THE HERPETOFAUNA OF GUADALUPE
MOUNTAINS NATIONAL PARK by JIM W. GRACE, B.S.
A THESIS IN RANGE SCIENCE (WILDLIFE SCIENCE) Submitted to the Graduate Faculty of Texas Tech University in Partial Fulfillment of the Requirements for the Degree of
MASTER OF SCIENCE
Approved
Accepted
May, 198 0 ACKNOWLEDGEMENTS
I wish to thank Dr. John D. Garcia, and the other members of my committee, Drs. Henry A. Wright, C. David Simpson, and John S. Mecham for their helpful comments. Upon the untimely death of Dr. Garcia, Dr. Fred S. Guthery graciously assumed the chair of the committee. This thesis benefited significantly from Dr. Guthery's comments. I am deeply grateful for the cooperation and hospital ity of the staff of Guadalupe Mountains National Park, par ticularly Roger Reisch, Mike and Joan Mayer, and Harry and Anne Steed. Meredith Rohde kindly prepared the base map used for the distribution maps. I would like to thank Don Phillips for his patient assistance with the computer analysis. Ed Hammock, and Glen and John Gilmore cordially provided access to remote areas through their property. Dr. Danny B. Pence prepared, and Dr. Richard B. Loomis identified, the ectoparasites. Drs. James F. Scudday and John S. Applegarth assisted with comments on the area herpetofauna. The following persons provided information on specimens from museum collections: R. Z. Zweifel, J. T. Collins, J. F Scudday, M. McCoid, D. Mosier, E. McGhee, F. I. McCullough, R. G. Webb, J. S. Applegarth, and B. C. Brown. ii I gratefully acknowledge the assistance of Dr. Gary Alstrand and the financial support of the National Park Service through contracts PX702900317 and CX702990008.
Ill CONTENTS
ACKNOWLEDGEMENTS ii LIST OF TABLES V LIST OF FIGURES vi I. INTRODUCTION 1 II. LITERATURE REVIEW 3 III. STUDY AREA AND METHODS 4 Description of Study Plots 15 IV. RESULTS AND DISCUSSION 2 0 Species Accounts 2 0 Analysis of Study Plots 119 Species Present 119 Environmental Factors 122 Effect of Operator on Count 126 Community Composition 127 Abundance 13 0 Age Ratios 13 6 Cover 13 9 Ectoparasites 144 Summary 14 6 LITERATURE CITED 14 8 APPENDIX: LIST OF SPECIMENS 157
IV LIST OF TABLES
1. Characteristics of study plots 16 2. Amphibian and reptile occurrence in plant communities of the GMNP area 22 3. Lizard species observed in plant communities 120 4. Univariate correlations (r) between environmental variables and lizard abundance. Lizard species are those seen most often 124 5. Indices of lizard abundance (maximum and mean number per hectare). Counts in which the species was not observed were excluded 132 6. Lizards observed on plots more than once,the cover they flushed to, and the diversity index for cover 141 7. Chiggers found on amphibians and reptiles from Guadalupe Mountains National Park 14 5
V LIST OF FIGURES
1. Landmarks of Guadalupe Mountains National Park and locations discussed in text 5 2. Location of study plots at Guadalupe Mountains National Park 11 3. Modified line transect technique used to census study plots 13 4. Distributional records for Ambystoma tigrinum in the immediate vicinity of Guadalupe Mountains National Park 21
5. Distributional records for Scaphiopus couchi in the immediate vicinity of Guadalupe Mountains National Park 2 6 6. Distributional records for Scaphiopus bombifrons in the immediate vicinity of Guadalupe Mountains National Park 28
7. Distributional records for Scaphiopus multiplicatus in the immediate vicinity of Guadalupe Mountains National Park 3 0 8. Distributional records for Bufo cognatus in the immediate vicinity of Guadalupe Mountains National Park 32 9. Distributional records for Bufo speciosus in the immediate vicinity of Guadalupe Mountains National Park 34 10. Distributional records for Bufo punctatus in the immediate vicinity of Guadalupe Mountains National Park 35 11. Distributional records for Bufo debilis in the immediate vicinity of Guadalupe Mountains National Park 38
12. Distributional records for Rana berlandieri in the immediate vicinity of Guadalupe Mountains National Park 4 0
VI 13. Distributional records for Kinosternon flavescens in the immediate vicinity of Guadalupe Mountains National Park 41
14. Distributional records for Terrapene ornata in the immediate vicinity of Guadalupe Mountains National Park 43
15. Distributional records for Coleonyx brevis in the immediate vicinity of Guadalupe Mountains National Park 4 6
16. Distributional records for Crotaphytus collaris in the immediate vicinity of Guadalupe Mountains National Park 47
17. Distributional records for Gambelia wislizeni in the immediate vicinity of Guadalupe Mountains National Park 4 9
18. Distributional records for Holbrookia texana in the immediate vicinity of Guadalupe Mountains National Park 51
19. Distributional records for Holbrookia maculata in the immediate vicinity of Guadalupe Mountains National Park 53
20. Distributional records for Sceloporus poinsetti in the immediate vicinity of Guadalupe Mountains National Park 55
21. Distributional records for Sceloporus magister in the immediate vicinity of Guadalupe Mountains National Park 56
22. Distributional records for Sceloporus undulatus in the immediate vicinity of Guadalupe Mountains National Park 58
23. Distributional records for Urosaurus ornatus in the immediate vicinity of Guadalupe Mountains National Park 60
24. Distributional records for Uta stansburiana in the immediate vicinity of Guadalupe Mountains National Park 61
Vll 25. Distributional records for Phrynosoma cornutum in the immediate vicinity of Guadalupe Mountains National Park 6 3 26. Distributional records for Phrynosoma douglassi in the immediate vicinity of Guadalupe Mountains National Park 64 27. Distributional records for Phrynosoma modestum in the immediate vicinity of Guadalupe Mountains National Park 66 28. Distributional records for Cnemidophorus exsanguis in the immediate vicinity of Guadalupe Mountains National Park 68 29. Distributional records for Cnemidophorus inornatus in the immediate vicinity of Guadalupe Mountains National Park 7 0 30. Distributional records for Cnemidophorus tesselatus in the immediate vicinity of Guadalupe Mountains National Park 71 31. Distributional records for Cnemidophorus tigris in the immediate vicinity of Guadalupe Mountains National Park 7 3 32. Distributional records for Eumeces obsoletus in the immediate vicinity of Guadalupe Mountains National Park 7 4 33. Distributional records for Eumeces multivirgatus in the immediate vicinity of Guadalupe Mountains National Park 76 34. Distributional records for Leptothyphlops dulcis in the immediate vicinity of Guadalupe Mountains National Park 78
