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diff --git a/34127.txt b/34127.txt new file mode 100644 index 0000000..37cd666 --- /dev/null +++ b/34127.txt @@ -0,0 +1,1272 @@ +The Project Gutenberg EBook of Morphological Variation in a Population of +the Snake, Tantilla gracilis Baird and Girard, by Laurence M. Hardy and Charles J. Cole + +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: Morphological Variation in a Population of the Snake, Tantilla gracilis Baird and Girard + +Author: Laurence M. Hardy + Charles J. Cole + +Release Date: October 24, 2010 [EBook #34127] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK MORPHOLOGICAL VARIATION--SNAKE *** + + + + +Produced by Simon Gardner, Chris Curnow, Joseph Cooper and +the Online Distributed Proofreading Team at +https://www.pgdp.net + + + + + +Transcriber's Notes + +This Plain Text version represents italic typeface with _underscores_ +and small caps typeface with ALL CAPS. + +Only a few changes were necessary to the text in the case of +typographical errors, as follows: + +Page 623 Table 3 column heading: changed "or" to "of" (Number of +specimens). + +Page 625: changed "percent" to "per cent" (92 per cent of the time). + + * * * * * + + + + + + UNIVERSITY OF KANSAS PUBLICATIONS + MUSEUM OF NATURAL HISTORY + + + Volume 17, No. 15, pp. 613-629, 6 figs. + May 14, 1968 + + + Morphological Variation in a Population + of the Snake, + Tantilla gracilis Baird and Girard + + + BY + + + LAURENCE M. HARDY AND CHARLES J. COLE + + + UNIVERSITY OF KANSAS + LAWRENCE + 1968 + + + + + UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + + Editors: E. Raymond Hall, Chairman, Frank B. Cross, Henry S. Fitch, + J. Knox Jones, Jr. + + + Volume 17, No. 15, pp. 613-629, 6 figs. + Published May 14, 1968 + + + University of Kansas + Lawrence, Kansas + + + PRINTED BY + ROBERT R. (BOB) SANDERS, STATE PRINTER + TOPEKA, KANSAS + 1968 + + 31-9422 + + + + +Morphological Variation in a Population of the Snake, Tantilla gracilis +Baird and Girard + +By + +LAURENCE M. HARDY AND CHARLES J. COLE + + + + +INTRODUCTION + + +Variation of selected taxonomic characteristics of flat-headed snakes +(_Tantilla gracilis_ Baird and Girard) from several midwestern states +was studied by Force (1935), but she did not attempt a comprehensive +evaluation. Taylor (1936) reported on variation in _T. gracilis_ from +various localities in Kansas; Kirn, Burger, and Smith (1949) studied +selected structures of specimens of _T. gracilis_ from throughout its +range. The present study was undertaken to determine the variation in +both currently used and potential taxonomic characteristics of _T. +gracilis_ from one locality. + +We are thankful to Charles W. Myers, Gorgas Memorial Laboratory, for +suggestions concerning characteristics examined. We are indebted to Drs. +William G. Degenhardt, University of New Mexico, Herndon G. Dowling, New +York Zoological Society, Charles H. Lowe, University of Arizona, and +Richard G. Zweifel, American Museum of Natural History, for criticizing +the manuscript. Dr. William E. Duellman permitted us to study specimens +in the University of Kansas Museum of Natural History. + + +Materials and Methods + +The specimens examined were donated to the Museum of Natural History, +University of Kansas, by the late Paul Anderson of Independence, +Missouri. All specimens (KU numbers 83435-83680; N = 246) were collected +in the vicinity of Winfield, Cowley County, Kansas, by Charles E. Burt +and students from Southwestern College in the period from 1938 to 1941, +inclusive. + +Both authors gathered data on most characteristics and examined each +atypical individual. Hardy determined characters of the maxillae (which +were removed; N = 20), body and total lengths, and the scale formula of +KU 83620. + +We examined 22 characteristics of external morphology and 10 of the +maxillae. All paired characteristics (excepting those of the maxillae) +were examined on both sides of each snake and are referred to from the +left side to the right side (for example, "preoculars 1-2" means there +is one preocular on the left side and there are two on the right). +Oviducts and oviducal eggs were observed on specimens designated as +females; hemipenes were examined on specimens designated as males. Total +length was measured by straightening each snake along a 300 mm. scale. +Tail length was measured only on snakes having complete tails, and body +length (snout-vent) was considered as total length minus tail length. +The body length was measured on snakes having incomplete tails. + +Supralabials, infralabials, preoculars and postoculars were counted, and +we followed Peters (1960:9) in determining the nature of the temporals. +Occasionally the primary (anterior) temporal is separated from the +postocular by contact of the parietal and a supralabial (Taylor, +1936:338). We