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Populational relationships in the corn snake Elaphe guttata (Reptilia: Serpentes).

ABSTRACT. -- A southeastern subspecies, Elaphe guttata meahllmorum subsp. nov., is distinct from a northwestern subspecies, E. g. emoryi, on the basis of having fewer dorsal blotches on body (44.5 or fewer, 100%, vs 45 or more, 98%, respectively), as well as less distinct differences in ventrals, caudals and ventral pattern. The former agrees with E. g. guttata in number of dorsal blotches on body, differing primarily in color from it (gray or brown vs reddish, respectively). E. g. emoryi agrees with E. g. meahllmorum in color, but differs from both of the other subspecies in having numerous blotches. The eastern, nominotypical subspecies appears to be dichopatric relative to the other subspecies, but if a contact occurs it is with E. g. meahllmorum, not with E. g. emoryi. The latter two subspecies have parapatric ranges and presumably intergrade, although conclusive evidence is lacking. One nominal species, Coluber maculatus Bonnaterre (1790), commonly referred to the synonymy of E. g. guttata, was found to be based partly also on Lampropeltis c. calligaster (Harlan, 1827). Since no type had previously been specified for C. maculatus, a lectotype is designated that maintains the current allocation of the name, thus avoiding replacement of Harlan's name were the Lampropeltis selected as lectotype.

RESUMEN. Una subespecie sureste, Elaphe guttata meahllmorum subsp. nov., se distingue de la subespecie noroeste, E. g. emoryi, por tener menos manchas dorsales sobre el cuerpo (44.5 o menos, 100%, vs 45 o m[sz]s, 98%, respectivamente), con tambien diferencias no tan prominente en el nomero de escamas ventrales y caudales, y en el patron ventral. La primera subespecie comporte con E. g. guttata el nomero de manchas dorsales sobre el cuerpo, pero se diferencia en color (gris o moreno vs rojizo, respectivamente). E. g. emoryi comporte con E. g. meahllmorum el color, pero se diferencia con ambos las otras en tener manchas m[sz]s numerosas. La subespecie nominotipical del sureste parece ser dicop[sz]trico con las otras subespecies, pero si contacto ocurre es con E. g. meahllmorum, no con E. g. emoryi. Las oltimas dos subespecies tienen distribuciones parap[sz]tricas y por presunci n son intergradiente, aunque no hay evidencia conclusiva. Una especie nominal, Coluber maculatus Bonnaterre (1790), antes referido a la sinonimia junior de E. g. guttata, fue descubrido fundida en parte sobre Lampropeltis c. calligaster (Harlan, 1827). Hasta ahora no tipo ha sido designado para C. maculatus; para evitar la posibilidad de asignar este nombre a la especie de Harlan, hacemos el lectotipo como un ejemplo de Bonnaterre que pertenece a la especie conocido como E. g. guttata, de cual C. maculatus definitivemente ahora es un sin nimo junior.


The field of herpetoculture, involving basic research on husbandry and propagation as well as sale of captive-produced animals, has become a multimillion dollar business (Collins, 1992). A species that has been extensively reproduced and that accounts for an important segment of commercial sales is the corn snake, Elaphe guttata. The nominate subspecies (E. g. guttata) has dominated the attention of herpetoculturists, and numerous color and pattern varieties have been produced by breeders. The western subspecies (great plains rat snake, E. g. emoryi), however, has also given rise to some unusual varieties (see below) and it is likely that herpetocultural attention will eventually expand to include this race. Anticipating this shift, we have reexamined the taxonomic status of E. g. emoryi, concluding that it should be split into two subspecies. This paper represents the evidence that leads to our conclusion together with an analysis of the nomenclatural history of the E. guttata complex.

Over 50 years ago, Woodbury and Woodbury (1942) differentiated a subspecies of snake, then known as Elaphe laeta (Baird and Girard, 1853), from the nominotypical and only other subspecies, E. l. laeta. They named it E. l. intermontanus (an incorrect ending; properly corrected to an -a ending by Dowling, 1951a) in reference to the isolated population, from which the type was chosen, occurring in central eastern Utah and adjacent Colorado, between the Wasatch and Rocky Mountain ranges. Comparisons were made primarily between the intermontane group and the populations of southern Texas, although specimens from Nebraska, Kansas and New Mexico that they considered intermediate between the nominotypical subspecies and E. l. intermontanus were also studied. Excluding those specimens, the Woodburys found a sharp distinction with very little (males) or no (females) overlap between the compared populations in numbers of ventrals and dorsal blotches.

Dowling (1951a) reported that the name Scotophis laetus of Baird and Girard (1853), erroneously applied by subsequent workers to the Great Plains Rat Snake, actually is a junior synonym of Coluber obsoletus Say (1823), now Elaphe obsoleta, and revived the next earliest name, Scotophis emoryi Baird and Girard (1853) for the species formerly known as E. laeta. Scotophis having been synonymized with Elaphe (Stejneger, 1907: 307), the proper nomenclatural combination became E. emoryi (Baird and Girard, 1853). Dowling recognized both E. e. emoryi and E. e. intermontana.

A year later, however, in a checklist of American members of Elaphe, Dowling (1952) lumped E. emoryi with the taxon described earlier (Linnaeus, 1758) as Coluber guttatus, retaining the former as a subspecies of the latter, E. guttata emoryi. With the latter he synonymized E. emoryi intermontana, without explanation. Such was his authority that without further documentation Woodbury and Woodbury's E. l. intermontana has been accepted as invalid by most authors ever since then; exceptions are Wright and Wright (1957), Maslin (1959), Miller (1961) and Weir (1993).

An unpublished master's degree dissertation (Thomas, 1974) did, however, document the untenability of E. l. intermontana, but at the same time refuted the validity of all other subspecies of E. guttata--E. g. rosacea (Cope, 1860), which had commonly been accepted as a valid species or subspecies since Barbour's (1920) resurrection of it, as well as E. g. emoryi. Duellman and Schwartz (1958: 298-300) had already rejected E. g. rosacea as a valid subspecies, and that arrangement, with but two subspecies of E. guttata, has almost universally been accepted since then, in spite of Thomas' (1974) conclusion.

The original intent was to determine whether the variation occurring in E. g. emoryi was based on artifacts of sampling, or significant geographic variation. To the authors' surprise, it was found that the separation envisioned by the Woodburys (1942) and reiterated by Weir (1993) was basically sound, even though the numbers of ventrals, caudals and tail blotches proved to be only marginally useful. The number of dorsal blotches on body, however, was found to vary in a geographically consistent, non-clinal pattern indicative of subspecific status of two groups of populations, although not exactly as conceived by the Woodburys or Weir. The isolated population of eastern Utah and western Colorado is not taxonomically distinct on the basis of morphological characters, limited to small size and usual dark gray background color--so dark in some that the dorsal blotches are difficult or impossible to count. Isozyme evidence would be useful in reconsideration of the isolate's taxonomic status.

Although an increase in number of dorsal blotches toward the northern part of the range of E. g. emoryi (sensu lato) has long been noted (e. g. Thomas, 1974; Conant and Collins, 1991: 196), and in some cases interpreted simply as a phenotypic effect of a temperature cline, in reality the race with few blotches (to the southeast) reaches nearly to the northernmost limit of the range of E. g. emoryi (s.l.), in southwestern Illinois, and the race with more numerous blotches (to the northwest, E. g. emoryi, sensu stricto) extends southward to about 29[degrees] N latitude, south of San Antonio, Texas (Fig. 1). Temperature effects are therefore essentially ruled out (see also statistical analyses reported below).


410 specimens were studied referable to E. g. emoryi (s.l.), representing all parts of its range except Louisiana, extreme eastern Texas and southern Arkansas. Range limits (Fig. 1) have been conceived on the bases of those specimens, Conant and Collins (1991) and Dundee and Rossman (1989). All specimens in the 13 acknowledged collections have been examined except for a few duplicates in UNM. Standard scutellational and mensural data were taken on 178 specimens randomly representative of the geographic range of the subspecies (s.l.) as a whole (Tables 3, 4). On the remaining 232 the only scutellational data taken were ventral and caudal counts (Tables 1, 2). Ventrals were counted by the Dowling (1951b) system.


Five sets of data were taken on the pattern of all 410 specimens (Tables 1, 2). The dorsal blotches on the body were counted beginning with the first blotch on the neck posterior to the two cranionuchal stripes, but because of the frequent occurrence of transversely Y-shaped blotches, two sides were counted separately and statistically analyzed for the mean figures for each specimen, resulting in some fractional counts (e.g., 35.5). Such aberrations were more numerous in populations with more numerous blotches (E. g. emoryi sensu stricto) than in those with fewer.

The number of dorsal blotches on the tail was also recorded for all specimens.

The number of ventrals with submedial spots appeared early in the study to be greater, especially on the anterior part of the venter, in E. g. emoryi (s.s.), hence two counts were made on all specimens to reflect that difference: (1) the number of ventrals with submedial dark spots among the first (anterior) 50 ventrals, and (2) the first (anterior) ventral bearing one or more submedial spots. Care had to be exercised to disregard the lateral blotches that involve the lateral ends of the ventrals; they are regularly present whereas the more medial spots, usually to one side of midline, are highly variable, occurring on anywhere from no ventrals at all to almost all of them. Evidently variation of the two sets of spots is under different genetic control.

In addition to the specimens personally examined, data on 60 specimens of E. g. guttata in the files originally created by Frank N. Blanchard were utilized. Those files were passed on to Howard K. Gloyd and ultimately were given to HMS.