35. Distributional records for Thamnophis cyrtopsis in the immediate vicinity of Guadalupe Mountains National Park 8 0 36. Distributional records for Diadophis punctatus in the immediate vicinity of Guadalupe Mountains National Park 8 3
Vlll 37. Distributional records for Masticophis flagellum in the immediate vicinity of Guadalupe Mountains National Park 84
38. Distributional records for Masticophis taeniatus in the immediate vicinity of Guadalupe Mountains National Park 8 5
39. Distributional records for Salvadora grahamiae in the immediate vicinity of Guadalupe Mountains National Park 8 7 40. Distributional records for Elaphe guttata in the immediate vicinity of Guadalupe Mountains National Park 8 9 41. Distributional records for Elaphe subocularis in the immediate vicinity of Guadalupe Mountains National Park 90 42. Distributional records for Arizona elegans in the immediate vicinity of Guadalupe Mountains National Park 92 43. Distributional records for Pituophis melanoleucus in the immediate vicinity of Guadalupe Mountains National Park 93 44. Distributional records for Lampropeltis getulus in the immediate vicinity of Guadalupe Mountains National Park 95 45. Distributional records for Lampropeltis mexicana in the immediate vicinity of Guadalupe Mountains National Park 96 46. Distributional records for Rhinocheilus lecontei in the immediate vicinity ol: Guadalupe Mountains National Park 98 47. Distributional records for Sonora semiannulata in the immediate vicinity of Guadalupe Mountains National Park 99 48. Distributional records for Gyalopion canum in the immediate vicinity of Guadalupe Mountains National Park 101
IX 49. Distributional records for Hypsiglena torquata in the immediate vicinity of Guadalupe Mountains National Park 102
50. Distributional records for Tantilla nigriceps in the immediate vicinity of Guadalupe Mountains National Park 103 51. Distributional records for Tantilla atriceps in the immediate vicinity of Guadalupe Mountains National Park 105
52. Distributional records for Crotalus atrox in the immediate vicinity of Guadalupe Mountains National Park 106
53. Distributional records for Crotalus scutulatus in the immediate vicinity of Guadalupe Mountains National Park 107
54. Distributional records for Crotalus viridis in the immediate vicinity of Guadalupe Mountains National Park 109 55. Distributional records for Crotalus molossus in the immediate vicinity of Guadalupe Mountains National Park 111 56. Distributional records for Crotalus lepidus in the immediate vicinity of Guadalupe Mountains National Park 113
57. Number of lizard species, and the total number of lizards per hectare seen in the ten plant communities 12 9
58. Monthly change in lizard abundance in creosotebush, mesquite, fourwing saltbush, and the mean for all plant communities 131
59. Monthly changes in the relative proportion of juveniles to adults of Cnemidophorus tigris and Uta stansburiana 137 CHAPTER I INTRODUCTION
As a major administrator of North American wilderness, the National Park Service is charged by the National Parks Act of 1916 with conserving scenery, natural and historic objects, and wildlife; and providing for the enjoyment of the same in such a manner and by such means as will leave them unimpaired for the enjoyment of future generations. Accomplishment of this objective, as the National Park Ser vice is aware, requires holistic, ecosystem management. In dealing with ecosystems, managers must know what they are managing; thus sufficient baseline data must be available. While faunal surveys have been prepared for most older National Parks, gathering these data remains a priority for the recently designated Guadalupe Mountains National Park (GMNP). Amphibians and reptiles are important parts of the eco system. Their role as consumers, food for predators, and competitors with other animals cannot be overlooked. These animals are also of aesthetic value to park visitors. The Guadalupe Mountains have a rich herpetofauna, with rare and peripheral species. Although herpetofaunal research has been conducted in the area, particularly the eastern slopes, com prehensive data are lacking for the park lands. The lower elevation west side, approximately 33% of the area of the park, has never been examined. The objectives of this study were to synthesize the scattered herpetofaunal information for GMNP and environs, determine the species of amphibians and reptiles that occur in the park and their general distribution, and develop and compare abundance estimates for selected species in widely scattered locations and habitats. CHAPTER II LITERATURE REVIEW
Bailey (1905, 1928) presented early accounts of the vertebrate fauna of the Guadalupe Mountains and Carlsbad Caverns area. The first herpetologist to do serious work in the Guadalupe Mountains was Mosaurer (1932), who recorded 20 species during his month-long stay in Dark Canyon, Eddy County, New Mexico, and near Frijole, Culberson County, Texas. Mecham (1955) investigated the biogeographic relation ships of the Guadalupe Mountains herpetofauna. His research in the area resulted in three additional papers (Mecham 1956, 1957, 1979). The latter is basically a synopsis of his 1955 work with a brief updated species account pertinent to the newly designated GMNP. Gehlbach (1964) produced the first checklist of amphib ians and reptiles of the Guadalupe Mountains, listing 42 species known to occur in Carlsbad Caverns National Park, and later papers resulting from his work in the area (Gehl bach and McCoy 1965, Gehlbach and Holman 1974, Gehlbach 1979) Raun and Gehlbach (1972) compiled an updated checklist of the Texas herpetofauna, listing 199 species for the state.