followed Peters (1964:219-220) in determining +characteristics of the nasal and we looked for contact of the prefrontal +with particular supralabials (Taylor, 1936:338). We determined which +supralabials are in contact with the eye and whether the mental is in +contact with the anterior pair of chin-shields. + +The number of rows of dorsal scales was determined at five locations: a) +around the neck--counts were made from each side diagonally and +posteriorly from the anteriormost dorsal scale in contact with the first +ventral (see below)--counts from the left and right sides are the first +and second counts, respectively, recorded in the scale formula; b) one +head length posterior to head--counts were made either diagonally and +posteriorly or in a zig-zag pattern, beginning at the end of the second +count around the neck--this is the third count recorded in the scale +formula; c) midbody (approximated)--counts were made either diagonally +and posteriorly or in a zig-zag pattern--this is the fourth count in the +scale formula; d) anterior to anus (preanal)--counts were made in a +zig-zag pattern beginning on one side and ending on the other at the +first dorsal scale in contact with the anal plate--this is the fifth +count in the scale formula. + +We followed the method of Dowling (1951:98-99) in counting ventrals, and +the anal plate was determined as entire, divided, or partly divided. +Subcaudals were counted on only the right side beginning at the first +scale that contacts a corresponding scale from the opposite side; fusion +of particular pairs of subcaudals was determined. The "spine" at the tip +of the tail was not counted. The head scutellation of each specimen was +examined for abnormalities. + +We determined the number of fangs and the number of maxillary teeth +anterior to them and recorded the presence or absence of a diastema +between the anterior fang and the maxillary tooth immediately preceding +it. The diastema, if present, is a space distinctly wider than the +spaces separating the prediastemal teeth. + +The point of intersection of two perpendicular lines of an ocular grid +determined which tooth-socket occurs directly opposite the distal tip of +the suborbital process (Fig. 5b), but if that point occurs between two +sockets then the number recorded was the number of the tooth immediately +anterior to that point plus one-half. + +The anterior and posterior edges of the suborbital process were +determined as parallel or not at any point by aligning them with +parallel lines on an ocular grid (Fig. 5b). + +The angle of the posterior edge of the lateral flange was determined as +an acute, obtuse, or right angle by fitting it to a pair of +perpendicular lines of an ocular grid (Fig. 5b). + +The number of lateral anterior foramina (Fig. 4b) was determined. If +only two are present, their lengths (anterior to posterior) were +compared by measuring them with an ocular micrometer. Also, if two +lateral anterior foramina are present, the point of intersection of two +perpendicular lines of an ocular grid determined which tooth-socket +occurs directly below the center of the posterior foramen (Fig. 4b). + + + + +RESULTS + + +General Characteristics + +From the data given by Force (1935), Taylor (1936), and Kirn _et al._ +(1949), and from our own data, we conclude that a typical specimen of +_gracilis_, meaning a specimen having the usual characteristics of the +species, has the following characteristics in combination: supralabials +6-6; infralabials 6-6; preoculars 1-1; postoculars 1-1; temporals 1 + 1 +on each side; nasal divided below naris; supralabials 3 + 4 entering +orbit; mental in contact with chin-shields; all five dorsal scale counts +15; and plate divided. (In the present work we use the words usual, +unusual, typical, and atypical with neither quotation marks nor +apology). + +Of the 244 specimens on which we could examine each of these +characteristics, only 125 (51.2%) have all of the typical +characteristics in combination. The 119 specimens that do not possess +all of the typical characteristics in combination include 31 (12.7% of +the 244) that lack at least two of the typical characteristics. + + +Sex + +The sample (N = 246) is composed of 107 (43.5%) females and 139 (56.5%) +males. + + +Measurements + +Females have total lengths from 96 to 244 mm. (mean, 173.3; N = 79). One +female (KU 83480) measuring 244 mm. may be the longest specimen known +(Conant, 1958, reports the longest as 9-1/8 inches--approximately 232 +mm.). Males have total lengths from 96 to 215 mm. (mean, 162.4; N = +109). Females have tail lengths from 16 to 50 mm. (mean, 34.6; N = 79), +and males have tail lengths from 21 to 53 mm. (mean, 37.8; N = 109). + +The ratio of tail length to total length in females is from 0.17 to 0.22 +(mean, 0.20; N = 79); in males it is from 0.21 to 0.27 (mean, 0.23; N = +109; Fig. 1). Juveniles (those less than 125 mm. in total length +according to Force, 1935: tables 1 and 2) have proportionally shorter +tails than do adults; the ratio of tail length divided by total length +is 0.17-0.21 (mean, 0.18; N = 14) in females and 0.21-0.23 (mean, 0.22; +N = 16) in males. Adult females (125 mm. or more in total length) have +ratios from 0.18 to 0.22 (mean, 0.20; N = 65) and adult males have +ratios from 0.21 to 0.27 (mean, 0.23; N = 93). + +[Illustration: FIG. 1. Ratio of tail length to total length in females +and males of _T. gracilis_.] + +Of the 246 specimens examined, 58 have incomplete tails. Of these 58 +specimens, six had freshly broken tails, so we assume that 52 (21.1% of +the 246) have incomplete tails resulting from natural causes; the +remaining six specimens could have had their tails broken at or soon +after capture, assuming that none of the snakes was retained in +captivity for an extended period. + + +Supralabials + +The supralabials are either six (usually) or seven on a side (Table 1). +When seven supralabials are present on either one or both sides, the +addition results from one of the following conditions: a) 5 is divided +to form 5 and 6 (N = 6); b) 2 and 3 form 2, 3, and 4 (N = 1); c) 5 and 6 +form 5, 6, and 7 (N = 12); d) 4, 5, and 6 form 4, 5, 6, and 7 (N = 1); +e) 1 and 2 form 1, 2, and 3 (N = 1); f) 1, 2, and 3 form 1, 2, 3, and 4 +(N = 1); g) 2 is fused with the nasal, and 3 and 4 are reduced to +granules (N = 1); h) type of addition unknown (N = 5). + +TABLE 1. Variation in Number of Supralabials (N = 245) and +Infralabials (N = 246) of Tantilla gracilis. + + ------------+----------+-----------+------------+---------+---------- + Number |Number |Per cent |Number |Number |Per cent + of |of |of |of |of |of + Supralabials|specimens |occurrence |infralabials|specimens|occurrence + ------------+----------+-----------+------------+---------+---------- + 6-6 | 221 | 90.2 | 5-5 | 5 | 2.0 + 6-7 | 10 | 4.1 | 5-6 | 4 | 1.6 + 7-6 | 10 | 4.1 | 6-5 | 5 | 2.0 + 7-7 | 4 | 1.6 | 6-6 | 229 | 93.1 + | | | 6-7 | 1 | 0.4 + | | | 7-6 | 2 | 0.8 + ------------+----------+-----------+------------+---------+---------- + +In three of the four specimens with 7-7 supralabials, the type of +addition is the same on each side, whereas the other specimen has a +different type of addition on each side. Of the 23 examples for which +the type of addition is known, 19 (82.6%) have the increase in number on +the posterior half of the series. + +In one specimen the third supralabial on the right is diagonally divided +and the resulting upper part does not contact the edge of the lip; hence +the specimen has 6-6 supralabials. + + +Infralabials + +The infralabials are five, six (usually), or seven on a side (Table 1). +When five infralabials are present on either one or both sides, the +reduction results from one of the following conditions: a) 2 and 3 are +fused (N = 3); b) 1, 2, and 3 form 1 and 2 (N = 6); c) 2, 3, and 4 form +2 and 3 (N = 3); d) 5 is separated from the edge of the lip by a +projection from an adjacent infralabial (N = 1); e) 1, 2, 3, and 4 form +1, 2, and 3 (N = 2); f) type of reduction unknown (N = 4). + +Of the specimens with 5-5 infralabials, two have the same type of +reduction on each side, one has a different type of reduction on each +side, and the type of reduction is unknown for the other two specimens. + +When seven infralabials are present on either one or both sides, the +addition can result from infralabials 2 and 3 forming 2, 3, and 4 (N = +2); the type of addition is unknown in one specimen. + +In one specimen the second infralabial on the right side is reduced to a +small wedge-shaped granule and in another specimen several infralabials +on both sides are irregularly divided horizontally, although both +specimens have six infralabials on the sides having the abnormalities. + + +Preoculars + +Of the 245 specimens examined, 242 (98.8%) have one preocular on each +side. Both preoculars are absent from two specimens, and one specimen +has 1-0. In each case when a preocular is lacking, the "preocular" and +prefrontal are fused. One specimen having 1-1 preoculars has both of +them reduced to small vertical slivers. + + +Postoculars + +Of the 245 specimens examined, 242 (98.8%) have one postocular on each +side. Postoculars are 2-1 in two specimens and 1-2 in one. In one +specimen the lower of the two postoculars on each side is very small, +being approximately 1/6 the diameter of the upper one (measured with an +ocular micrometer). + + +Temporals + +Of the 245 specimens examined, 235 (95.9%) have 1 + 1 temporals on each +side. Four specimens exhibit addition of temporals: one specimen has two +secondary temporals on the left, and one has a tertiary temporal on the +left. Two specimens have partly healed head injuries that probably +caused