All museum alphabetisms (they are not, as a rule, acronyms; Algeo and Algeo, 1991:9) are conformant with Leviton et al., 1980, insofar as given; others are: ASUMZ, Arkansas State University Museum of Zoology, State University, Arkansas; EAL, Ernest A. Liner, Houma, Louisiana; UTEP, University of Texas at El Paso; and WTAM, West Texas A & M University, Canyon, Texas.


No characters were found to provide acceptable diagnostic differentiation of the two subspecies except for the number of dorsal blotches on the body, although several other characters exhibited significant differences (see Variation and Comparisons for statistical analyses). The pattern of variation nevertheless is indicative of the validity of two subspecies in the populations formerly referred to E. g. emoryi. Since the race with fewer blotches has never received a distinctive name, the authors here call it Elaphe guttata meahllmorum, subsp. nov.

Holotype. UCM 46009, El Salto, San Luis Potosi, Mexico, June 13, 1964, T. Paul Maslin et al. (the UCM "Yucatan Expedition").

Paratypes. Forty-one; states (and counties in the United States) as follows (see list of Specimens Examined for more precise locality data). Arkansas: Cleburne (MPM 17000, 19100), Saline (ASUMZ 18304), Washington (UAM 68-735-1833). Illinois: Monroe (UIMNH 50963). Missouri: Taney (KU 19093). Texas: Cameron (UCM 15232, 15234, UTA 8404), Duval (UTA 17179), Jim Hogg (KU 174799, UTA 10439), Karnes (WTSU 8766), LaSalle (UTA 16780), McMullen (UTA 10428, 16662), Nueces (KU 61001), San Patricio (EAL 255), Starr (UTA 17784), Van Zandt (UTA 15809), Webb (KU 145867). Chihuahua: EAL 2535, UNM 3430. Coahuila: EAL 7, 2391, 2420, KU 47100, UTEP 6336. Nuevo Leon: EAL 669, 1187, 2603, KU 87743, UCM 47378, UO 33466. Queretaro: UTEP 9116. San Luis Potosi: EAL 323, KU 67652, UTA 4674. Tamaulipas: KU 61008-9, UCM 50118.

Definition/Diagnosis. A member of the E. guttata complex, having "parallel neck stripes crossing the parietals and normally uniting on the frontal" (Thomas, 1974:57); no ontogenetic change in pattern; a postorbital stripe extending onto or beyond infralabials; ventrals and subcaudals not exceeding 232 and 86 in males, 240 and 83 in females, respectively (Thomas, 1974:98); venter usually distinctly checkerboard-like in pattern, light areas between dark rectangles usually white, sharply contrasting with pigmented areas; usually two, largely continuous black stripes on subcaudal surface. Distinct from E. g. guttata in having dark gray, olive, or gray-brown dorsal blotches on a light gray background (vs reddish or orange blotches and ground color), and from E. g. emoryi in having 44.5 or fewer dorsal blotches on body (100%, vs 2%).

Description of holotype. A young adult female, s-v 517 mm, tail 113 mm, tail/total length ratio 0.1794. Specimen well preserved and in good condition except for a deep cut on right side of neck from midline between posterior genials to level of ventral 14, and a more superficial cut on left side between levels of ventrals 20 and 34.

Preoculars 1-1; postoculars 2-2; temporals 2-2-3 on both sides; supralabials 8-8; infralabials 12-12; scale rows 25-29-19; ventrals 223; subcaudals 71 (plus 2-4 missing at tip); anal divided; median dorsals weakly keeled, beginning with the vertebral scale row posterior to the anterior third of body (where all dorsals are smooth) and increasing within a span of 20 ventrals to the median eight rows; no keels on tail, others smooth or very faintly keeled; two apical pits on all dorsal scales of both body and tail, except erratic (present, absent or only one of the pair, usually the more medial one, present) on the first dorsal scale row.

Dark markings gray-brown, black-edged; a dark bar across internasals, extending onto supralabials and rostral; an interorbital dark bar continuous with a postocular dark bar ending on infralabials at rictus oris; a pair of longitudinal, short cranionuchal dark stripes beginning posteriorly at level of ventral five, uniting on frontal; right cranionuchal dark stripe narrowly connected with first blotch on trunk; a small, elongate, medial dark spot on posterior extremity of interparietal suture, extending posteriorly onto anterior three rows of nuchals; 35 dorsal dark blotches on body, 13 on tail; body blotches mostly about as broad as long, extending to 7th-9th-5th scale rows (anterior, middle, rear trunk regions), separated medially by spaces about 1.5 scale rows long; most of the anterior blotches with a transverse light area surrounding black-centered scales; 25th blotch more elongate than others and almost completely split by that transverse light area with black-centered scales (much like the 9th blotch shown in Fig. 2); lateral spots distinct, relatively large (0.25 size of dorsal blotches), mostly alternating with dorsal blotches, not reaching ventrals, but alternating or coinciding with small, more numerous sublateral spots (covering the equivalent of 2-5 dorsal scales) that encroach onto the ends of the ventrals.

First ventral with a median or paramedian dark spot, no. 17; 17 ventrals among the anterior 50 with one or more such spots; little scattered pigment on venter between spots; checkered ventral pattern moderately dense, approximately 2/3 of ventrals involved; paired subcaudal stripes continuous only on posterior 3/5 of tail.

Etymology. The subspecific name meahllmorum (pronounced as though spelled "meal/morum") is derived from the names of 11 associates of HMS to whom he is indebted particularly for absolutely vital aid in preparation for publication of volume seven of the series jointly authored by himself and his wife and co-worker, Rozella B. Smith, entitled A Synopsis of the Herpetofauna of Mexico. The new name is a collective, proposed in honor of the following individuals, now or formerly at the University of Colorado, listed with their designated letters in the order of occurrence in the name: m, Dr. Michael J. Preston, Department of English; e, Mary E. Marcotte, EPOB (Department of Environmental, Population and Organismic Biology) secretary; a, Ann E. Carrington, do.; h, Laura J. Heigl, do.; l, Linda K. Bowden, do.; l, Dr. William M Lewis, Chairman, EPOB; m, Dr. Michael D. Breed, EPOB, former Chairman; o, Phyllis A. O'Connell, EPOB secretary; r, Dean Charles R. Middleton, College of Arts and Sciences; u, Dr. Shi-Kuei Wu, Curator of Zoology, University of Colorado Museum; and m, Dr. Michael C. Grant, EPOB, former Chairman.

Variation and Comparisons. Discriminatory variation in E. g. emoryi and E. g. meahllmorum, based on our data on 410 specimens, and in E. g. guttata, based on data for 60 specimens in the Blanchard files, is summarized in Table 1. Statistical tests comparing E. g. emoryi and E. g. meahllmorum, the primary focus of this paper, are presented in Table 2.

Because the subspecies we call E. g. meahllmorum was not recognized in the past, the literature (Thomas, 1974; Raymond and Hardy, 1983; Dundee and Rossman, 1989) indicates that no significant differences in scalation exists between the subspecies of E. guttata, but with segregation of the present subspecies we have found them in both ventral and caudal counts. In a number of respects E. g. meahllmorum agrees with E. g. guttata, or is more or less intermediate between the latter and E. g. emoryi (s.s.); lumping E. g. meahllmorum and E. g. emoryi (s.s.) as E. g. emoryi (s.l.) thus tends to eliminate the scalational differences, leaving the reddish coloration of E. g. guttata as its sole distinction from the other subspecies.


The number of dorsal blotches on body remains the most reliable distinction between E. g. emoryi (s.s.) and the other two subspecies, the former having 45 or more in 98% of examined specimens, as opposed to 100% having fewer than 45 in the others. However, E. g. guttata has 35 or fewer in 86% of specimens examined, E. g. meahllmorum 35.5 or more in 78%. Likewise, 74% of E. g. emoryi have 18.5 or more dorsal caudal blotches, as opposed to 25% of E. g. meahllmorum and 0% of E. g. guttata. In the latter subspecies 88% have fewer than 15 caudal blotches, whereas in E. g. meahllmorum only 15% do, and of course only 2% of E. g. emoryi. Thus the number of dorsal blotches on body and tail serves as a rather strongly reliable or even infallible criterion for distinguishing all three subspecies from each other.

As shown in Figs. 2-6, considerble variation exists in the markings of both E. g. emoryi and E. g. meahllmorum. Fig. 2 illustrates the highest degree observed of secondary splitting of dorsal blotches, yielding a possible count of 47 in an animal basically with fewer, certainly no more than 44; because of this ambiguity its dorsal blotch count was not entered in the statistical analyses. Although several other specimens exhibited divided blotches that required separate count, no other was rejected for statistical analysis, because their totals did not exceed the standard limit (44.5) for the subspecies. The key to subspecies is adjusted to extremes exceeding that limit by incorporating notation of the split-blotch tendency.

Fig. 2 also illustrates an example of the extensive reduction in number of ventral spots that is common in E. g. meahllmorum, rare in E. g. emoryi. Such poorly defined ventral spots, and such abundant, scattered ventral pigment, are rare variants observed only in E. g. meahllmorum, although universal in sympatric E. obsoleta. The absence or great reduction of the subcaudal stripes occurs much more frequently in E. g. meahllmorum than in E. g. emoryi; complete absence was noted only in material from Texas and Mexico (37 in 129, 29%, higher in Mexico, with 20 in 58, 34%, than in Texas, 17 in 71, 24%) in the former, whereas in E. g. emoryi only 4% (11 in 248) had no subcaudal stripes, although the percentage was higher to the south (Texas 5 in 54, 9%, vs 6 in 194 from western Colorado (1), eastern Kansas (3) and northeastern New Mexico (2), 3%.


