodeira</i>. Univ. Kansas Sci. Bull.,
+25:315-355, pls. 30-34, figs. 1-7.</p></div>
+
+
+<p><i>Transmitted March 11, 1958. Contribution No. 11 from the Department of
+Biology, Wayne State University, Detroit 2, Michigan.</i>
+</p>
+
+
+
+
+
+
+
+
+
+<pre>
+
+
+
+
+
+End of the Project Gutenberg EBook of Systematic Status of the Colubrid
+Snake, Leptodeira discolor Gunther, by William E. Duellman
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+The Project Gutenberg EBook of Systematic Status of the Colubrid Snake,
+Leptodeira discolor Gunther, by William E. Duellman
+
+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: Systematic Status of the Colubrid Snake, Leptodeira discolor Gunther
+
+Author: William E. Duellman
+
+Release Date: January 14, 2011 [EBook #34954]
+
+Language: English
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+*** START OF THIS PROJECT GUTENBERG EBOOK SYSTEMATIC STATUS OF THE ***
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+ UNIVERSITY OF KANSAS PUBLICATIONS
+ MUSEUM OF NATURAL HISTORY
+
+ Volume 11, No. 1, pp. 1-9, 4 figs.
+ July 14, 1958
+
+
+ Systematic Status of the Colubrid Snake,
+ Leptodeira discolor Guenther
+
+ BY
+
+ WILLIAM E. DUELLMAN
+
+
+ UNIVERSITY OF KANSAS
+ LAWRENCE
+ 1958
+
+
+
+ UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY
+
+ Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
+ Robert W. Wilson
+
+
+ Volume 11, No. 1, pp 1-9, 4 figs.
+ Published July 14, 1958
+
+
+ UNIVERSITY OF KANSAS
+ Lawrence, Kansas
+
+
+ PRINTED IN
+ THE STATE PRINTING PLANT
+ TOPEKA, KANSAS
+ 1958
+
+ 27-6708
+
+
+
+
+ Systematic Status of the Colubrid Snake,
+ Leptodeira discolor Guenther
+
+ BY
+
+ WILLIAM E. DUELLMAN
+
+
+At the time of completing my study of the genus _Leptodeira_ (1958) I
+had seen no specimens of _Leptodeira discolor_, a species described by
+Guenther in 1860 and subsequently referred to the genus _Hypsiglena_ by
+Cope (1887), Boulenger (1894), and Mocquard (1908), and to the genus
+_Pseudoleptodeira_ by Taylor (1938). Guenther's description was based on
+two syntypes (British Museum of Natural History numbers 1946.1.23.67
+and 68) collected in Oaxaca, Mexico, by Auguste Salle. Information
+concerning the scutellation and coloration of the syntypes was provided
+by J. C. Battersby; in my revisionary study (_op. cit._) this
+information was included in a short discussion of the species, which
+was referred to _incerta sedis_ until specimens could be examined and
+the relationships of the species determined.
+
+Through the courtesy of John M. Legler of the Museum of Natural History,
+University of Kansas, I have been able to study a specimen of
+_Leptodeira discolor_ obtained six miles southeast of Tamazulapam,
+Oaxaca, Mexico, by J. R. Alcorn on June 22, 1955. Superficial
+examination of the external characters of this snake shows a striking
+resemblance to _Leptodeira_. The specimen has a vertical pupil, divided
+anal, 21 scale rows, and two apical pits. The enlarged posterior
+maxillary teeth are without a trace of a groove. Examination of the
+hemipenis revealed that the organ was bifurcate and had a forked sulcus;
+these penial characteristics are diagnostic of the subfamily
+Xenodontinae and not the subfamily Colubrinae that includes the genera
+_Hypsiglena_ and _Leptodeira_.
+
+Examination of all available xenodontine genera indicates that this
+snake belongs to a heretofore unnamed genus. In recognition of the
+mental torment customarily suffered by workers attempting to ascertain
+the relationships of the many genera of colubrid snakes, I propose the
+generic name
+
+
+ _Tantalophis_, new genus
+
+ _Leptodeira_ (in part), Guenther, Proc. Zool. Soc. London, pp.
+ 317-318, 1860; Garman, Bull. Essex Inst., vol. 16, p. 23, January
+ 9, 1884; Dunn, Proc. Nat. Acad. Sci., vol. 22, pp. 697-698,
+ December, 1936; Duellman, Bull. Amer. Mus. Nat. Hist., vol. 114
+ (1), pp. 95-96, February 24, 1958.
