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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
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+
+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
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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 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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