See Figs. 3-6 for examples and explanations of other pattern variants.

Most important is the fact that clinal variation in both body and tail blotches cannot account for these taxonomic differences (Table 2). Tests of this matter focussed exclusively on E. g. emoryi and E. g. meahllmorum. Data were pooled over taxa and regressed on longitude and latitude. Both geographical factors were associated with variation in body and tail blotches, but for neither character could clinal effects account for as much variation as was visible between the taxa (Table 2). In this analysis, longitude and latitude were treated as covariates with respect to taxa. Variation in, for example, body blotches was partitioned into the proportions associated with longitude (0.06) and latitude (0.46). When these values were substracted from the proportion of variance associated with taxa (0.69), the difference (0.17) was significantly greater than zero, implying differentiation (presumably from genetic sources) between taxa that cannot be accounted for by clinal effects. The same outcome arose from our analysis of tail blotches.



Ventrals and caudals also exhibit significant differences between E. g. emoryi (s.s.) and E. g. meahllmorum (see Tables 1 and 2). However, clinal variation is generally equal to or greater than the taxonomic variation. Therefore, we are reluctant to use these characters for diagnostic purposes. The same conclusion was reached for the last two characters in Tables 1 and 2.

Other variation is summarized in Tables 3 and 4. None of the variables listed here distinguish between E. g. emoryi and E. g. meahllmorum (all p's < 0.05).

Although it is clear that clinal effects cannot account for all of the variance between E. g. meahllmorum and E. g. emoryi in either body blotch counts or tail blotch counts, nevertheless it is acknowledged that substantial clinal variation exists in these and other characters. It therefore seems likely that the phenotypic clinal variance within the subspecies of E. guttata will be found to correlate either directly or indirectly with physiographic and climatological forces, by studies that apply the methods of Owen and Dixon (1989) or Ward et al. (1994) to geographic variation in E. guttata. Indeed, such efforts are encouraged by the authors because they will lead ultimately to a precise partitioning of phenotypic variation into direct environmental and genetic components, a result that is sorely needed in modern systematics, particularly in applications of the Evolutionary Species Concept.


Only one example of the striped variant of the species has been seen by the authors. It is a small (322 mm total length), female E. g. emoryi, captive bred, from Ft. Stockton, Pecos Co., Texas (KU 174798). It is anomalous also in having many ventrals divided medially and partially fused, to such an extent that ventral and caudal counts and other measurements were not recorded.

Key to the Subspecies of Elaphe guttata

1. Dorsal ground color and dorsal blotches reddish or pinkish; dorsal blotches on body fewer than 44.5 (100% ?); dorsal tail blotches fewer than 15 (88%); male ventrals more than 217 (67%); female ventrals more than 224 (69%); caudals fewer than 72 in males (75%) and females (96%); east of the Mississippi River........ E. g. guttata

Dorsal ground color gray; dorsal blotches gray, olive or gray-brown, 24-73 on body, 11-28.5 on tail; male ventrals fewer than 218 in at least 72%; female ventrals fewer than 225 in at least 73%; caudals fewer than 72 or not; west of the Mississippi River........ 2

2. Dorsal blotches on body 45 or more (98%), and none tending to be split transversely by a light streak; dorsal tail blotches 18.5 or more (74%); male ventrals fewer than 213 (72%); female ventrals fewer than 220 (71%); male caudals fewer than 72 (77%); female caudals 72 or more (75%)................................... E. g. emoryi

Dorsal blotches on body 44.5 or fewer (100%), unless some show evidence of a transverse split; dorsal tail blotches fewer than 18.5 (75%); male ventrals 213 or more (65%); female ventrals 220 or more (71%); male caudals 72 or more (76%); female caudals fewer than 72 (68%) ....................................... E. g. meahllmorum

Name allocations

In order to be certain that no name previously applied to E. guttata pertains to the subspecies here described, it has been necessary to examine all of them. They are here reviewed.

That the name Scotophis emoryi Baird and Girard (1853) is properly associated with the northwestern subspecies with numerous dorsal blotches is assured by (1) its type locality (Howard Springs, [approximately equal to]20 mi SW Ozona, Crockett Co, Texas, fide Dowling, 1951a: 43), which lies within, although near the edge of, the known boundaries of that subspecies; (2) color ("Ground color grayish ash," dorsal blotches "olivaceous brown"), and (3) pattern ("transverse quadrate blotches, 70 in number, the 50th opposite the anus. These are ten or 12 scales broad, two or three long, and separated by intervals of one to two scales. They are narrowly margined with black.") (quotations from the original description, p. 157). The type is lost (Dowling, loc. cit.), but the original description conclusively confirms the allocation here adopted.

Two junior synonyms of E. g. emoryi exist. The holotype of E. l. intermontanus Woodbury and Woodbury (1942), UU 271 from Moab, Utah, came from the western edge of the range of E. g. emoryi, and conforms in color with other material of that subspecies. It has fewer dorsal blotches (43, fide the original description) than average (51.2, Table 1), but 43 is within the known range of variation (39.5-73) in E. g. emoryi (s.s.). The dorsal blotch count was very likely not made the same way by the Woodburys as by us, and could be three or four fewer than a count made by our system. We were unable to borrow material from the UU collection.

The other junior synonym of E. g. emoryi is Coluber rhinomegas Cope (1860), a substitute for Scotophis calligaster: Kennicott (1859) (nec Harlan). Unfortunately neither Cope nor Kennicott noted the number of dorsal blotches, but the hologype is USNM 2259 (Cochran, 1961:166). Dr. Kevin de Queiroz very kindly examined it and reported (pers. comm.) its dorsal blotch count as 47, above the 44.5 upper limit for E. g. meahllmorum, and within the range of E. g. emoryi (s.s.). Our allocation of it to E. g. emoryi (s.s.) is based on the type locality. Hyatt, Anderson Co., Kansas, well within the known range of that subspecies, and upon variation of specimens examined from Anderson county and the surrounding area. Three specimens (KU 1057-8, 30072) from that county have 39.5, 47, 47 dorsal blotches on body, and seven from the next county east (Linn, KU 192203, 207170) and from four counties to the south (Bourbon, KU 192204; Montgomery, KU 154029, 154514; Wilson, KU 188691; Woodson, KU 170627) have 47-59.5 (mean 51.1) blotches on body. Therefore the authors regard allocation of C. rhinomegas to E. g. emoryi as conclusively confirmed.

The history of Kennicott's material from Hyatt, Anderson Co., Kansas--the type locality of both C. rhinomegas and Diadophis arnyi--is of some interest, inasmuch as no such place name is to be found in recent gazetteers or atlases, even in the extremely detailed U.S. Geological Survey 1:24,000 (7.5 X 7.5 minutes) topographic maps of Anderson county, although numerous names for places no longer in existence are given there. The possibility of error of locality citation for these names, perhaps involving ranges of adjacent subspecies, led us to seek expert help. It was forthcoming from Dorothy Kipper Licktieg of Greeley, Anderson Co., Kansas, president of the Anderson County Historical Society and author of a nearly completed book, the first of three, on the history of the county (Licktieg, 1993). She sent us a copy of a map showing Hyatt on the Missouri Pacific Railway about 3-4 miles southwest of Garnett. According to her information, Hyatt was a Massachusetts colony founded by Thaddeus Hyatt, who befriended and was tainted by association with the then infamous abolitionist John Brown, who in turn was active in eastern Kansas in the late 1850's. The colony existed from about 1856 to 1860, no more than three or four years, during which time it competed vigorously with Garnett for designation as county seat. Garnett won, and shortly thereafter the colony at Hyatt ceased to exist. Its leader in 1856 was W. F. M. Arny, of special interest since a "Mr. Samuel Arny" collected and sent to Kennicott, at some time before 1859, the specimens that became the holotypes of both C. rhinomegas Cope and Diadophis arnyi Kennicott. The latter is still recognized as D. punctatus arnyi.

According to Murphy (1972, reference and copies of pertinent pages courtesy Dorothy Licktieg), William Arny married Selina Craft in 1836, and they had three sons: Albert in 1837, Samuel in 1839, and William in 1846. Only the latter survived his parents, and Samuel was in his late teens when he sent the snakes to Kennicott that immortalized his family's name.

Actually, one other name, Elaphe quivira Burt (1946), has been applied to E. g. emoryi (s.s.), but it is a nomen nudum, existing in print only in the title of a presentation scheduled for the April 12, 1946, 78th annual meeting of the Kansas Academy of Science. As a frequent visitor to Kansas in those times, HMS learned that Burt intended to apply the name to one of the rare striped variants from Kansas of E. g. emoryi, but before doing so he realized what it was, so the description never appeared. It was undoubtedly a great disappointment to him, because he obviously had hoped initially to immortalize the name Quivira, which he had adopted for his biological supply company (Quivira Specialties Company, of Topeka, Kansas), although it would have been more for the Indian tribe of that name than for his company. As he stated (Burt, 1948), Quivira "is an old Indian name referring to villages or tribes of this Mid-West area as of some 400 to 500 years ago. Some of the ruins are said to have been found in central Kansas near Lyons and at other points and there are reports printed by the Nebraska State Historical Society indicating the probable existence of "Quivira" ruins in South-central Nebraska. By general usage Quivira, like Jayhawk, has become a synonym of Kansas; just as Cibola signifies New Mexico."

No other names previously applied to this species can be contrued as based upon E. g. meahllmorum, as reviewed in the following accounts. The holotype of Coluber guttatus Linnaeus (1766) is still in the Zoological Museum of Uppsala, Sweden, no. 147, according to Wallin (1992:226). Its original description, although brief, suffices to assure as correct the association long accepted for it and adopted most critically by Dowling (1952:2), who restricted the type locality, originally "Carolina," to Charleston, South Carolina.