+
+ _Hypsiglena_ (in part), Cope, Bull. U. S. Nat. Mus., no. 32, p. 78,
+ 1887; Guenther, Biologia Centrali-Americana, Reptilia, pp. 137-138,
+ pl. 49, fig. A, October, 1894; Boulenger, Catalogue Snakes British
+ Museum, vol. 2, p. 211, September 23, 1894; Mocquard, _in_ Dumeril
+ and Bocourt, Mission Scientifique Mexique l'Amerique Centrale, vol.
+ 3, p. 871, 1908; Amaral, Mem. Inst. Butantan, vol. 4, p. 183, May,
+ 1930.
+
+ _Pseudoleptodeira_ (in part) Taylor, Univ. Kansas Sci. Bull., vol.
+ 25, no. 15, p. 343, June 1, 1938.
+
+ _Type Species._--_Leptodeira discolor_ Guenther, Proc. Zool. Soc.
+ London, pp. 317-318, 1860.
+
+ _Diagnosis._--A xenodontine colubrid snake having a bifurcate
+ hemipenis with a forked sulcus spermaticus, many longitudinal folds
+ on basal portion, and small spines and calyces on distal part; 12
+ or 13 maxillary teeth followed by short diastema and two somewhat
+ enlarged maxillary teeth lacking grooves; small parotid gland;
+ normal colubrid skull; no hypapophyses on posterior vertebrae;
+ elliptical pupils; two apical pits; smooth scales; normal colubrid
+ head shields; divided anal; paired caudals.
+
+ The generic name comes from the Greek Tantalos, a mythological
+ character symbolic of eternal torment, and from the Greek ophis
+ for snake.
+
+
+ _Tantalophis discolor_ (Guenther) New comb.
+
+ The synonymy for the species is indicated in the account of the
+ genus. The description below of the species is based on an adult
+ male from 6 miles southeast of Tamazulapam, Oaxaca, Mexico
+ (University of Kansas Museum of Natural History No. 40143).
+
+ _Scutellation._--Head shields normal; upper labials 7-7 (third and
+ fourth entering orbit); lower labials 9-9 (1-4 in contact with
+ anterior chin-shield, 4 and 5 in contact with posterior
+ chin-shield); preoculars 1-1 and not in contact with frontal;
+ postoculars 2-2; temporals 1-2-3, 1-2-3; nasals divided by a
+ distinct groove below nostril and faint groove above; portion of
+ rostral visible from above, one-third length of internasals;
+ internasals pentagonal and one-half as long as prefrontals;
+ parietal suture approximately as long as frontal; ventrals 178;
+ anal divided; caudals 80. Scales in 21 rows at midbody and showing
+ the following reduction:
+
+ 2 + 3 (130) 8 + 9 (162)
+ 21 --------------- 19 -------------- 17 (178)
+ 2 + 3 (130) 8 + 9 (152)
+
+ _Coloration._--Dorsal ground-color light brown and extending onto
+ edges of ventrals; transverse body blotches numbering 50, each
+ 1-1/2 to 3 scales long and separated by light interspaces 1-1/2 to
+ 2 scales long; blotches brownish black and extending onto second
+ scale row; lateral intercalary spots forming dark smudges on rows 1
+ and 2. Top of head black, flecked with tan; nape cream, followed by
+ dark band six scales long; dark nape stripe from posterior edges of
+ parietals to first dark body band. Venter cream-tan; throat and
+ labials cream; posterior margins of all upper labials and of lower
+ labials 1-3 black-edged (Figure 1).
+
+ _Size and Proportions._--Head and body 312 mm. long; tail 118 mm.,
+ amounting to 37.8 per cent of length of head and body.
+
+ _Variation._--Data on the syntypes of _Leptodeira discolor_
+ furnished by J. C. Battersby give some indication of the variation
+ in the species. Both are males, and they have 175 and 180 ventrals,
+ 88 and 89 caudals, 1 preocular, 2 postoculars, 1-2-3 temporals, 7
+ and 8 upper labials, 9 lower labials. They have body lengths of
+ 365 and 402 mm., total lengths of 509 and 555 mm., tail/body ratios
+ of 38.0 and 39.4. They have 51 and 54 dark blotches on the body.
+
+ [Illustration: Fig. 1. Dorsal and lateral views of the head of
+ _Tantalophis discolor_ (Guenther). (KU No. 40143). x 7.]