Additional relevant literature will be discussed in the species accounts. CHAPTER III STUDY AREA AND METHODS
Encompassing 3 0,412 ha, GMNP lies in Culberson and Hudspeth Counties in northern Trans-Pecos Texas, bordered on the north by New Mexico (Fig. 1). Trans-Pecos Texas receives 75% of its total rainfall during the summer and fall, and 50% during July, August, and September (Hinckley 1944). Average annual precipitation increases from 250 mm or less in the basins and 3 00 to 380 mm in the lower mountains, to 500 mm or more in the high mountains, where daily summer showers are common (Bunting 1978) . Year-to-year variation is great and localized heavy rains of short duration are common.
Winds, common all year, reach greatest velocities and are most consistent during March, April, and May, the driest part of the year (Hanks and Dick-Peddie 1974). Wind speeds exceeding 15 0 kph are not uncommon at Pine Springs during these months, particularly March. Winters are mild and summers hot. Regionally the area can be classified as having a continental semiarid climate (Fish and Dvoracek 1980). The Guadalupe Mountains are composed almost entirely of Permian Limestone that extends 17 km into Texas from central New Mexico in the shape of a wedge (Bunting 1978), and com pose one of the most extensive fossil barrier reef systems in JIEWJ4EXIC0 sF'TEXA S
Fig. 1. Landmarks of Guadalupe Mountains National Park and locations discussed in text. Legend for Localities: 1. McKittrick Canyon entrance road 2. Parkers Tank (1478 m) 3. Ship of the Desert (McKittrick) Ranger Station (1597 m) 4. Pratt Lodge, Junction of North and South McKittrick Canyon (1585 m) 5. North McKittrick Canyon 6. South McKittrick Canyon, Turtle Rock area 7. Nickel Creek (1556 m) 8. Choza Spring (1609 m) 9. Nipple Hill (1732 m) 10. Manzanita Spring (1683 m) 11. Smith Spring C1817 m) 12. Frijole Ranch and Ranger Station C168 2 m) 13. Highway 62/180 14. Pine Springs Texas (1719 m) 15. Pine Springs Canyon 16. The Bowl, earthen tank area (2353 m) 17. Dog Canyon Ranger Station (1917 m) 18. West Dog Canyon, Marcus Cabin (1900 m) 19. Lost Peak (2386 m) 20. Cox Tank (2006 m) 21. Bush Mountain (2631 m) 22. Lower Guadalupe Spring, Guadalupe Canyon (1646 m) 23. Signal Peak housing area (1443 m) 24. Junction highways 62/180 and 54 (1280 m) 25. Williams Ranch Road entry, park boundary (1323 m) 26. Bone Spring, Bone Canyon (1676 m) 27. Williams Ranch House (1524 m) 28. Notch in east rim of Patterson Hills (Little Sandy) (1353 m) 29. Lewis Well (1124 m) 30. Red Quartz Sand Dunes (ca. 1158 ra) 31. Eclipse Well (1119 m) 32. Abies Well (1137 m) 33. Northeast corner of Gypsum Sand Dunes (1125 ra) 34. East edge of Salt Lake (1102 m) Selected elevational contours are in meters above mean sea level converted from 7.5 minute U.S. Geological Survey topographical maps (1973). 8 the world (Hayes 1964). On the west, the mountains rise abruptly over 1500 m above the Diablo Bolson and surrounding desert. From 2668-m Guadalupe Peak, they downslope an aver age of 24.6 m/km to 1341 m at Carlsbad Caverns (Hoberg 1949).
Bunting (1978) found the soils generally thin and stony. The mountain soils are derived principally from limestone residuum, while the soils developed in the alluvium from the mountains are characteristically immature, light colored and low in organic matter (Bunting 1978). The basin and plains soils are developed from material that eroded from nearby mountains and range from course gravels to very fine clays (Carter and Cory 193 0) . Deep sand deposits, salt flats, and gypsum deposits also occur in the bolsons (Rowell 1962). Gehlbach (1967) studied the overstory vegetation of the eastern Guadalupe escarpment and presented the first quanti tative data for the area. Glass et al. (1974) identified and mapped 11 plant communities from an interagency survey in Guadalupe Mountains National Park. Bunting (1978) reviewed additional literature and pro vided the first extensive quantitative data on the vegetation of both the eastern and western escarpment, predicting plant communities based on specific site variables. To determine the herpetofauna present in GMNP, I con ducted extensive searches in all major areas of the park from March through October, both in 1978 and 1979. During 1978 the most extensive field work was conducted from May through August, whereas in 1979 over 150 days from March to September were spent in the area. These searches were made by backpacking into the rugged backcountry, day hiking areas of easier access, and driving area roads. These activities were conducted both day and night, but road driving was emphasized at night because herpetofauna are more readily found on roads at this time. Amphibians and reptiles encoun tered were identified, and the plant community (Glass et al. 1974) in which the animal was found and microhabitat condi tions were recorded.
Specimens necessary to document a park checklist were captured by hand; captured with nooses; shot with rubber bands or 22 cal. dust shot; and captured with dip nets, seines, drift fences, and pit-fall traps. Species which had been collected numerous times in the past, or which were rare or uncommon were identified, examined and released. Because all animals were not collected, great care was exercised in their field identification. No individuals were released until positive identification had been made. Specimens were preserved following Duellman (1962) and Pisani (1973), and catalogued and deposited in the Herpetology Collection of The Museum of Texas Tech University. Queries were made to curators at forty herpetological collections nationwide for information on specimens from 10 the Guadalupe Mountains. These data, coupled with the results of this study were used to compile distribution maps Collections or observations made within 0.5 km of each other were combined and mapped as one. Museum records were mapped as accurately as possible, although locations were sometimes vague. Those with extremely vague or improbable locations were excluded. Mapped herpetofaunal remains are from Pratt Cave at the mouth of McKittrick Canyon (Gehlbach and Holman 1974) and Upper Sloth Cave on the western escarpment (Logan and Black 197 9) . The specimen records on which the maps are based, in part, are listed in the Appendix.