atypical temporal conditions: one specimen has two primary +temporals on the left, and one has two secondary temporals on the left +and two upper and one lower secondary temporal on the right. + +Six specimens exhibit reduction of temporals: one lacks a primary +temporal on the left ("temporal" is fused with sixth supralabial), two +lack secondary temporals on the right, and three lack secondary +temporals on the left. The right secondary "temporal" of one specimen is +separated from the supralabials by a posterior extension of the primary +temporal. + +Of the 246 specimens examined, 200 (81.3%) have both primary temporals +in contact with their adjacent postoculars. Thirteen specimens (5.3%) +have only the left primary temporal in contact with the postocular and +eight (3.3%) have only the right primary temporal in contact with the +postocular. Twenty-five specimens (10.2%) have both primary temporals +separated from the postoculars. Separation of the primary temporal from +the postocular, whenever it occurs, results from contact of mutual +extensions from the adjacent parietal and supralabial (usually the +fifth). + + +Nasal + +Of the 243 specimens examined, 237 (97.5%) have the nasal divided below +the naris on both sides. Three specimens have the nasal completely +divided (above and below the naris) on only the left side; one specimen +has a completely divided nasal on only the right side. A groove, but not +a complete division of the nasal, is present above the left naris of one +specimen. One specimen lacks the typical division below the naris on +both sides, but a complete suture is present posterior to the naris and +the resulting postnasal scales are, according to position, loreals. + + +Contact of prefrontal with supralabials + +Of the 243 specimens examined, 147 (60.5%) have the prefrontal separated +from the labials on each side. Of the remaining 96 specimens, 53 (21.8% +of the 243) have the prefrontals in contact with the second labial on +each side; five other variations of this characteristic were observed +(Table 2). + +TABLE 2. Variation in Contact of the Prefrontals and the Labials in 243 +Specimens of Tantilla gracilis. + + ----------------------------------------------------+---------+---------- + |Number |Per cent + CHARACTERISTIC |of |of + |specimens|occurrence + ----------------------------------------------------+---------+---------- + No contact | 147 | 60.5 + Contact of prefrontal with second labial on right | 26 | 10.7 + Contact of prefrontal with second labial on left | 10 | 4.1 + Contact of prefrontal with each second labial | 53 | 21.8 + Contact of prefrontals with labial two on left and | | + labial three on right | 1 | 0.4 + Contact of prefrontals with labial three on left and| | + labial two on right | 3 | 1.2 + Contact of prefrontals with labial two on left and | | + labials two and three on right | 3 | 1.2 + ----------------------------------------------------+---------+---------- + +The specimen having the prefrontal in contact with labial 2 on the left +and labial 3 on the right has seven supralabials on the right, the extra +labial being added anterior to the eye. In two of the three specimens +with the prefrontals in contact with labial 3 on the left and labial 2 +on the right, the preoculars and prefrontals are fused on each side; the +third specimen has each preocular reduced to a small vertical sliver. Of +the three specimens having the prefrontals in contact with labial 2 on +the left and labials 2 and 3 on the right, two are typical with respect +to other characteristics that might result in this atypical condition, +but one has the prefrontals and preoculars fused. + + +Supralabials entering orbit + +Of the 246 specimens examined, 233 (94.7%) have the third and fourth +supralabials entering each eye (Table 3). All snakes that have the +fourth and fifth labials entering the orbit on either side also have +seven supralabials on the same side. One specimen having only the fourth +labial entering the eye on the right side also has six supralabials on +that side, but the third supralabial is split diagonally and the part +contacting the eye does not contact the lip and hence is not a +supralabial. The remaining three atypical conditions (4/3 + 4; 3 + 4/3 + +4 + 5; 3 + 4 + 5/3 + 4) occur only one time each and on snakes having +6-6 supralabials. + + +Contact of mental with chin-shields + +Of the 246 specimens examined, 190 (77.2%) have the mental in contact +with both anterior chin-shields. The mental is in contact with only the +left anterior chin-shield in 15 specimens, only the right anterior +chin-shield in 2 specimens, and the mental does not contact either +anterior chin-shield in 39 specimens. Separation of the mental and an +anterior chin-shield always results from the posterior elongation of a +first infralabial. + +TABLE 3. Variation in Number of