An amazing eight names, aside from those now referred to E. g. emoryi (s.s.), were assigned by Boulenger (1894:39-40) to junior synonymy with Coluber guttata, and he assigned one other, Coluber rosaceus Cope (1888), to the synonymy of C. laetus.

Cope's C. rosaceus is the most familiar of all, because it was long recognized as a valid species or subspecies (of E. guttata) following Barbour (1920). Its holotype is USNM 14418, Key West, Monroe Co., Florida. It has seldom been recognized as valid since Duellman and Schwartz (1958:298-300) rejected it, although Staszko and Walls (1994: 70, 82-83) recognize it as a "weak" subspecies..

Cope (1888) also at the same time described a C. g. sellatus (syntypes USNM 6507 fide the original description {5507 fide Cochran, 1961: 165}, Palatka, Putnam Co., Florida, and USNM 9692, Arlington, Duval Co., Florida) that has never been recognized as valid. It was distinguished from C. g. guttatus on the basis of color and pattern features, as well as having 29 instead of 27 scale rows, all within the range of variation of sympatric E. g. guttata.

None of the other seven names applied to the species has ever been accepted as valid. The earliest, Coluber maculatus Bonnaterre (1790:19), was based in part on a specimen from Louisiana, and in part on Catesby's (1743) prelinnean description of the "corn snake" from Carolina, part of Linnaeus' hypodigm of C. guttatus. Location of the Louisiana type is unknown, although it may be in the Paris Museum. Its ventrals and caudals were given as 119 and 70, respectively; the former is no doubt a lapsus for 219. However, inasmuch as Bonnaterre explicitly attributed some (not all) ventral and caudal counts to other authorities (e. g. Crotalus horridus to Linnaeus and Tyson, C. dryinus to Linnaeus and Seba, and C. durissus to Linnaeus and Lacepede, two sets of counts for each), and must have done so for counts on other taxa with no explicit attribution but of which he apparently saw no specimens, it is concluded that the counts for his C. maculatus also were drawn from the literature, as noted, and do not in reality pertain to his specimen from Louisiana, because that specimen was described as having a whitish venter (no mention is made of any spotting), and the reddish, black-edged dorsal blotches as split along the midline, alternating on the two sides and fused to form a zigzag stripe. The reddish blotches are of regular occurrence in young L. calligaster (Harlan, 1827), and common in adults; and the unmarked venter and split dorsal blotches occur occasionally (Blanchard, 1921: 177-8) in that species, whereas they do not in any other species, including E. guttata, to which the Louisiana specimen might be referred. The ventrals and caudals of L. c. calligaster are 196-215 and 38-57 respectively (Blanchard, 1921: 117), much different from the supposed 219 and 70, respectively, given by Bonnaterre. As argued previously, those figures are not certainly applicable to Bonnaterre's Louisiana specimen. Because of the uncertainty of identity of Bonaterre's Louisiana specimen, and its possible representation of L. calligaster, we here designate, from Bonnaterre's hypodigm, the specimen on which Catesby (1743) based his description, not the specimen described by Bonnaterre, as lectotype of Coluber maculatus Bonnaterre (1790), in conformance with Art. 74 (c) of the Code. Thus that name definitively becomes a junior synonym of E. g. guttata (Linnaeus, 1766).

Eight years later Donndorff (1798: 206) named and described a Coluber compressus, without locality, based on a description and illustration in a 1790 work by Merrem (1790, fasc. 2: 49, pl. 11) in which no scientific names were given, although the vernacular "die zusammengedruckte Natter" was applied. Curiously, Daudin (1803: 318-321, pl. 77, fig. 2) also named the same "zusammengedruckte Natter" as Coluber pantherinus, reproducing Merrem's illustration and translating his description into French. Both the description and the figure are commensurate with assignment to E. g. guttata, of which both C. compressus Donndorff and C. pantherinus Daudin are incontestably junior synonyms. No type is known.

Shaw's (1802:460-1, 1 fig.) Coluber carolinianus was another name based on Catesby's prelinnean account, hence is clearly referable to E. g. guttata.

Coluber molossus Daudin (1803: 269-271) was based on reports by Bosc on specimens he observed in "Carolina." Presumably no types exist, although ventral and caudal counts are given on two specimens (220 and 60, 226 and 64), whether given by Bosc or taken on snakes seen by Daudin. Assignment to E. g. guttata is assured.

Coluber floridanus Harlan (1827: 360) was very briefly described, from "East Florida," but appears to be correctly referred to the synonymy of E. g. guttata. The type was stated in the original description to be in the Academy of Natural Sciences of Philadelphia, but it appears to have been lost since it is not entered in Malnate's list (1971) of the Academy types.

The remaining name, Elaphis rubriceps Dumeril, Bibron and Dumeril (1854: 270-2), was based on a specimen in the Paris Museum from an uncertain locality, thought by the authors to be in North America, probably New York. Its description agrees with variation known in E. g. guttata.

It is of interest to note that, as pointed out by Boulenger (1894: 40), Jan and Sordelli (1867: livr. 24, pl. 2) illustrated under the name Elaphis alleghaniensis (Holbrook, 1836) not only an Elaphe obsoleta (Say, 1823), of which Holbrook's name is a junior synonym, but also E. g. guttata.


Character-state dispersal in E. guttata leads the authors to hypothesize that E. g. meahllmorum is ancestral to the other two subspecies. In the course of evolution E. g. guttata retained the ancestral large and few dorsal blotches, but became reddish, whereas E. g. emoryi retained the ancestral gray color but experienced a marked increase in number of dorsal blotches.

All three subspecies of E. guttata now have isolated, northern relictual populations, none of which have evolved taxonomic distinction. As many as five dichopatric populations of E. g. guttata exist north of the main body of the subspecies' population (Conant and Collins, 1991: map 167). One widely disjunct population of E. g. emoryi exists in central eastern Utah and central western Colorado (Fig. 1). At least one and possibly two apparently disjunct populations of E. g. meahllmorum exist northeast of its main body, in (1) eastern Texas, western Louisiana and southern Arkansas, and (2) eastern Oklahoma, northwestern Arkansas and southern Missouri.

These several relictual populations collectively suggest a wider distribution of the species to the north in the past, during a hypsithermal period, and a partial withdrawal during the current hypothermal period, occurring not long enough ago to have permitted subsequent taxonomic differentiation.

Roth et al. (1989) hypothesized that the isolated population of Heterodon n. nasicus in northwestern Colorado arrived there via the Wyoming Corridor between the northern and southern Rocky Mountains during the early Holocene Climatic Optimum some 8500-5000 years ago. Probably the West Slope population of E. g. emoryi arrived there by the same route at the same time, when the entire species was more widely distributed than now. Bury's (1983) record from extreme northeastern Utah provides support for the hypothesis of northern, not southern, derivation of the West Slope population.

Both E. g. emoryi and E. g. meahllmorum are conspicuously partial, throughout most if not all of their ranges, to the vicinity of water, either lentic or lotic, and are not to be found distant from it. This strong affinity could well have been an isolating factor in some historic stage of xeric hypsithermy, affecting the relationships of all three currently recognized subspecies.

The exclusively northern relic populations suggest a late Holocene hypothermal period as being responsible for the range retraction that all three subspecies now exhibit. The southward confinement of E. g. guttata encountered no obstacle in the form of differentiated populations of its own species, unless the Keys population was then in existence. On the contrary, a southward retreat of E. g. emoryi did encounter E. g. meahllmorum, and the extensive interdigitation of the ranges of these two taxa, and the apparent isolation of two northern enclaves from the body of the latter subspecies, are very likely the result of that retreat.

Those phenomena also suggest that the two western subspecies evolved in dichopatry, E. g. emoryi to the northwest, E. g. meahllmorum retaining its position to the southeast. Their response to each other in renewed contact is typical of secondary subspecies.


No intergradation between E. g. guttata and its nearest relative, E. g. meahllmorum, is known to take place at the present time (Dundee and Rossman, 1989), although certainly, as shown by their pl. 16, the latter subspecies in Louisiana is distinctly less gray than elsewhere. A relatively recent contact is indicated, but none appears to exist now.

On the other hand, intergradation almost certainly occurs between the two western subspecies in scattered areas of contact, but their integrity is surprisingly well maintained in other areas. For instance, in El Paso Co., Texas, the dorsal body blotch counts of 39.5, 40, 40.5, 42 and 54 occur; all are readily assignable to one subspecies or the other, the latter to E. g. emoryi. On that basis, however, occurrence of E. g. emoryi in adjacent Chihuahua, not now known, is to be expected. Indeed, the northernmost specimen from Chihuahua (BYU 13918, Casas Grandes) has the highest number (44.5) of dorsal blotches of all 57 specimens seen from Mexico, except for two (KU 67653, 37 mi S Matehuala, San Luis Potosi, with 46.5 blotches; and UTA 4856, 10 mi S Cd. Mante, Tamaulipas, with 44.5 blotches), some of whose blotches are secondarily divided transversely; they are consequently not comparable with others without such secondarily subdivided blotches.