To develop abundance estimates, I permanently marked 35 study plots in nine plant communities (Glass et al. 1974) and the quartz sand association of Bunting (1978) (Fig. 2). Although Glass et al. include the quartz sand area in the desert shrub community, I agree with Bunting that it should be separated based on its obvious dunal soils and vegetation. One grassland plot was placed in a grassland area of the mountain shrub community of Glass et al. near Frijole. Two plots were placed immediately outside the park in the gypsum sand dunes west of Eclipse Well. Bunting (1978) includes these dunes in the gypsum association. The community desig nations of Glass et al. seemed descriptive of the other park areas visited. Plot locations were selected within the mapped community subjectively to alleviate their occurrence 11
Fig. 2. Location of study plots at Guadalupe Mountains National Park. The first two digits of the numeric c ode correspond to the plant communities as follows: 01-grassland, 05-mountain shrub, 06-conifer, 09-pinon-juniper^ 10-hardwood, 11-creosotebush, 12- mesquite, 13-fourwing saltbush, 16-desert shrub, and 17-quartz sand. The third digit is the within commu- nity plot designation. 12 on roads, active hiking trails, unwalkable slopes, or atypical areas such as springs or arroyos. Efforts were also made to place the plots at widely scattered locations within each community. Once the general area for placement was determined, plots were placed at random. Two plot sizes were used. One-fourth hectare plots were used in the more dense vegetation of the grassland, mountain shrub, conifer, pinon-juniper, hardwood, and desert shrub communities. One- half hectare plots were used in the more open creosotebush, mesquite, fourwing saltbush, and quartz sand communities, where animals could be seen at greater distances and flush ing distances were greater CDegenhardt 1966). Plots were permanently marked in one corner with white plastic pipe. Additional stakes were placed in the other corners and along the sides to insure proper delineation while plots were being examined. Each plot and its soil surface were photographed in the summer of 1979. A modified form of the line transect (Fig. 3) was used to count lizards as this approach maximized the amount of information gathered per unit of time and effort (Pianka 1970). Similar methods were used by Degenhardt (1966, 1977) in Big Bend National Park and Gehlbach (1979) in the Guada lupe Mountains. I slowly walked the transects of each plot recording data for any reptile seen on the plot. When feasible, checks 13
® 0 © ® [ I I I 0 0 I • t A A T T A
A V FINISH START
Fig. 3. Modified line transect technique used to census study plots. Operator begins at the start arrow and crosses the plot in six equally spaced transects, resulting in three rounds as indi cated by the circled numbers. 14 were made beneath ground cover, and the cover was returned to its original position. The elapsed time for counting a plot was usually 15 to 2 0 minutes. Immediately following the plot count, the soil temperature 1 mm below the soil surface and open shade air temperatures 1 cm and 1.5 m above the surface (Schall 1977) were taken, each time at the same corner of the plot. The date, time since sunrise when the count began, relative humidity, wind speed and direction, and cloud cover (clear, partly cloudy, or overcast) were also recorded. No plots were counted immediately before, during, or after a rain, or before 3 hours or after 11 hours after sunrise. Plots were examined twice monthly April through August and once in September 1979. Statistical tests were conducted following Snedecor and Cochran (1967) and the Statistical Analysis System pack age at Texas Tech University. Multiple regression was used to test the effect of the environmental factors on lizard abundance. The effect of transect round and changes in seasonal abundance were tested by analysis of variance. Changes in numbers of juveniles and adults were examined with Chi-square procedures.
Niche breadth (H) for cover was calculated using
Shannon's (1949) information index, as used by MacArthur
and MacArthur (1961): 15 n H = - Z p^logp^ where p^ is the proportion of the ith resource used. Niche overlap (0) for cover was calculated using the measure of Pianka (1973): n ' P.. P., 0., = O, . = -^ ^ ^^ jk kj n 2 n 2 ^ P.. ^ P 1 i: 1 ik where P^. and P. are proportions of the ith resource used by the jth and kth species respectively. This measure gen erates values less than one and greater than zero. Species arrangement follows Conant (1975); names follow Collins et al. (1978, 1979). Plant nomenclature follows Correll and Johnston (1970).
Description of Study Plots Grassland study plots were established on the southeast side of the mountains (Fig. 2) (Table 1). The vegetation was mainly grama grasses (Bouteloua spp.), purple threeawn (Aristida purpurea), and New Mexico feathergrass (Stipa neomexicana). Interspersed shrubs included broom snakeweed (Xanthocephalum sarothrae), cholla (Opuntia imbricata), soap- tree yucca (Yucca elata), honey mesquite (Prosopis glandulosa) and redberry juniper (Juniperus pinchotii). 16
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>i Ti (U O "^ o in 0 04 ,-^ o> in ro m in vo n iH CM in •P 0 OlP 1 1 1 1 V V V V 1 V in VD rH w rH ^•^ Ln cr» M-l 0 m w o • 4J •H 0 0 CO "«^ m n ro in ro ^ ^ ro -P z rH U] PL4 •H M 0) +J ^ o cn (TJ :3 ^ ^ ^ fC P 5-1 +J ^ ^ Q) ^ iH Xi CO fd 3 13 u •H cn 1 u I >1 T3 C Xi x: fd • 4J C C P ^3 Q) Q) cn cn cn rH •H (0 •H ^-1 •D 0 4J +J c c; rH nj QJ 1 0 0 •H •H +j N (x] :3 CO +J "4-1 c: ^ cn ;3 15 u 4J J W C •H 0 TJ 0 D^ 5-1 d) M g td !3 C c: }-l 0) cn 3 cn fd 9 0 M 0 0 •H fd S-l 0) 0 OJ :3 E-t u o S U CU K u 2 PM Q a 17 Mountain shrub plots were placed in widely scattered areas (Fig. 2) (Table 1). The vegetation was diverse and included oaks (Quercus spp.), junipers (Juniperus spp.), smallseed sacahuista (Nolina micrantha), desert ceanothus (Ceanothus greggii), mountain mahogany (Cercocarpus raontanus), New Mexico agave (Agave neomexicana), lechuguilla (A lechuguilla), and smooth sotol (Dasylirion leiophyllum). Grass composition was variable but included grama grasses and muhlys (Muhlenbergia spp.).