+
+ _Skull._--The skull is typically colubrid and shows no
+ modifications. The quadrate has both a median and a lateral
+ depression, forming a strong lateral flange on the anterior edge;
+ the columellar process is elliptical, and the supra-columellar
+ crest is robust. The posteroinferior vomerine process extends
+ directly posteriorly and then angles sharply posterodorsally,
+ enclosing an elliptical vomerine fenestra. The lateral processes of
+ the premaxillary are slightly pointed; the median spine is
+ relatively thin and high. The pterygoids bear 23 and 21 teeth that
+ decrease in size posteriorly; the transpalatine articulating
+ process of the pterygoid is rounded, not robust; the lateral crest
+ is high and moderately robust; the depression in the ventral
+ surface of the pterygoid is moderate. There are 12 and 13 maxillary
+ teeth that increase in size posteriorly; these are followed by a
+ short diastema and two larger, solid teeth. The prediastemal teeth
+ are slightly curved and slender. The maxillary is laterally
+ compressed; the posterior knob is not robust; there is one foramen
+ in the lateral face of the bone (Figure 2). The 10 palatine teeth
+ are almost uniform in size; the dentary bears 19 teeth that
+ decrease in size posteriorly.
+
+ [Illustration: Fig. 2. Lateral view of the left maxillary of
+ _Tantalophis discolor_ (Guenther). (KU No. 40143). x 17.]
+
+ A thin and otherwise small parotid gland or "venom sac" extends
+ posteriorly from beneath the eye to about the angle of the jaw; a
+ minute duct connects with the anteromedian surface and extends to
+ the fleshy part of the mouth at the posterior end of the maxillary
+ (Figure 3).
+
+ [Illustration: Fig. 3. Lateral view of the head of _Tantalophis
+ discolor_ (Guenther), showing the position and relative size of the
+ parotid gland. (KU No. 40143). x 3.]
+
+ _Hemipenis._--In _situ_ the hemipenis extends to the posterior edge
+ of the thirteenth caudal. The unforked part of the organ is
+ bedecked with numerous heavy longitudinal folds alternating with
+ thinner folds. The basal parts of the two heads are covered with
+ moderate sized spines, those closest to the base and the sulcus
+ being the smallest. The distal parts of the heads are covered with
+ calyces. The sulcus bifurcates on the unforked part of the organ at
+ a point about two-thirds of the distance from the base to the
+ division of the organ. The sulcus is a deep groove between heavy
+ folds proximally and is a shallower furrow distally (Figure 4).
+
+ [Illustration: Fig. 4. Hemipenis of _Tantalophis discolor_
+ (Guenther). The organ was cut on the ventral surface and opened.
+ (KU No. 40143). x 4.]
+
+_Relationships._--Using Dunn's (1928) arrangement of the American
+colubrid snakes as a guide permits the taxonomist to group _Tantalophis_
+with several genera, some of which occur in South America and others in
+the West Indies. Although the significance of such generic characters as
+scale pits and nature of the hemipenis is not clear, these characters
+must, of necessity, be utilized in attempting to ascertain the
+relationships of _Tantalophis_ to other colubrid snakes. Assuming that
+the primary divisions of the American colubrids into subfamilies based
+on the nature of the sulcus spermaticus and the presence or absence of
+hypapophyses on the posterior vertebrae have some reality, _Tantalophis_
+must be placed in the subfamily Xenodontinae comprising genera chiefly
+South American in their distribution, but with several genera in Middle
+America and a few in North America and the West Indies. In order to
+limit the number of genera to be compared with _Tantalophis_, only those
+xenodontines having apical pits and bifurcate hemipenis are considered.
+These include _Cyclagras_, _Drepanoides_, _Hypsirhynchus_, _Ialtris_,
+_Leimadophis_, _Pseudablabes_, _Siphlophis_, _Tachymenis_, _Tomodon_,
+and _Trypanurgos_. Aside from differences in scutellation,
+_Leimadophis_, _Siphlophis_, and _Trypanurgos_ have the heads of the
+hemipenes terminating in a disc, and _Ialtris_ has a plicate hemipenis.
+_Tomodon_ has basal spines on the hemipenis. The hemipenes of the other
+genera have proximal folds, distal spines, and distal calyces, not
+greatly unlike the condition found in _Tantalophis_. Of these,
+_Cyclagras_, _Hypsirhynchus_, and _Pseudablabes_ have round pupils and
+certain differences in scutellation. _Drepanoides_ and _Tachymenis_ have
+elliptical pupils like those of _Tantalophis_, but _Tachymenis_ has only
+one apical pit, and _Drepanoides_ has one apical pit or none. In the
+above characters no especially close relationship between _Tantalophis_
+and any one of these genera is apparent.