In 29 specimens from New Mexico, all have 45.5-65 dorsal blotches except for three. One, with 42 blotches (UNM 7501, 1.5 mi SW Pilar, Taos Co, Hy 64) was rejected as a probable released or escaped pet, inasmuch as all others from nearby areas had typically numerous blotches. Inasmuch as this species is extremely popular as a pet, a number of distributional may be due to escapes or releases in exotic (from original sources) localities. Another New Mexico specimen, with 43 blotches (UNM 13195, Bottomless Lakes, Chaves Co.), may similarly have been imported, because six other specimens from the same or adjacent localities in Chaves Co. had 46.5-56.5 (mean 50.25) blotches. The specimen from New Mexico referred to E. g. meahllmorum has 44 dorsal blotches, and is from Del Rio Ditch, 0.2 mi E Hy 28, Dona Ana Co. (NMSU 3136). Others from the same county (one from the same locality as NMSU 3136) have 48, 48.5 and 50 dorsal blotches, hence are clearly referable to E. g. emoryi. Since both subspecies occur in nearby El Paso Co., however, both may well also occur in adjacent Dona Ana Co. On the other hand, this general area may be one of intergradation; sufficient series are not yet available to resolve the question definitively.

Another area of possible intergradation is the vicinity of the Davis Mts. in Jeff Davis and Reeves counties, Texas, where counts of 43, 45.5, 48.5, 53 and 53 are available; the first is assigned to E. g. meahllmorum, the others to E. g. emoryi. The two counts from Pecos Co. (45.5, 47.5) are clearly exemplary of the latter subspecies, as is the count of 50 for the holotype of the subspecies from nearby Crockett Co. (Baird and Girard, 1853: 157), and the count of 46.5 for one from Tom Green Co. (KU 81987, 4 mi NW Christoval). However, one from Coke Co. (KU 81984, 6 mi N Water Valley) has but 42 blotches, and one from Irion Co. (KU 81985, 2 mi W Mertzon) has 44.5; these hence fall with E. g. meahllmorum, ranging northward from the Val Verde-Terrell counties area, where the counts are 36, 36.5, 37, 37.5, 39, 41, 42, 43, and 44 (mean 39.6). Even there, one count of 47 occurs (UCM 56667, 6.7 mi N Comstock), but it in turn fits geographically with the large southward extension of E. g. emoryi that circumscribes the Edwards Plateau (Bandera Co., 50; Bastrop, 46; Brown, 49; Callahan, 45; Coleman, 47; Frio, 49.5; Taylor, 47.5). These counts are too consistently referable to E. g. emoryi, without exception, to regard them as meaningless or intergrades.

A Brown Co. count of 44 (UTA 16759, Hy 377 100 m N Hy 1176) is consistent geographically with the extensive distribution of E. g. meahllmorum in eastern Texas, the nearest records being for Eastland Co. (43, 43.5) and Erath Co. (44, 44.5).

One other from Eastland Co. (KU 1139, nr Cisco) has 45 blotches and is assigned to E. g. emoryi; the other two (KU 1054, 1809), here assigned to E. g. meahllmorum, are also from Cisco (not "near").

To the northwest, the Stonewall (42), Knox (44) and Throckmorton (44) county records justify extension for E. g. meahllmorum into that area; one E. g. emoryi with 52.5 dorsal body blotches (UTA 482, 20 mi S Throckmorton) is, however, from the same locality as the specimen referred to the other subspecies. Intergradation in that area may well occur.

There are only three other specimens on the basis of which the range of E. g. meahllmorum is depicted on our map to extend into northern central Texas. One is from Wise Co., with 44 blotches (UTA 16338, LBJ Natural Grassland); others from that county, including another from the same locality with 62 blotches, have 52.5 (2) and 57.5 blotches. The other two, with 44 and 44.5 blotches, are from Dallas Co. (UTA 26462, S Cedar Hill, jct Texas Plume Rd & Lakeridge Rd; UTA 30226, Texas Plume Rd, 0.6 mi W Anderson Rd); others from the same and adjacent counties (Hood, Johnson, Palo Pinto, Parker, Shackelford, Tarrant) all have E. g. emoryi counts (45.5, 46, 47.5(2), 48, 48.5(3), 49(2), 50, 50.5, 51.5, 52, 52.5, 53, 56.5, 59; mean 50.2).

In Oklahoma, the only unexpected record is of an E. g. emoryi with 48.5 blotches from Le Flore Co. (UO field 3212; Holson Valley Rd, 0.5 mi S Cedar Lake), whereas others from that county have 35.5, 36, 39, 40 blotches, consistent with E. g. meahllmorum. The exceptional specimen may, or may not, be an import; intergradation is not evident. It is of interest that the highest blotch count ever recorded for the species (73) occurs on a specimen from Cimarron Co., Oklahoma (UO 33993, 1.5 mi E Kenton).

In Missouri, four specimens of E. g. emoryi are from surprisingly far south and east: KU 81980-1, McDonald Co., Noel, with 50.5 and 54 blotches; KU 81983, Taney Co., Kissee Mills, with 48.5 blotches; and UCM uncat., Crawford Co., 6 rd mi E Steelville, with 48 blotches. Yet KU 19093 from 1 mi N Branson, Taney Co., has 39.5. The two taxa may be sympatric in these areas.

The preceding analysis leaves no E. g. meahllmorum with 45 or more dorsal blotches on the body. On the contrary, 251 of 257 E. g. emoryi (98%) have 45 or more. Of the six exceptions, one is from Nebraska (KU 52221, Jefferson Co., 4 mi S, 0.5 mi W Reynolds, with 44), one from New Mexico (UNM 13195, Chaves Co., Bottomless Lakes State Park, with 43), one from Texas (WTAM 6078, Potter Co., Lake Meredith, with 43), and the other three from Kansas (KU 1058, Anderson Co., with 39.5; KU 206488, Greenwood Co., T28S, R12E, Sec. 3, with 43; and KU 158009, Marion Co., Marion Co. State Lake, with 43.5). How many, if any, of these exceptions may be due to importation is conjectural.

The over-all picture of the contacts between E. g. emoryi and E. g. meahllmorum is of possible sympatry in some areas (especially in Missouri, and possibly in Oklahoma), parapatry in others, and probable intergradation in others. More material will be required to establish definitively what the interaction is in any given area. Extensive range interdigitation in Texas is clearly evident, but to what extent, if any, it is artifactual remains to be clarified. It will be of much interest to determine whether the area where the species appears to be absent in southeastern coastal Texas (see Fig. 1) contacts the range of E. g. emoryi, thus isolating the population of E. g. meahllmorum in eastern Texas, western Louisiana and southern central Arkansas. Also much to be desired is confirmation of the isolation of that subspecies in southeastern Oklahoma, northwestern Arkansas and southern Missouri. That the three isolated enclaves of E. g. meahllmorum may differ in their interaction with E. g. emoryi, where their ranges are in contact, is not likely.

Taxonomic Ranks

Because E. g. guttata, as the authors understand it, now has no contact with the other two subspecies, its taxonomic rank is not established by the Biological Species Concept, which is limited in application to sympatric and possibly also to parapatric populations. The Evolutionary Species Concept does apply in such cases of allopatry, however, although with considerable subjectivity in determining whether or not an allopatric population in sufficiently differentiated to be regarded as having established a unique evolutionary trajectory. In the case of E. g. guttata, its reddish coloration is indeed distinctive, at least in degree, as compared with the other subspecies, and possibly the distinction may only become firmly established and even significantly supplemented in the future, but the subspecies' similarity to and apparent origin from adjacent E. g. meahllmorum in Louisiana (see especially Dundee and Rossman, 1989: pl. 16) convinces the authors that under the Evolutionary Species Concept E. g. guttata is best retained at a subspecific rank. The descriptions of Louisiana specimens (Raymond and Hardy, 1983) of what the authors now know to be E. g. meahllmorum, with pink in their coloration, bespeaks incomplete differentiation. So also does the occurrence in E. g. guttata (Schnitzler, 1990: 161) of the same, curious, light transverse areas across the dorsal body blotches as occurs regularly in southeastern E. g. meahllmorum, often splitting the blotches. There are too many shared peculiarities of these two populations, especially where their ranges are approximated, to regard them as safely launched on separate evolutionary tangents.

That they may be regarded as secondary subspecies, as suggested by Raymond and Hardy (1983), however, is a tenable hypothesis, perhaps presupposing an even more extensive range retraction having occurred at some time in the past than at present.

Despite the considerable evidence of possible sympatry of the other two subspecies in some (especially northeastern) areas of contact, the high probability of interbreeding (hence intergradation) elsewhere likewise imposes a subspecific, not specific, ranking as the most parsimonious assumption at the present time. Only with explicit evidence of sympatry, inviolate parapatry or dichopatry throughout the areas of juxtaposition of ranges, with or without hybridization, would assumption of specific rank be justified. As argued elsewhere, these two subspecies, as well as E. g. guttata, appear to be secondary.

The apparent phylogeny of these three taxa is also more harmonious with the concept of one species with three subspecies than with that of two or three separate species.

Descriptions in the literature of southern Florida populations frequently referred in the past to E. g. rosacea suggest that a reappraisal is in order, particularly in view of the separation of E. g. meahllmorum.


(counties in italics)

1. Elaphe guttata emoryi (252)

COLORADO (UCM; 38). Baca: Pritchett (56055); SE Pritchett (14968); SW Pritchett (52057). Bent: John Martin Reservoir (15233). Delta: Delta (field 4950); Eckert (56187, 56189, 56193-4, 56200-1, 56219, field 4951); 2 mi E Fruit Growers' Res. (56214); Hart's Basin (field 4944, 4947, 4949); Orchard City (56566, field 4999); Redlands Mesa (56212). Garfield: Rifle (19712). Las Animas: Cottonwood Creek (7643); SW Pritchett (9734). Mesa: Colorado National Monument (18108); Fruita Canyon (23948); Gateway (6700); Grand Junction (18109, 314689); Molina (56215). Montrose: Escalante Canyon (50713); Nucla (10700). Otero: La Junta (51876). Prowers: Lamar (1226). Pueblo: Lime (31466); Pueblo (2669-70, 31467).