Conifer study plots were established in the Bowl (Fig. 2) (Table 1). The overstory vegetation, which was dense on all plots, was dominated by douglas-fir (Pseudotsuga menziesii), ponderosa pine (Pinus ponderosa), southwestern white pine (P. strobiformis), and gambel oak (Q. gambelii). The sparse under- story had pine muhley (M. dubia), pringle needlegrass (S. pringlei), and Arizona fescue (Festuca arizonica). Overstory species on pinon-juniper plots (Fig. 2) (Table 1) included pinon pine CP. edulus), alligator juniper (J. deppeana), and oneseeded juniper (J. monosperma). The under- story contained smallseed sacahuista, skunkbush (Rhus aromatica), broom snakeweed, and purple threeawn. Hardwood forest plots were placed on stream terraces in McKittrick Canyon (Fig. 2) (Table 1). Overstory vegetation, often with dense canopies, included bigtooth maple (Acer qrandidentatum), chinkapin oak (Q. muhlenbergii) , wavyleaf 18 oak (Q. undulata), alligator juniper, and Texas madrone (Arbutus xalapensis). The understory varied from little vegetation under tree canopies to dense grasses, succulents, and shrubs in the broken areas where mountain muhly (M. montana) , New Mexico muhly (M. pauciflora), and pinon- ricegrass (Piptochaetium fimbriatum) were found. Other species were New Mexico agave and smooth sotol.
Plots were placed in the creosotebush community (Fig. 2) (Table 1). Some mounding occurred under the larger creosotebush (Larrea tridentata), the dominant plant. Addi tional species were honey mesquite, mountain pepperweed (Lepidium montanum), broom snakeweed, pricklypear (Opuntia spp.), and common dyssodia (Dyssodia pentachaeta). A few grasses such as fluffgrass (Erioneuron pulchellum), and black grama (B. eriopoda) were present as were several annual forbs. Plots were placed in the mesquite community west of the mountains, northeast of Lewis Well (Fig. 2) (Table 1). Honey mesquite occurred on the crests of mounds, often 3 m above the inter-mount areas, and 10 to 3 0 m apart. Mountain pepper- weed occurred sparsely at the base of the mounds, with the areas between the mounds largely barren. Broom snakeweed, grassland croton (Croton dioicus), soaptree yucca, and drop- seeds (Sporabolus spp.) were also present.
The fourwing saltbush community was bordered by the gypsum sand dunes west of Eclipse and Lewis Wells. Two 19 plots were placed on the flats and two on the dunes (Fig. 2) (Table 1). The soils of the first two plots were light colored with no rocks. Soils became sandy around areas of disturbance. Fourwing saltbush was prevalent, with thread- leaf sartwellia, mountain pepperweed, plains pricklypear (0. polycantha) dropseeds and grama grasses.
Soils of the active dunes were white gypsum sand. Vege tation was characterized by sweet rosemarymint (Poliomintha incana) , fourwing saltbush, giant dropseed (S^. giganteus) , and mesa dropseed (£. flexuosus). Interdunal troughs had a somewhat heavier soil with range ratany (Krameria gland ulosa) , rough coldenia (Coldenia hispidissima), torrey ephedra (Ephedra torreyana) , and chino grama (B^. breviseta) . Vegetation was diverse in the desert shrub community (Fig. 2) (Table 1). Dominants included oaks (Quercus spp.), redberry juniper, smooth sotol, and catclaw acacia (Acacia greggii). Also present were lechuguilla. New Mexico agave, ocotillo (Fouquieria splendens) , smallseed sacahuista, and desert ceanothus. Grasses included New Mexico muhly and blue grama (B. gracilis). Vegetation of the quartz sand association (Fig. 2) (Table 1) resembled that of the mesquite community. Char acteristic plants were honey mesquite, giant dropseed, mesa dropseed, grassland croton, fourwing saltbush, and soaptree yucca. Duning occurred at the base of the honey mesquite. CHAPTER IV RESULTS AND DISCUSSION
Species Accounts Ambystoma tigrinum.—Mosaurer (1932) first reported the tiger salamander from Dark Canyon of the Guadalupe Mountains, and noted that they were "... said to be common in the water tanks at all ranches in the region." A. tigrinum has since been reported in several areas of the Guadalupe Moun tains CMecham 1955, Gehlbach 1964). The first specimens of A. tigrinum reported from GMNP include one large adult (total length 195 mm) and one small newly metamorphosed in dividual (total length 105 mm), which were collected by George Newman (pers. comm.) on 24 October 1970 from the earthen tank in the coniferous forest of the Bowl (2353 m) CFig. 4) (Table 2). During this study additional specimens from this tank were examined and collected. On 22 October 1978, 12•individuals were seined from the tank (ca. 30 X 18 m surface water measurements) . All were larvae with well developed gills. The salamanders averaged 112 mm in total length, with a range of 100-120 mm. Sixty- four A. tigrinum that had apparently overwintered in the tank were examined 3 April 1979. The tank was nearly dry, and most of the salamanders were confined to a depression filled with ca. 4 1 of water. Four salamanders retaining
four developed gills were seen crawling around out of the 20 21
Fig. 4. Distributional records for Ambystoma tigrinum in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indi cated by circles. 22 TABLE 2.