+
+If the characteristics usually employed in distinguishing and relating
+genera are ignored and other more subjective criteria are used, the
+relationships of _Tantalophis_ still remain obscure. Of the xenodontine
+genera _Tantalophis_ approaches _Leimadophis_ in general physiognomy;
+perhaps it represents an early divergent stock of _Leimadophis_ that has
+undergone radical changes in the hemipenis and other characters. On the
+other hand, if the nature of the hemipenis is of no importance in
+defining supergeneric groups of colubrid snakes, _Tantalophis_ may have
+its relationships with _Leptodeira_ and _Hypsiglena_. Although
+herpetologists have been working intensively on American colubrids for
+many decades, the relationships of the majority of the groups are not
+well understood. Until the hemipenes and skulls of all of the forms have
+been studied and compared, and the evolutionary significance has been
+determined for the characters of the hemipenes, dentition, and apical
+pits, our knowledge of the relationships of these snakes will be
+incomplete.
+
+_Remarks._--The individual on which this paper is based is the only
+specimen of the species with definite locality data. It is from a
+locality six miles southeast of Tamazulapam in northwestern Oaxaca. This
+town lies at an elevation of about 6500 feet in the upper reaches of the
+Balsas Basin, an arid interior valley that expands in its upper end to
+form a broad basin of rolling and dissected terrain ranging from about
+4000 to 6800 feet in elevation. The countryside around Tamazulapam is
+arid and supports plants of the genera _Prosopis_, _Acacia_, _Ipomoea_,
+and _Cassia_, and also columnar cacti. Oaks and pines grow on the
+limestone hills rising above the rolling valley. _Tantalophis_ may be
+endemic to the Balsas Basin, as are many other species of reptiles.
+However, if the snake has its relatives to the south in lower Central
+America and South America, such a distribution seems unlikely, even for
+an apparent relict.
+
+_Acknowledgments._--For permission to study and report on this specimen
+I am indebted to Dr. E. Raymond Hall and Mr. John M. Legler. I am
+grateful to Dr. Laurence C. Stuart for many helpful suggestions and to
+Dr. Norman E. Hartweg for placing at my disposal the facilities of the
+Museum of Zoology at the University of Michigan.
+
+
+LITERATURE CITED
+
+ AMARAL, A. DO
+
+ 1930 Estudos sobre ophidios neotropicos XVIII--Lista remissiva dos
+ ophidios da regiao neotropica. Mem. Inst. Butantan, 4:129-275.
+
+ BOULENGER, G. A.
+
+ 1894 Catalogue of the snakes in the British Museum (Natural
+ History). London, 2:xi + 382, pls. 1-20, figs. 1-25.
+
+ COPE, E. D.
+
+ 1887 Catalogue of the batrachians and reptiles of Central America
+ and Mexico. Bull. U. S. Nat. Mus., 32:1-98.
+
+ DUELLMAN, W. E.
+
+ 1958 A monographic study of the colubrid snake genus _Leptodeira_.
+ Bull. Amer. Mus. Nat. Hist., 114:1-152, pls. 1-31, figs. 1-25,
+ maps 1-25.
+
+ DUMERIL, A. M., and BOCOURT, F.
+
+ 1870-1909 Etudes sur les reptiles. Mission scientifique au Mexique
+ et dans l'Amerique Centrale, Recherches zoologiques. Paris,
+ Pt. 3, 1:xiv + 1012, pls. 1-77.
+
+ DUNN, E. R.
+
+ 1928 A tentative key and arrangement of the American genera of
+ Colubridae. Bull. Antivenin Inst. Amer., 2 (1):18-24.
+
+ 1936 Notes on North American _Leptodeira_. Proc. Nat. Acad. Sci.,
+ 22 (12):689-698.
+
+ GARMAN, S.
+
+ 1884 The North American reptiles and batrachians. Bull. Essex
+ Inst., 16:1-46, 3 figs.
+
+ GUeNTHER, A. C. L. G.
+
+ 1860 On new reptiles and fishes from Mexico. Proc. Zool. Soc.
+ London, pp. 316-319.
+
+ 1885-1902 Biologia Centrali-Americana. Reptilia and Batrachia.
+ London, pp. xx + 1-326, pls. 1-76.
+
+ TAYLOR, E. H.
+
+ 1938 Notes on Mexican snakes of the genus _Leptodeira_, with a
+ proposal of a new snake genus, _Pseudoleptodeira_. Univ.
+ Kansas Sci. Bull., 25:315-355, pls. 30-34, figs. 1-7.
+
+_Transmitted March 11, 1958. Contribution No. 11 from the Department of
+Biology, Wayne State University, Detroit 2, Michigan._
+
+
+
+
+
+End of the Project Gutenberg EBook of Systematic Status of the Colubrid
+Snake, Leptodeira discolor Gunther, by William E. Duellman
+
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