KANSAS (KU except as noted; 103). No county: Flint Hills (OMNH 34313). Anderson (1057-8): 4 mi N Garnett (30072). Barber: 1 mi W Aetna (20005); 15.2 km W Medicine Lodge (204056); 5 mi S Sun City (19200); 7 mi S Sun City (18112). Bourbon: 6.4 mi W, 0.6 mi S Garland (192204). Butler: T25S, R6E, Sec. 25 (~2 mi NE El Dorado; 206489); T27S, R6E, Sec. 33 (~3 mi S Leon; 192411). Chase (188697): 17.6 km SE Bazaar (188690); 6.4 km E Cottonwood Falls (188698); 1.6 km S Saffordville (188694); 8 km S Saffordville (188686-9, 188693, 188695); 8 mi S Saffordville (75165); 3.2 km NE Strong City (188696). Clark: Clark Co. State Lake (21388, 207290). Clay: T10S, R4E, Sec. 29 (Milford Lake area; 218764). Cloud: 4.8 km N Glasco (206308); 8 km S Jamestown (207168). Comanche (1059): 4 mi W Aetna (18156). Cowley: 35.5 mi W Cedarvale, Hy 160 (176712); T35S, R5E, Sec. 20 (~Silverdale; 206173); Winfield (29983). Dickinson: T3S, R4E, Sec. 8 (~SE corner; 192106). Douglas (1073): Apponoose Creek (1063); Clinton (BYU 9138); 4 mi S Clinton (188701); 7.5 mi NE Lawrence courthouse (28679); Washington Creek (1077-8). Elk: 6.4 km S Fall River (193220). Ellis: Hays (16554-5); 4 mi E, 2 mi N Hays (159779). Ellsworth: 9.6 km E Lorraine (179026). Ford: 3.2 km S Hodgeman Co. line, Hy 283 (182289). Franklin (553778). Geary: Hy 177 5 mi S I70 (159778); Hy 177 8 mi S I70 (171162); T13S, R8E, Sec. 10 (~5 mi N Dwight; 159777). Gove: T15S, R26W, Sec. 12 (SE corner; 216123). Graham: 5.6 km S Bogue (192201-2). Greenwood: T28S, R12E, Sec. 3 (~1/2 way between Severy and Fall River; 206488). Hodgeman: Hodgeman Co. State Lake (182289); 14.4 km SW Jetmore (206309). Jewell: Jewell Co. State Lake (158019). Lane: T18S, R27W, Sec. 2 (~Alamota; 217222). Leavenworth: 7 mi NE Lawrence (2425). Lincoln: Wilson dam (174553). Linn: Cadmus (192203); 6.4 km SE Centerville (207170). Logan: T14S, R33W, Sec. 2 (~10 mi NW Elkader; 217224). Lyon: ESU Ross Nat. Hist. Reservation (192376). Marion: Marion Co. State Lake (158009). Marshall: T4S, R7E, Sec. 16 (~4 mi N Blue Rapids; 207291). Meade: SE corner (22842). Mitchell: 5.6 km WSW Beloit (206310). Montgomery: 4 mi W Sycamore (154029, 154514). Morris: 16 km S Council Grove (188692), Ness: T16S, R22W, Sec. 33 (~Brownell; 216189), Norton: T4S, R21W, Sec. 35 (~5 mi N Densmore; 217223). Osage: 0.8 km N Melvern (188699); Quenemo (1053). Osborne: T10S, R15W, Sec. 18 (SW corner; 216122). Ottawa: NW corner (188700); SW Ada (17033); 8 km W Delphos (207169). Pottawatomie: 6.5 mi N, 1.5 mi E Olsburg (49361). Republic: 8 km S Belleville (206311, 207167). Riley (55376): Hy 177 just N 170 (207189); 0.25 m W end Tuttle Creek dam (49360); Winkler (17094). Russell: 17.6 km SW Lucas (188702). Saline: T14S, R5W, Sec. 5, NNW Brookville (UTA 10934); 1.6 km S, 1.6 km E Salina (1823189). Shawnee: T12S, R14E, Sec. 10 (~3 mi SE Valencia; 207289). Smith: T5S, R15W, Sec. 17 (~Claudell; 203310). Trego (1472,2426-8): Cedar Bluffs Lake (191951); 1.6 km E Cedar Bluff dam (182320). Wabaunsee: 1 mi SW Alma (159824). Wilson: 4.8 km E Buffalo (188691). Woodson: Toronto Point, Toronto Lake (170627).

MISSOURI (KU except as noted; 8). Atchison: 1 mi N Rockport (81976). Bates: Drexel (81982). Cass: 0.5 mi NE Drexel (81978). Crawford: 6 rd mi E Steelville (UMC uncat.). Hickory: 5 mi SE Cross Timbers (81979). Jackson: 4 mi S Blue Springs (81975). McDonald: Noel (81980-1). Taney: 6 mi NE Kissee Mills (81983).

NEBRASKA (KU;2). Gage: Barneston (52222). Jefferson: 4 mi S, 0.5 mi W Reynolds (52221).

NEW MEXICO (UNM except as noted; 27). Bernalillo: 2 mi N San Antonito, Hy 14 (36910); Embudo Canyon, 10 mi E Albuquerque (4745); Hy 14 1 mi N I40, Sandia Mts. (32896); Pino Canyon, Sandia Mts. (415); 1 mi W Tijeras post office (48749); nr Tijeras Canyon, La Mura Rd (36574). Chaves: Bottomless Lakes State Park (13195, 16467, 17227); E side Mirror Lake, Bottomless Lakes State Park (15055, 15129); 7 mi E Roswell, Hy 380 (14589); Hy 380 at Rio Hondo, E Roswell (36943). Dona Ana: Del Rio Ditch 0.2 mi E Hy 28 (NMSU 3135); East Picacho School, Las Cruces (NMSU 2740); 3 mi NW Las Cruces, Hy 292 (NMSU 4959). Eddy: Black River, 4 mi NE Black River Village (43669); T25S, R24E, Sec. 35 (~7 mi SW Whites City; 54828); Cottonwood Spring, Cottonwood Draw, T25S, R25E, Sec. 36, SW 1/4 (~5 mi S Whites City; 51986); Dark Canyon Rd, T23S, R23E, Sec. 14, 4100' (49056); Hy 31, 0.5 mi W Pecos River, nr Carlsbad (19372). Guadelupe: Puerto de Luna, Hy 3 (36909). Quay: Logan, 3800' (31664). Sandoval: Placitas Fish Hatchery, N Sandia Mts (18275). San Miguel: 9.5 mi W Tucumcari, Hy (16629). Santa Fe: 2 mi S Madrid, Hy 14 (48072). Union: Dry Cimarron River, Folsom Falls (52022).

OKLAHOMA (UO except as noted; 20). Alfalfa: 7 mi N Jet (25291). Beaver: nr Gate (13742). Blaine: 20 mi SW Okeene (8454); 4 mi N Watonga (29266). Cimarron: 1.5 mi E Kenton (33993). Comanche: 0.5 mi SW Mt Quanah, Ft. Sill Military Res. (field 2295); Lake Lawtonka (12901); E Lake Lawtonka (12900); W Cache Creek (8137, 8213); Wichita Mt. Game Refuge (26889). Garvin: Pauls Valley (8860). Greer: 1.7 mi S Salt Fork Red River (25825). Harmon: NW Reed (29278). Kay: 3 mi N, 5 mi E Ponca City (25976). Kiowa: 10 mi NW Meers (13187). Le Flore: 0.5 mi S Cedar Lake, Holson Valley Rd (field 3212). Major: E Glass Mts (22890). Roger Mills (26438). Seminole: Bowlegs (10289).

TEXAS (UTA except as noted; 53). Bandera: Hy P 37 0.5 mi NE Hy 1283, 13 air km W San Geronimo, 350 m (UTEP 8988). Bastrop: 100 m S of Hy 969, 2.4 km SE Hy 1704 (15810). Brown: Hy 279, Brownwood State Park (WTAM field DC 92-R22). Callahan: I20, 1.5 m E Hy 2228 (5261). Coleman: 10 mi SE Santa Anna, Hy 1176 (5064). Cooke: Gainsville (KU 13726). Dallas: Duncanville, Greenhills Environmental Center (14097); E Mt. Creek Lake (2126); Plume Rd at Lakeridge Rd, S Cedar Hill (26463). Denton: Keller (22271). Eastland: nr Cisco (KU 1139). El Paso: 1 mi E Clint, exit to I10 (UTEP 124). Foard: 5 mi N Crowell, Hy 6 (9295). Frio: 8 km W Derby (22272). Hall: 2 mi SW Klondike (30678). Hansford: Gibbner Ranch, 10 mi N Spearman, Hy 760 (WTAM 5004). Hardeman: 3 mi N Copper Breaks State Park, Hy 6 (9293). Hood: Benbrook-Aledo Rd at Hy 2376 (11148); Hy 171 at Hy 377 (14228); Hy 179, 2.5 km NW Hy 377 (11149). Hutchinson: 2 mi N Stinnett (WTAM 5979). Jeff Davis: 41.6 km N Ft. Davis (KU 187741); Limpia Canyon, 5 mi NNW Ft. Davis (KU 45355). Johnson: I35, 0.6 mi SE Hy 917, 7.9 mi SE Tarrant Co. line (2614). Lubbock (5535). Palo Pinto: 7 mi W Graford (30680); 4 mi N Palo Pinto (2072). Parker: Aledo (266); 1 mi E Aledo (2125); 8.2 mi SE Weatherford (EAL 2317). Pecos (UTEP 14715): Ft. Stockton (KU 174798); 20 mi E Ft. Stockton (KU 81986). Potter: 6 mi W Amarillo (2058); Lake Meredith (WTAM 6078). Randall: 0.75 mi E Canyon, Hy 217 (WTAM 8379). Reeves: Hy 17, 4.2 km S Hy 290 (28891); 1 mi S Toyahvale, Hy 17 (UTEP 6119). Shackelford: 6.6 mi N Albany, Hy 293 (2057); 16 km W Albany, Hy 351 (14075). Tarrant: Benbrook-Aledo Rd (11368); Hy 1187, 1 mi W Hy 1902 (2912); Mary's Creek, 200 yds W jct Ridgmar Blvd and Dakae St. (1156). Taylor: Camp Berkeley (BYU 6060). Throckmorton: 20 mi S Throckmorton (UTA 482). Tom Green: 4 mi NW Cristoval (KU 81987). Val Verde: 6.7 mi N Comstock (UMC 56667). Wise (30677): 4 mi NNW Decatur (8639); 6.4-8 km N Decatur, off Hy 287 (15312); LBJ National Grassland (16337). Young: 1.3 mi E Graham, Hy 380 (9437).