—Amphibian and reptile occurrence in plant communities of the GMNP area. *Denotes locality record of specimen collected previous to this study. us h XJ ^ JJ a Ambystoma tigriniun X X X Scaphiopus couchi X X X X Scaphiopus bombifrons X* X* Scaphiopus multiplicatus X X X X X X X X X Bufo cognatus X X Bufo speciosus X Bufo punctatus X X X X X X Bufo debilis X X Rana berlandieri X X X Kinosternon flavescens X X Terrapene ornata X X X X Coleonyx brevis X* X Crotaphytus collaris X X X X X Gambelia wislizeni X X X X Holbrookia texana X X X Holbrookia maculata X Sceloporus poinsetti X X X X X X Sceloporus magister X* X* Sceloporus undulatus X X X X X X X Urosaurus ornatus X X X X Uta stansburiana X X X X X Phrynosoma cornutum X X X X X X Phrynosoma douglassi X X X Phrynosoma modestum X X X X X Cnemidophorus exsanguis X X X X X X Cnemidophorus inornatus X X Cnemidophorus tesselatus X X X Cnemidophorus tigris X X J\ A Eumeces obsoletus X X X Knmeces multivirgatus X X 23 TABLE 2—Continued x: cn P ^ p M +J u qj x: T-^ ^ Xi ft cn m P cn •H P cn ^ C ^ ^ a c P •O 0) o Di cn td •H •H •n 0 •p -p G c (T3 0) 1 0 0 •H •H -P cn c -p C 15 cn P > SH cn c •H 0 13 0 D^ ^ (U u u p c U (U cn P cn u (D O c •H (d u (U 0 O 0> ft M-l g 8 ft -C o e M-l TJ Leptotyphlops dulcis X* Thamnophis cyrtopsis X X X X Diadophis punctatus X X X X Masticophis flagellum X X X Masticophis taeniatus X X X X X Slavadora grahamiae X X X X X X X Elaphe guttata X X X Elaphe subocularis X X Arizona elegans X Pituophis melanoleucus X X X X X X X X Lampropeltis getulus X* Lampropeltis mexicana Rhinocheilus lecontei Sonora semiannulata Gyalopion canum X Hypsiglena torquata X X Tantilla nigriceps X Tantilla atriceps X* X* X* Crotalus atrox X X X X X X Crotalus scutulatus X X X* Crotalus viridis X X X X X X Crotalus molossus X X X X X X X X X X Cortalus lepidus X X* X 24 water at its edge. All but one of the A. tigrinum examined retained the larval form. The exception was a recently met- morphosed individual with flecked spotting and vestigial gills. The salamanders had a mean total length of 12 9 mm, with a range of 63-145 mm. The earthen tank dried up 2 days later and no salamander remains could be found, even by dig ging in the mud of the tank. However, following summer rains on 2 October 1979, six A. tigrinum were found in a rapidly drying 5 x 12 m tank. Only one had recently under gone metamorphosis. The animals had an average total length of 98 mm with a range of 28-120 mm. The question of how this salamander population was established at the high altitude tank is puzzling, and the answers speculative. The depression has always been a water catchment according to local ranchers and was enlarged and deepened by ranchers in 1953 (Roger Reisch pers. comm.). People possibly introduced the salamanders, but this seems less likely than natural propagation. One large adult A. tigrinum was collected 3.2 km south west of Pine Springs on highway 62/18 0 on the rainy night of 26 May 1979. The Glover Tank, in a grassland community, is 100 m to the north, and although this tank was regularly checked, no A. tigrinum were found there. A road-kill of this species was found 21 km south of Dell City, Texas, on highway 62/180 on the rainy night of 19 May 1978. 25 In a report of zoological field investigations con ducted in August, 1961 on file at GMNP, Frederick Gehlbach records A. tigrinum in McKittrick Canyon, but he does not remember the observation (pers. comm.). With the lack of seemingly suitable ponds or tanks in McKittrick Canyon, the record seems doubtful. Scaphiopus couchi.--A resident of deep alluvial soils CApplegarth 1979), Couch's spadefoot was found from 1100 to 1730 m, but was much more common below 1300 m (Fig. 5). This species is reported here from GMNP for the first time CTable 2) . Mecham (1955) reported S^. couchi from the southern end of th^ Guadalupe Mountains during an August rain. During the summers of 1978 and 197 9, these toads were found on the roads at night from May to September, following afternoon rains showers, but were most common during the warm summer rains of July and August when breeding was taking place in temporary pools. Three pools used by S_. couchi, Parker's Tank (Fig. 1), a tank near the signal peak housing area, and Cottonwood Tank, were found in the immediate area. The salt lake and its side pools in the basin immediately to the west of the mountains were also used by this species. Parker's Tank, the smaller and older one just outside the park ca. 2.4 km east of the Ship of the Desert Ranger Station has been used by several toad species. Frederick 26 Fig. 5. Distributional records for Scaphiopus couchi in the immediate vicinity of Guadalupe Moun- tains National Park. Collection localities from this study or museum or literature records are indicated by circles. Triangles represent personal observations 27 Gehlbach found Bufo debilis, B. puntatus, S. couchi, and S. multiplicatus there on 26 July 1962. These species, with the exception of B. debilis, were found calling from the tank during a heavy rain the night of 18 August 1979. Sur prisingly, no toads were heard calling from the new and larger tank to the south dug in 1977. Scaphiopus bombifrons.—The plains spadefoot is uncommon in the Guadalupe Mountains with one record from Eddy County, New Mexico (Applegarth 1979), and three from the southern end of the mountains in Culberson and Hudspeth Counties (Appen dix) . None was found during this study. S_. bombifrons from the white sand dunes west of Eclipse well (Fig. 1) were col lected by J. F. Scudday (pers. comm.). Although this loca tion is 2.5 km west of the park boundary (Fig. 6), this species has not been found in the park. The habitat looks favorable on the lower west side, except for the lack of tem porary pools necessary for breeding. Many kilometers were walked in this area following summer rains and no pools were found as the substrate of the area allows rapid infiltration. Scudday (pers. comm.) reported that the specimens he collected from the gypsum sand dunes were "exceedingly pale," but tended to darken somewhat after exposure to daylight. Scaphiopus multiplicatus.—The New Mexico spadefoot is recorded here for the first time from GMNP and was found to an altitude of 2353 m (Bowl Tank) , but, like S^. couchi, was 28 > ^ • mi i« » 3 1 0 •• Z N Fig. 6. Distributional records for Scaphiopus bombifrons in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indi cated by circles. 