Supralabials Entering the Eye in 246 +Specimens of Tantilla gracilis. + + ----------------------+-----------+------------ + SUPRALABIALS | Number of | Per cent of + ----------------------+ specimens | occurrence + Left | Right | | + ----------+-----------+-----------+------------ + 3 + 4 | 3 + 4 | 233 | 94.7 + 4 + 5 | 3 + 4 | 4 | 1.6 + 4 | 3 + 4 | 1 | 0.4 + 3 + 4 | 4 + 5 | 5 | 2.0 + 3 + 4 | 4 | 1 | 0.4 + 3 + 4 | 3 + 4 + 5 | 1 | 0.4 + 3 + 4 + 5 | 3 + 4 | 1 | 0.4 + ----------+-----------+-----------+------------ + + +Dorsal scale rows + +Of the 245 specimens examined for this characteristic, 228 (92.7%) have +scale counts of 15-15-15-15-15. The number of scale rows around the neck +and one head length behind the head are 15 except in a single specimen +that has 14 scale rows at these three places. The number of scale rows +around midbody is 15 in 244 of 246 specimens (99.2%); two specimens have +14 scale rows around midbody. The number of preanal scale rows is more +variable; counts of 14 through 17 were recorded. Fourteen specimens have +14 preanal scale rows (5.7%), 229 have 15 preanal rows (93.1%), two have +16 preanal rows (0.8%), and one has 17 preanal rows (0.4%). Known causes +for atypical numbers of preanal scale rows are listed in Table 4. A +preanal count at one head length anterior to the anus would have omitted +most of the above variation in numbers of preanal scale rows. + +TABLE 4. Atypical Numbers of Preanal Scale Rows in Specimens of Tantilla +gracilis. + + -------+---------+--------------------------------------------+---------- + Preanal|Number | |Per cent + scales |of | CAUSE |of + |specimens| |occurrence + -------+---------+--------------------------------------------+---------- + | |Scale rows seven and eight on left fuse | + 14 | 10 |within the last five ventrals | 4.1 + | | | + | |Scale row two on the right is lost adjacent | + 14 | 1 |to the penultimate ventral | 0.4 + | | | + | |Scale row eight on the left divides adjacent| + 16 | 1 |to the last ventral | 0.4 + | | | + | |A scale row is added on the right adjacent | + 16 | 1 |to and anterior to the anal plate | 0.4 + | | | + | |A scale row is added on each side adjacent | + 17 | 1 |to and anterior to the anal plate | 0.4 + -------+---------+--------------------------------------------+---------- + +The complete dorsal scale formula for KU 83620, a male, is as follows: + + 6 + 7(2) + 7(4) 6 + 7(11) + 15 --------- 13 --------- 14 --------- 15 ---------- 13 --------------, + 6 + 7(2) +7(5) 6 + 7(11) 6 = 6 + 7(17) + + 14 ------------- 13---------, + 6 + 7(18) +7(25) + + + 7(26) 6 + 7(38) +7(39) + 14 ---------- 15 ---------- 14 ------------ 13---------- 14----------, + 6 + 7(36) +7(42) + + 15 --------- 14 ---------, + -6(45) +7(53) + + 7 + 8(67) + 7(69) + 15 ------------ 14 ----------- 15 ----------- 14 ---------- 15----------, + 6 + 7(90) +7(93) 6 + 7(99) + + 14 ------------ 15(122). + + 7(100) + +Specimen 83620 is the only one for which we have determined the complete +dorsal scale formula. Obviously the formula is complex, and possibly the +specimen is atypical in this regard. Nevertheless, addition and +reduction of scale rows is apparent on other specimens and can be easily +observed by examining specimens under a dissection microscope. + + +Anal plate + +Of the 246 specimens examined, 232 (94.3%) have the anal plate divided. +Of the 14 specimens having undivided anal plates, four have an +incomplete groove anteriorly in the plate. + + +Ventrals + +[Illustration: FIG. 2. Number of ventrals in females and males of _T. +gracilis_.] + +Females have from 122 to 137 (mean, 130.8 +- 0.26; N = 107) ventrals; +males have from 117 to 131 (mean, 122.3 +- 0.21; N = 139; Fig. 2). Of the +246 specimens examined, 29 (11.8%; 11 females and 18 males) have an +added half-ventral immediately anterior to the anal plate. Of these +specimens, 27 have the added half-ventral on the left side and two have +it on the right. Five specimens have the last ventral divided similar to +the normal division of the anal plate. One specimen has the last ventral +incompletely divided transversely. Excluding the last ventral, seven +specimens have divided ventrals (half-ventrals). Four of these specimens +have added half-ventrals on the right side (two specimens with one +half-ventral each, one specimen with two half-ventrals, and one specimen +with three half-ventrals), and four have added half-ventrals on the left +side (one half-ventral on each specimen). + + +Subcaudals + +Females have from 37 to 47 (mean, 42.0 +- 0.20; N = 79) subcaudals; males +have from 43 to 55 (mean, 48.4 +- 0.23; N = 109; Fig. 3). Of the 187 +specimens examined, 13 (7.0%) have the last pair of subcaudals fused. +One specimen has left subcaudal 28 fused with both adjacent subcaudals +(also fused) on the right. While reading proof we found two omissions in +Fig. 3; there should be