UTAH (1). Grand: Moab (BYU 18896).

2. Elaphe guttata meahllmorum (157)

ARKANSAS (12). Cleburne: Big Creek Natural Area MPM (19100); Big Creek, 5 mi N Pickens (MPM 17000); W Pangburn (MPM 15703). Garland: Montgomery Co. line (ASUMZ 6221). Marion: Promised Land Ridge, Bull Shoals Lake (MPM 20306). Montgomery: 3 mi W Albert Pike, Hy 106 (ASUMZ 18478). Polk: Ouachita National Forest, Hy 88, 19 km E Oklahoma border (UTA 12717). Pulaski: Murray Park, Little Rock (ASUMZ 7823). Saline: Rattlesnake Mt. (ASUMZ 18304). Washington: Devil's Den State Park, 6 mi WNW Winslow (ARK 68-735-1833). White (MPM 18641).

ILLINOIS (2). Monroe: Bluff Rd, 7 mi N Valmeyer (UIMNH 50963); Fults (UIMNH 50837).

MISSOURI (KU except as noted; 4). Cedar: El Dorado Springs (81977). Greene: Hy 123 W Harold (UNM 10089). McDonald: 2.4 mi NE Noel (30030). Taney: 1 mi N Branson (19093).

NEW MEXICO (1). Dona Ana: Del Rio Ditch, 0.2 mi E Hy 28 (NMSU 3136).

OKLAHOMA (UTA except as noted; 6). Delaware: 8 mi S, 1.4 km W Jay (KU 154481). Le Flore: Cedar Lake, Holson Valley Rd (OMNH field 3215); Ouachita National Forest, Ark. Hy 1 (22270); Rich Mt. Crest, Hy 1, 13 km W Arkansas line (26118); 3.2 km S Zoe, Hy 259, 3.2 km N Hy 1 (10933). McCurtain: Hy 259, 0.6 km S Boktukolo Creek, 15.3 km S Le Flore Co. line (26461).

TEXAS (UTA except as noted; 76). Anderson: Hy 860, 0.6 km N Hy 2330 (16928); Pert, Hy 155, 5.5 km S Hy 2267 (16807). Aransas: 2.8 mi N Lamar (EAL 512). Atascosa: Benton (KU 8438). Bexar: San Antonio (KU 13779). Brazos: 5 mi NE College Station, Elmo Weedon Rd, 1 mi N Hy 30 (UTEP 10486); Hy 30, 8.2 mi E Hy 168 (UCM 41824). Brown: Hy 377, 100 m N Hy 1176 (16759). Cameron: Bayview, General Store Hy 2480, 0.6 km S Hy 510 (26699); Bayview, Hy 2480, 100 m S Hy 510 (26698); Brownsville (UCM 15234); Hy 77, 2 mi NW El Camino Real (8389); Los Fresnos (UCM 15232); Los Fresnos, downtown, Hy 1847 (8391, 8402); Hy 510, 1 mi W Hy 1847 (8404). Coke: 6 mi N Water Valley (KU 81984). Dallas: Texas Plume Rd, 0.6 mi W Anderson Rd (30226); Texas Plume Rd and Lakeridge Rd, S Cedar Hill (26462). Duval: Hy 16, 13.2 km S McMullen Co. line (17179). Eastland: Cisco (KU 1054, 1809). El Paso: Auburn Drive, Ascarate Park area, El Paso (UTEP 13784); 416 Lisbon St., SE El Paso (UTEP 9989); Hys 20 and 793, 4.8 mi SE Fabens (UTEP 12339); Upper Valley nr Canutillo, Gato Rd off Doniphan St. (UTEP 3562). Erath: Cedar Point, 1.6 km W jct Hys 2157, 3106 (14731-2). Irion: 2 mi W Mertzon (KU 81985). Jeff Davis: 5 mi S Toyahvale, Hy 17 (UNM 35798). Jim Hogg: 6.4 km SW Hebbronville (KU 174799); 18.8 mi SW Hebbronville (10439). Jim Wells: 14 mi S Alice, Hys 141, 281 (UTEP 5955). Karnes: 5.6 mi N Choate, Hy 239 (WTAM 8766). Kenedy: 8 mi N Armstrong, Hy 77 (EAL 2374); 19.1 mi S Riviera (EAL 520); 26.2 mi S Riviera (EAL 731); 34.5 mi S Riviera (EAL 732). Kleberg: 4.5 mi W Riviera (EAL 3918). Knox: Hy 6, 19 km N or S Benjamin (14078). La Salle: Hy 624, 41.5 km W Hy 16 (16664); Hy 624, 21.9 km W La Salle Co. line (16780); Nueces River nr Cotulla (KU 13780-1). McMullen: 16.7 mi N Freer, Hy 16 (10428); Hy 624, Nueces River (16662); Hy 16, 3.7 km N Hy 624 (16691); Hy 16, 11.1 km S Hy 72 (18499); Hy 624, 15.4 km W Hy 16 (16806); Hy 624, 23.3 km W Hy 16 (16663). Nueces: 3.5 mi W Chapman Ranch (KU 61001). Presidio: 38.1 km W Lajitas, Hy 170 (14629). Refugio (UTEP 14031). San Patricio: 1.9 mi S Odem (EAL 255). Starr: Rd on S border of Federal Property, behind Falcon Dam (17784); Rio Grande City (KU 8439); Hy 83, 5.6 km W Hy 755 (14733). Stonewall: Hy 380, 4.3 mi W Hy 83 (2903). Terrell: 1 mi W Pecos River (KU 45354). Throckmorton: 20 mi S Throckmorton (481). Val Verde: 3 mi SE Comstock (KU 129721); 3.2 km N Comstock, Hy 163 (10931); 8 km N Comstock, Hy 163 (10932); 6.7 mi N Comstock (UCM 56666); Hy 163, Juno rd (KU 174941); Hy 163, Baker's Crossing (KU 174942); Langtry (UNM 53328); Evans Creek, 1325 ft, Hy 90, 5.3 mi NNW spur 349 to Amistad Dam (UTEP 9157). Van Zandt: I20, 66 km E Dallas (15809). Webb: 10 mi W Bruni, Hy 359 (10339); Hy 59 betw. Laredo and Hy 2050 (10338); Hy 649, 12.4 km N Hy 3073 (19312); Hy 2050 17.3 mi S Hy 59 (KU 145867). Wise: LBJ Natural Grassland (16338).

CHIHUAHUA (6). 0.7 mi E Aldama Pemex, Hy 16 (UNM 34310); Casas Grandes (BYU 13918); 2 km E Gral. Trias, Hy 16 (UNM 34308); 9.2 km E Gral. Trias, Hy 16 (UNM 34309); 32 km E Gral. Trias, Hy 16 (UNM 34311); 16.1 mi S Jimenez (EAL 2535).

COAHUILA (9). 9 mi N, 5 mi W Castillon (KU 33872); 6 mi W Cuatro Cienagas (KU 47100); 6.9 mi S Cuatro Cienagas (EAL 2420); 30 mi N Monclova (EAL 7); 10.5 mi S Nueva Rosita (EAL 2387); 9.4 mi E Paila (UTEP 3797); 11.8 mi S Sabinas Cachuila (EAL 2391); 7.8 mi W Sacramento (EAL 2623); Santa Cruz, 3250 ft, Hy 57 47 km N Saltillo (UTEP 6336).

DURANGO (2). 7 mi N Donato Guerra (EAL 2704); 2.6 mi S San Juan del Rio (EAL 2705).

NUEVO LEON (12). 8 km NW Allende, Hy 85 (UTA 16126); Aramberri, 3600 ft. (KU 87743); 2.6 mi SW China (UTA 3386); 0.6 mi N Cienaga de Flores (EAL 2603); Garcia (UCM 47378); 23.8 mi NE Gral. Bravo (EAL 1187); 8 mi N Royal Courts, Monterrey (UO 33466); Sabinas Hidalgo (EAL 669, UTA 12282); 7.9 mi W Sabinas Hidalgo (EAL 4250); 17 mi S Sabinas Hidalgo, 1350 ft (KU 67650); Hy 54, 7 mi SW Hy 23 (UNM 37745).

QUERETARO (2). Hy 120, 2 km S Penamiller turnoff (UTA 16127); Old Mill at Hda. San Nicolas Conca (Sharpton's Finca), 10.4 mi S Arroyo Seco (UTEP 9116).