29 more common at lower elevations. The tadpoles of S_. multiplicatus were collected from the Bowl Tank, Cox and Coyote Peak Tanks in West Dog Canyon, Indian Meadow Tank in Dog Canyon and were observed in several tanks in Lincoln National Forest near Queen, New Mexico. The discovery of S^. multiplicatus from the high eleva tion tanks, especially in the Bowl, is surprising. These ephemeral tanks are far removed from the lowlands where the toad is common. Cox Tank was built sometime between 1916 and the early 19 20's, but like the Bowl Tank, was always a natural water catchment according to local ranchers (Roger Reisch pers. comm.). Zwiefel (1968) indicated that S^. multiplicatus had a temperature range for embryologic development similar to that of S^. bombifrons; however, Applegarth (1979) concluded on the basis of distribution in Arizona and the northern Great Plains that the optimal temperature was probably inter mediate between S^. couchi and S. bombifrons. The ability of S. multiplicatus to tolerate cooler temperatures may allow it to survive at the higher elevations in the Guadalupe Mountains, while S. bombifrons, which can also tolerate cool temperatures, may be restricted to lower elevations by fac tors such as substrate (Applegarth 1979). Adult S. multiplicatus were common on the roads at night following summer rains, and in numerous tanks and temporary pools at the lower elevations (Fig. 7). 30 Fig. 7. Distributional records for Scaphiopus multiplicatus in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indi cated by circles. Triangles represent personal observations. 31 I estimated the number of ^. multiplicatus tadpoles (relatively large with well developed hind legs) in the out side 61 cm of the perimeter of Cox Tank on 27 August 1979. The tadpoles in four random plots were counted, averaged, and multiplied by the tank perimeter, ca. 128 m. The outside 61 cm appeared to have the same tadpole density. This re sulted in a conservative estimate of 16,8 00 tadpoles in 78 2 2 m of the tank perifery, or 215 tadpoles m . The tadpole population, like the tanks area, was undoubtedly much larger. Similar densities were observed in the Bowl Tank. Bufo cognatus.—The Great Plains toad was previously reported from one record south of Dell City (Mecham 197 9) and three records from southeastern Eddy County, New Mexico (Applegarth 1979). During this study B. cognatus was a com mon toad of the basin area west of the Guadalupe and Dela ware Mountains, from Dell City to Van Horn, Texas. B. cognatus was found from 1100 to 1342 m (Fig. 8) (Table 2). Large breeding choruses were found at the edge pools of the salt lake (Fig. 1) in the summers of 1978 and 1979. One of these choruses, 3.2 km east and 3.2 km south of Dell City along Texas road 157 6 on 2 June 1978 was deafening. Numerous B. cognatus were seen crossing the road to pools on each side, following heavy afternoon and evening rains that created flooding conditions. 32 V %. mi I* • 0 1 2 kilam«e«r I N Fig. 8. Distributional records for Bufo cognatus in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indicated by circles. Triangles represent personal observations. 33 The great plains toad is recorded near the GMNP boundary by two records near the Signal Peak housing area, 9.7 km southwest of Pine Springs, and from breeding choruses at the east edge of the salt lake, 4.8 km west of the Culberson-Hudspeth County line. B. cognatus has not been found within the boundaries of the park, but may occur there at the lower elevations. Bufo speciosus.—The Texas toad is documented here from the immediate vicinity of GMNP for the first time by three records (Fig. 9). However, the toad has been found nearby to the south toward Van Horn, Texas, and more commonly near Carlsbad, New Mexico, to the northeast (pers. observ.). The three individuals were found 2 to 3 hr after sunset on the McKittrick Canyon entrance road following afternoon and during evening rainshowers. This road was driven at least once almost every night from June to September in 1978 and 197 9. The few B. speciosus found indicate the relative scarcity of this species. Morafka (1974) found that B. speciosus occurred in lowland mesquite-grassland and pe ripheral Chihuahuan Desert situations. Bufo punctatus.—The red-spotted toad is the commonest bufonid of the Guadalupe Mountains, and has been found from 1105 to 1936 m (Fig. 10) (Table 2). This toad seems to prefer rocky regions and open grasslands (Conant 1975). 34 "I?. mil** 0 1 ki lomclar * Fig. 9. Distributional records for Bufo speciosus in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indicated by circles. Triangle represent personal observations. 35 Fig. 10. Distributional records for Bufo punctatus in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indi cated by circles. Triangles represent personal observations. 36 B. punctatus was common on the roads at night follow ing summer showers, but, unlike the other anurans, was also active, though less abundant, on the roads during clear weather. B. punctatus ranges widely in the park. Collecting locations include the north end of El Centre in the hot and dry Patterson Hills (dessicated adult beside a dried pool), tadpoles in stagnant pools in North McKittrick Canyon, Middle McKittrick Canyon (adult under a large boulder in a dry wash), and tadpoles in Lower Guadalupe Spring, Bone Spring, and pools in Shumard Canyon (Fig. 10). On 4 August 1979, all pools of Bone Spring had many tadpoles of two distinct sizes indicating two breeding periods. One small Chead-body length 9 mm) metamorphosed toad was collected as it hopped about on rocks beside the stream (12 00 hr, 2 9" C and sunny). Lower Guadalupe Spring had a tremendous num ber of tadpoles of B. punctatus in several pools on 6 August 197 9, despite heavy use of the spring by cattle. Rains the following week washed all but a few of the tad poles away. B. puncatatus used only Bone Spring and Lower Guadalupe Spring (two of the many available springs) for breeding, al though adults were found near other springs. In these two springs, Rana berlandieri was absent, while it was present 37 in other springs. In North McKittrick Canyon where both R. berlandieri and B. punctatus tadpoles were found, the leopard frog tadpoles were in the clear running spring water while the red-spotted toad tadpoles were found only in stag nant rain-catch pools. Near Choza Spring, an adult B. punctatus was found under a rock 100 m above the spring in a wash, and tadpoles were found in temporary pools of the wash; however, no tadpoles were found in Choza Spring where R. berlandieri adults and tadpoles were common. Bufo debilis.