illustrated two males with 43 subcaudals and +three males with 44 subcaudals. + +[Illustration: FIG. 3. Number of subcaudals in females and males of _T. +gracilis_.] + + +Ventrals minus subcaudals + +The value for number of ventrals minus the number of subcaudals varies +from 80 to 96 (mean, 88.8 +- 0.39) for 79 females, and from 67 to 83 +(mean, 73.8 +- 0.28) for 109 males. + + +Additional scale characteristics + +Of the 246 specimens examined, four (1.6%) have the rostral in contact +with the left prefrontal, and hence the internasals are separated from +one another. One snake has approximately one-fourth (anteriorly) of the +suture between the parietals fused. One specimen has the posterior edge +of the frontal fused with the left parietal. One specimen has the +parietal shield on the right side abbreviated posteriorly. One specimen +has both prefrontals fused with the frontal, although an incomplete +suture is present. + + +Maxillary characteristics (_Figs. 4 and 5_) + +Maxillary teeth anterior to fangs 10-14 (mean, 12.1; N = 20); +tooth-socket number 6.5-8.5 (mean, 7.2; N = 20) directly opposite distal +tip of suborbital process; lateral anterior foramina 1-3 (mean, 1.8; N = +20); tooth number 6-9 (mean, 6.9; N = 12) directly below center of +posterior foramen (if only two foramina present). Three specimens lack a +diastema and 17 specimens have a diastema. The anterior and posterior +edges of the suborbital process are parallel in two specimens and not +parallel in 14; the process is broken in four specimens. The lengths +(anterior to posterior) of the lateral anterior foramina (if only two +are present) are approximately equal in six specimens, and in six the +anteriormost one is the longest; six specimens have one foramen and two +specimens have three foramina. All specimens examined have two fangs, +with the posterior edge of the lateral flange forming an obtuse angle. + +[Illustration: FIG. 4. Lateral view of right maxilla of _T. gracilis_ +(KU 83484), x 39. Actual length of bone is 2.44 mm. A. Detail of bone. +B. Outline sketch of bone to illustrate characteristics examined. +Lateral anterior foramina (LF); fangs (F); "prediastemal" maxillary +teeth (MT); lateral flange (PLF); perpendicular lines, A-B and C-D, are +used to determine which tooth-socket occurs directly below the center of +the posterior foramen (the specimen illustrated is atypical because +there are three foramina present).] + + +Discussion and Conclusions + +Generic characteristics of _Tantilla_ are as follows: size small, body +slender, head not noticeably distinct from body; preoculars one; +postoculars one or two; nasals two; loreal absent; smooth dorsal scales +in 15 rows; anal plate usually divided but sometimes single; subcaudals +paired; usually two enlarged, posterior, grooved fangs on the maxilla +separated by a small diastema from the other maxillary teeth (Baird and +Girard, 1853:131; Cope, 1900:1110; Dunn, 1928:24; Blanchard, 1938:369; +Schmidt and Davis, 1941:268; Wright and Wright, 1957:722; Conant, +1958:180). Of these characteristics, only the following were invariable +in our sample: size small, body slender, head not noticeably distinct +from body; postoculars one or two; smooth dorsal scales; posterior +grooved fangs on maxilla. The dorsal scales, anal, and subcaudals were +"correct" 92 per cent of the time; the preoculars and nasals were +"correct" 97 per cent of the time. + +[Illustration: FIG. 5. Ventral view of right maxilla of _T. gracilis_ +(KU 83484), x 39. Actual length of bone is 2.44 mm. A. Detail of bone. +B. Outline sketch of bone to illustrate characteristics examined. +Lateral flange (PLF) forms an obtuse angle (ADC) with main axis of bone +(AB); a reference line (2), perpendicular to the main axis of the bone, +is used to locate the tooth-socket directly opposite the tip of the +suborbital process (SOP); the anterior and posterior edges of the +suborbital process are not parallel at any point (reference lines 1 and +3).] + +_Tantilla_ reportedly has two nasals (Baird and Girard, 1853:131; Cope, +1900:1110; Wright and Wright, 1957:722). _Tantilla gracilis_ usually has +a single nasal that is divided below the naris; variation in this +characteristic is discussed above. + +In addition to the generic characteristics, specific characteristics for +_T. gracilis_ are as follows: supralabials 5-8 (usually 6); supralabials +3 and 4 entering orbit; infralabials 5-7 (usually 6); temporals 1 + 1; +ventrals 115-138 in females, 106-132 in males; subcaudals 33-53 in +females, 40-57 in males; tail length 13-27 per cent total length in +females, 15-30 per cent in males (Baird and Girard, 1853:132; Cope, +1900:1111-12; Force, 1935:653-54; Taylor, 1936:337-38; Blanchard, +1938:371-72; Kirn, Burger, and Smith, 1949:240-49). Excepting the number +of temporals and the supralabials entering the orbit, the +characteristics of the specimens in our sample are within the ranges of +variation mentioned above. Of the 10 maxillary characteristics studied, +no variation was observed in number of fangs or angle of lateral flange. + +Because our data have some bearing on the problem of geographic +variation in _T. gracilis_ and the recognition of subspecies, we comment +briefly on the status of subspecies in _T. gracilis_. Kirn, Burger and +Smith (1949) proposed the recognition of two subspecies of _Tantilla +gracilis_ (_T. g. gracilis_ Baird and Girard and _T. g. hallowelli_ +Cope). These subspecies were diagnosed on the basis of differences in +ventrals, subcaudals, and the ratio of tail length to total length; +sexual dimorphism in each characteristic was considered. We do not +recognize these subspecies for the reasons given below. + +The data of Kirn _et al._ (1949) appear to be presented inaccurately in +part. Both of us independently recalculated the mean given for each +characteristic for each subspecies, using the data in figures 2-4 of +Kirn _et al._ (1949:242, 244-245). Of their 12 means presented (table 1, +p. 247) only two agree with our recalculated means, although the means +calculated by each of us independently are in complete agreement (Table +5). Also, we independently calculated the percentages of specimens of +each "subspecies" that are included in the ranges of variation given in +their diagnoses (Table 5); again, our independent calculations are in +complete agreement. In our opinion the differences between the +populations for the characteristics analyzed do not warrant recognition +of subspecies (Fig. 6). + +TABLE 5. Means of "Diagnostic" Characteristics of T. g. gracilis Baird +and Girard and T. g. hallowelli Cope. + + ============================================================================= + | _T. g. gracilis_ | _T. g. hallowelli_ + -----------------+-----------------------------+----------------------------- + | | |Per cent | | |Per cent + CHARACTERISTICS | | Mean |specimens | | Mean |specimens + | Mean | (Kirn, |of Kirn, | Mean | (Kirn, |of Kirn, + |(Kirn, |_et al._)|_et al._ | (Kirn, |_et al._)|_et al._ + |_et al._|recalcu- |included |_et al._ |recalcu- |included + | 1949) | lated |their |in 1949) | lated |in their + | | by us |diagnosis | | by us |diagnosis + -----------------+--------+---------+----------+---------+---------+--------- + Ventrals--females|125.67 | 125.71 | 69.4 | 130.07 | 130.07 | 79.6 + Ventrals--males |115.97 | 116.61 | 70.0 | 121.22 | 120.87 | 69.7 + | | | | | | + Caudals--females | 40.99 | 40.82 | 62.8 | 46.79 | 43.82 | 77.4 + Caudals--males | 47.75 | 48.29 | 78.3 | 51.67 | 50.29 | 51.0 + | | | | | | + Tail L./total L. | | | | | | + females | .1976| .1976| 69.0 | .2084| .2076| 74.2 + Tail L./total L. | | | | | | + males | .2336| .2362| 56.1 | .2477| .2423| 79.6 + ----------------------------------------------------------------------------- + +The data presented by Kirn _et al._ do not demonstrate intergradation +between two populations. Moreover, the diagnostic ranges of the number +of subcaudals in males of _hallowelli_ and the ratio of tail length to +total length in males of _gracilis_ do not include the recalculated +means for those characteristics. Furthermore, the means for the +following characteristics are dangerously close to being excluded from +their respective "diagnostic" ranges: in _hallowelli_, the number of +ventrals in males; in _gracilis_, the number of subcaudals in females, +and the ratio of tail length to total length in females. It is +incongruous that Kirn _et al._ state (p. 243) that "the coefficient of +geographic divergence is .6 per degree latitude" for ventrals, and on +the same page they state that "The average number of ventrals gradually +increases toward the north in _Tantilla gracilis_ except in southern +Oklahoma and central Arkansas (the area of intergradation between _T. g. +gracilis_ and _T. g. hallowelli_) where the change seems to be more +abrupt." The data presented in Kirn _et al._ (1949) do not demonstrate +an abrupt change. + +The present sample of _T. gracilis_ from Kansas is clearly within the +geographic range of _T. g. hallowelli_ as mapped by Kirn _et al._ +(1949:241). However, for the sample from Kansas, the mean number of +subcaudals of males is well within the diagnostic range for _T. g. +gracilis_ (Table 6), the mean number of subcaudals of females is closer +to the mean of _gracilis_ than it is to the mean of _hallowelli_ (Table +5), and the mean of the ratio of tail length to total length of both +sexes is within or very close to the diagnostic range for _gracilis_ +(Table 6). + +[Illustration: FIG. 6. Frequency polygons presenting ratios of tail +length to total length for males of _T. g. gracilis_ and _T. g. +hallowelli_ from Kirn, _et al._ (1949). The dotted vertical lines +represent the