SAN LUIS POTOSI (8). 20.1 mi S Antiguo Morelos (EAL 323); El Salto (UCM 46009); 19 mi S Matehuala, 4700 ft (KU 67651-2); 37 mi S Matehuala, 4650 ft (KU 67653); 39 mi S Matehuala, 5000 ft (KU 67654); 30 mi N Valles, Hy 85 (UTA 4674); 17 mi NE Villa Hidalgo, 5000 ft (KU 67655).

TAMAULIPAS (18). 6 mi N, 6 mi W Altamira (KU 33995); 3 mi S Cd. Victoria (KU 61002); 6 mi S Cd. Victoria (KU 61004); 9 mi S Cd. Victoria (KU 61003); 15.3 mi S Cd. Victoria (EAL 535); 19.1 mi S Cd. Victoria (EAL 536); 26.7 mi S Cd. Victoria (UTA 2457); 36 km S Cd. Victoria, Hy 85 (KU 140057); 39.1 mi S Cd. Victoria (EAL 537); 8 mi W Gonza'lez, Hy 80 (UTA 4855); 10 mi S Mante (UTA 4856); 9 mi S Matamoros (KU 61008); 10 mi SSW Matamoros (KU 61007); 12 mi S Reynosa (UCM 50118); 9 mi S San Fernando (KU 61009); 17 mi NE Santa Teresa (KU 61005); 22 mi NE Santa Teresa (KU 51006); 29 mi S Soto La Marina (UTEP 5100).
TABLE 1. Comparisons of the subspecies of Elaphe guttata on continuously
distributed characters*.

Character emoryi meahllmorum

Body blotches
N (range) [bar.x] 256 (39.5-73) 51.2 154 (27.5-44.5) 37.3
<45,% 2 100
<35.5,% 0 22
Tail blotches
N (range) [bar.x] 223 (12-28.5) 20.4 152 (11-23) 15.9
<18.5,% 26 75
<16,% 4 25
<15,% 2 15
Male ventrals
N (range) [bar.x] 159 (197-225) 210.3 102 (201-233) 214.4
<213,% 72 35
<215,% 91 54
<218,% 96 72
Female ventrals
N (range) [bar.x] 86 (203-228) 216.0 51 (206-236) 221.6
<220,% 71 29
<222,% 83 47
<225,% 97 73
Male caudals
N (range) [bar.x] 139 (62-77) 69.3 89 (65-83) 75.0
<72,% 77 24
Female caudals
N (range) [bar.x] 77 (58-74) 64.2 44 (60-77) 69.1
<72,% 25 68
Spotted ventrals in
 1st 50
N (range) [bar.x] 245 (0-47) 26.9 153 (0-50) 16.4
<24,% 31 70
1st spotted ventral
N (range) [bar.x] 246 (1-205) 12.5 155 (1-226) 26.7
<11,% 62 40

Character guttata

Body blotches
N (range) [bar.x] 56 (27-40) 31.8
<45,% 100
<35.5,% 86
Tail blotches
N (range) [bar.x] 32 (10-16) 12.3
<18.5,% 100
<16,% 97
<15,% 88
Male ventrals
N (range) [bar.x] 27 (211-230) 220
<213,% 7
<215,% 15
<218,% 33
Female ventrals
N (range) [bar.x] 26 (208-238) 227
<220,% 12
<222,% 15
<225,% 31
Male caudals
N (range) [bar.x] 24 (58-76) 69.3
<72,% 75
Female caudals
N (range) [bar.x] 23 (57-73) 64.0
<72,% 96
Spotted ventrals in
 1st 50
N (range) [bar.x]
1st spotted ventral
N (range) [bar.x]

*The data for E. g. guttata are from the Blanchard files; see Materials
and Methods. Data for the remaining subspecies are based on 410
specimens, exclusive of damaged individuals on which certain counts
could not be made.

TABLE 2. Student t tests comparing Elaphe guttata emoryi (s.s.) and E.
g. meahllmorum for each character shown in Table 1. Sample sizes are
listed in Table 1. Also shown are proportions of variance in each
character that are associated with taxa treated as an independent
variable and with longitude and latitude treated in the same manner.

 Proportion of variance
 ([r.sup.2]) associated with:
Character t P taxa longitude latitude

Body blotches 29.74 <0.01 0.69* 0.06 0.46
Tail blotches 11.35 <0.01 0.26* 0.11 0.05
Male ventrals 6.22 <0.01 0.13 0.12 0.16
Female ventrals 5.38 <0.01 0.17 0.17 0.17
Male caudals 11.68 <0.01 0.37* 0.04 0.28
Female caudals 7.59 <0.01 0.32 0.21 0.28
Spotted ventrals in 1st 50 9.43 <0.01 0.18 0.17 0.16
1st spotted ventral 6.58 <0.01 0.09 0.10 0.03

*Clinal effects singly or in combination are significantly smaller than
the variance between taxa.

TABLE 3. Non-discriminatory, non-sexually variable meristics in the
western subspecies of Elaphe guttata*

 emoryi meahllmorum emoryi meahllmorum

postoculars post.
 2-? 0 1 2-4 0 1
 2-2 102 66 3-3 20 11
 2-3 2 2 3-4 22 19
 3-3 0 1 3-5 1 2
ant. temporals 4-4 53 32
 1-2 1 0 4-5 7 3
 2-2 62 31 5-5 0 1
 2-3 15 17 preoculars
 3-3 24 21 1-1 101 70
 3-4 1 0 2-2 2 0
med. temporals supralabials
 2-2 12 5 7-8 1 0
 2-3 9 5 8-? 1 1
 3-3 71 49 8-8 97 58
 3-4 10 8 8-9 5 8
 4-4 1 1 9-9 0 3
 4-5 0 1
 emoryi meahllmorum 12-13 20 15
infralabials 13-? 0 2
11-? 1 0 13-13 26 14
11-11 6 8 13-14 10 9
11-12 11 1 14-? 1 1
11-14 0 1 14-14 6 6
12-? 1 1 15-16 0 1
12-12 20 9

*Counts were made on 178 specimens, exclusive of damaged individuals on
which certain counts could not be made.

TABLE 4. Non-discriminatory, sexually variable meristics in the western
subspecies of Elaphe guttata*

 Males Females
 emoryi meahllmorum emoryi meahllmorum

scale rows, ant.
23 8 5 3 0
24 2 1 1 0
25 53 43 38 18
26 0 0 0 1
27 0 1 0 0
N 63 50 42 19
Scale rows, midbody
25 2 1 1 0
26 0 0 1 0
27 54 31 35 11
28 1 1 1 1
29 8 17 4 8
N 65 50 42 20
scale rows, post.
18 0 1 0 0
19 57 35 22 6
20 2 4 4 2
21 7 10 16 12
N 66 50 42 20

*Counts were made on 178 specimens, exclusive of damaged individuals on
which a few counts could not be made.


We are much indebted to the custodians of the collections from which specimens were borrowed for study, viz.: Nancy Glover McCartney, University of Arkansas Museum; Dr. Stanley E. Trauth, Arkansas State University Museum of Zoology; Ernest A. Liner, private collection, Houma, Louisiana; John E. Simmons and Dr. William E. Duellman, University of Kansas Museum of Natural History; Paul Hyder and Dr. Joseph L. LaPointe, New Mexico State University; Dr. Shi-Kuei Wu, University of Colorado Museum; Dr. Daniel B. Blake and Aine Shiozaki, University of Illinois Museum of Natural History; Allan J. Landwer and Dr. Howard L. Snell, Museum of Southwestern Biology, University of New Mexico; Dr. Laurie J. Vitt, Oklahoma Museum of Natural History, University of Oklahoma; Dr. Jonathan A. Campbell and Carol K. Malcolm, Department of Biology Museum, University of Texas at Arlington; Dr. Robert G. Webb, Laboratory for Environmental Biology, University of Texas at El Paso; and Dr. Kathleen Blair, Department of Biology and Geology Museum, West Texas A & M University.

We are also very grateful for the assistance in various ways of David Kizirian, Joseph T. Collins, Dr. Carl S. Lieb, Dr. John D. Lynch, and Dr. James M. Walker; for the treasured use of word-processing equipment, through the kindness and generosity of Dr. Michael C. Grant; for examination of the holotype of Coluber rhinomegas by Dr. Kevin de Queiroz; for the invaluable support by Dr. William M. Lewis; for permission by Dr. Robert A. Thomas, author of a highly useful athough unpublished review of E. guttata, to use data incorporated in his dissertation; for a critical specimen from Missouri, now in UCM, kindly donated by Dr. John M. Matter; for a vital xerox of a pertinent page in Donndorff (1798), generously supplied by Van Wallach and Garth Underwood; for a copy of Weir's important article (1993), which we would not otherwise have seen, kindly provided by Peter Strimple; and for fascinating historical information about the Arny family and their home town, Hyatt, Kansas, furnished through the kindness of the historian Dorothy Kipper Licktieg of Greeley, Kansas.


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Department of EPO Biology, University of Colorado, Boulder, CO 80309-0334 (HMS), Department of Psychology, University of Colorado, Boulder CO 80309-0345 (DC), Colorado Division of Wildlife, Central Region, 6060 Broadway, Denver, CO 80216 (JRS), Bushmaster Reptiles, 1818 Pine Street, Boulder, CO 80302 (KT)
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Author:Smith, Hobart M.; Chiszar, David; Staley, James R., II; Tepedelen, Kamuran
Publication:The Texas Journal of Science
Date:Aug 1, 1994
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