means given by Kirn, _et al._ (1949) for _gracilis_ (left) +and _hallowelli_ (right); the solid vertical lines represent the +recalculated means for _gracilis_ and _hallowelli_, respectively, using +the data of Kirn, _et al._ The hatched bars represent the range included +by Kirn, _et al._ (1949) in their diagnoses of _gracilis_ (left) and +_hallowelli_ (right).] + +TABLE 6. Comparison of the Means of Some Characteristics of the Sample +of _Tantilla gracilis_ from Kansas (Our Data) with Some of the +Diagnostic Characteristics (Kirn _et al._, 1949:240) of _Tantilla +gracilis gracilis_. + + ============================================================ + |Diagnostic range for | Mean of the sample + CHARACTERISTIC | _T. g. gracilis_ |("_T. g. hallowelli_") + | | from Kansas + ---------------+---------------------+---------------------- + No. of | | + subcaudals | 40-50 | 48.4 (N = 109) + males | | + | | + Tail L./ | | + Total L. | .160-.200 | .20 (N = 79) + females | | + | | + Tail L./ | | + Total L. | .200-.235 | .23 (N = 109) + males | | + ---------------+---------------------+---------------------- + +Smith and Sanders (1952:218) pointed out an error in the range of +subcaudals in female _T. g. gracilis_ as stated in table 1 (p. 247) of +Kirn _et al._ We add some additional corrections that should be made on +the same page. In table two, column two ("Sex"), the words "male" and +"female" are reversed for each characteristic. Also, in table two, +column four ("Means from data of Force"), the means for caudals should +be corrected from 33.61 to 43.61 in females and from 39.99 to 50.32 in +males, as based on our recalculations from Force's data; we did not +recalculate the means of Force's data for the other characteristics. + +The discussion above demonstrates that the number of subcaudals and the +ratio of tail length to total length are weak characteristics for +diagnosing _T. g. gracilis_ and _T. g. hallowelli_. The only remaining +diagnostic characteristic given by Kirn _et al._ (1949) is the number of +ventrals. Nevertheless, a step-cline in the number of ventrals, if one +exists, is yet to be demonstrated. Therefore, we completely agree with +Dowling (1957:32), who stated that "... the overlap in each character is +so great that any subspecific differences appear to be as yet undefined. +It is here suggested that _T. gracilis_ be retained as binomial...." + + +LITERATURE CITED + +BAIRD, S. F. and GIRARD, C. + + 1853. Catalogue of North American Reptiles in the museum of the + Smithsonian Institution. Part I.--Serpents. Smithsonian Misc. + Colls., 2:1-172. + +BLANCHARD, F. N. + + 1938. Snakes of the genus _Tantilla_ in the United States. Zool. + Ser. Field Mus. Nat. Hist., 20(28):369-376. + +CONANT, R. + + 1958. A field guide to reptiles and amphibians. Houghton Mifflin + Co., Boston, 366 pp. + +COPE, E. D. + + 1900. The crocodilians, lizards, and snakes of North America. U. S. + Nat. Mus. Rept. (1898), 153-1270. + +DOWLING, H. G. + + 1951. A proposed standard system of counting ventrals in snakes. + British J. Herpetology, 1(5):97-99. + + 1957. A review of the amphibians and reptiles of Arkansas. Occas. + Paps. Univ. Arkansas Mus., No. 3:1-51. + +DUNN, E. R. + + 1928. A tentative key and arrangement of the American genera of + Colubridae. Bull. Antivenin Inst. Amer., 2 (147):18-24. + +FORCE, E. R. + + 1935. A local study of the opisthoglyph snake _Tantilla gracilis_ + Baird and Girard. Papers Michigan Acad. Sci., Arts and + Letters, 20:645-659. + +KIRN, A. J., BURGER, W. L., and SMITH, H. M. + + 1949. The subspecies of _Tantilla gracilis_. Amer. Midl. Nat., + 42(1):238-251. + +PETERS, J. A. + + 1960. The snakes of the subfamily Dipsadinae. Misc. Pubis. Mus. + Zool., Univ. Michigan, No. 114:1-224. + + 1964. Dictionary of herpetology. Hafner Publ. Co., New York, 392 pp. + +SCHMIDT, K. P. and DAVIS, D. D. + + 1941. Field book of snakes. G. P. Putnam's Sons, New York, 365 pp. + +SMITH, H. M. and SANDERS, O. + + 1952. Distributional data on Texan amphibians and reptiles. Texas J. + Sci., 4(2):204-219. + +TAYLOR, E. H. + + 1936. Notes and comments on certain American and Mexican snakes of + the genus _Tantilla_, with descriptions of new species. Trans. + Kansas Acad. Sci., 39:335-348. + +WRIGHT, A. H. and WRIGHT, A. A. + + 1957. Handbook of snakes. Comstock Publ. Assoc, Cornell, Vol. II, + pp. 565-1105. + +_Department of Biology, The University of New Mexico, Albuquerque, +87106, and Department of Zoology, The University of Arizona, Tucson, +85721._ + +_Transmitted July 11, 1967._ + + * * * * * + +Transcriber's Notes + +The following corrections were made to the text: + + + + + + + +End of the Project Gutenberg EBook of Morphological Variation in a +Population of the Snake, Tantilla gracilis Baird and Girard, by Laurence M. Hardy and Charles J. 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