—The green toad is documented in the imme diate vicinity of GMNP by three specimens (Appendix). The two collecting localities, Parker's Tank, 2.4 km east of the Ship of the Desert Ranger Station, and Bear Canyon Ranger Station (i.e. Ship of the Desert Ranger Station (Fig. 11), suggest an upper elevation of 1585 m. The toad is more common at lower elevations at night following summer rains. An intense search was made of the area of the previous collection locations during this study. Although Parker's Tank had breeding choruses of §. multiplicatus, S_. couchi, and a few B. punctatus, no B. debilis were found. Rana berlandieri.—Mosaurer (1932) first reported Rana pipiens (probably R. berlandieri) from Dark Canyon, and a ". . . restricted number at the spring near Frijole" (Man zanita or Smith Spring). The Rio Grande leopard frog is 33 Fig. 11. Distributional records for Bufo debilis in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indicated by circles. 39 common in spring fed pools including the McKittrick Canyon spring system. Smith Spring, Choza Spring, and Manzanita Spring at the eastern base of the mountains (Fig. 12), but is absent from Lower Guadalupe Spring and Bone Spring to the south and west of the mountains. R. berlandieri has also been reported to me by park personnel from the pools of Devils Den Canyon. During the summer rainy season this frog ventures to temporary pools and was found far from permanent water sources on rainy nights. Various size tadpoles of this species were found at all seasons, and egg clusters were observed in the vegetation at the waters edge of quiet pools from April to September. Leopard frog abundance in North and South McKittrick Canyons appeared to be greater above the pools where fish occurred. Two species of fish, rainbow trout (Salmo gairdneri) and sunfish (Lepomis spp.), were introduced in McKittrick Canyon in the early 1950's. Both species read ily ate free-swimming R. berlandieri tadpoles and aggres sively fed on small tadpoles thrown into the water. Sunfish are also sympatric with R. berlandieri in Choza and Manzanita Spring. Kinosternon flavescens.—The yellow mud turtle is docu mented in the immediate vicinity and from GMNP for the first time by two records, both from near Pine Springs (Fig. 13). The first turtle was found by Mike Mayer of the National 40 Fig. 12. Distributional records for Rana berlandieri in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indi cated by circles. Triangles represent personal observations. Squares denote herpetofaunal remains. 41 •y^^ mil* « 0 1 Fig 13 Distributional records for Kinosternon flavesceAs in the iitraiediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indi cated by circles. Triangles represent personal observations. 42 Park Service on 2 July 1978 at 0745 hr on highway 62/180 at the Pine Springs Station (1720 m) following a rainy night. The second K. flavescens appeared in an earthen tank 0.67 km southwest of Choza Spring (1634 m). This tank had not held water in several years and was being filled by the National Park Service for use in road construction. The yellow mud turtle probably reaches its upper eleva tional limit near Pine Springs, and is more common at lower elevations around tanks, muddy pools, and river floodplains. Degenhardt and Christiansen (1974) found all reliable records for K. flavescens in New Mexico below 1524 m. Both the above records are above this elevation. The turtle found at Pine Springs could have been released by tourists; however, this is highly unlikely in the case of the turtle found in the tank near Choza Spring. Iverson (1979) also suggests K. £. flavescens occurs to at least 1500 m, and incorrectly maps the range as not including the southern Guadalupe Moun tains area. Terrapene ornata.—The western box turtle is the common turtle of GMNP, and was found from 1105 to 1650 m (Fig. 14) CTable 2). Two subspecies, T. o. ornata and T. o. luteola, intergrade in the area (Gehlbach 1964); however, turtles from west of the mountains at lower elevations more closely resemble the desert box turtle (luteola), while turtles in 43 Fig. 14. Distributional records for Terrapene ornata in the immediate vicinity of Guadalupe Moun tains National Park. Collection localities from this study or museum or literature records are indicated by circles. Triangles represent personal observations 44 the mountain foothills and to the east more closely resemble the ornate box turtle (ornata). Box turtles were found during the day following rains or early in the morning by driving the roads. Several tur tles were found under the same circumstances while I hiked the lower west side of the park, while I was waiting out a lightning storm in a rocky wash ca. 4 km west of Bush Moun tain on the lower west side of the park, a T. ornata walked past in the rain. It raised its head periodically and opened its mouth. As it did, a drop of water from accumu lated rainfall on the turtle's head would fall into its mouth. Harry Steed of the National Park Service observed and photographed a T. ornata in upper Dog Canyon (1982 m) in October 1979. The turtle had fresh blood and teeth marks on it, as if it had been chewed on, or possibly transported by a predator. Steed also reported this species from West Dog Canyon near New Mexico. Coleonyx brevis.—The secretive Texas banded gecko is documented from the area by five records (Appendix). Only one of these lizards was found during this study, and after it was photographed the specimen escaped. The individual was found under an exfoliated rock slab on a nearly barren south facing slope, desert shrub community, in the Patterson Hills. C. brevis seems restricted to rocky outcrops at lower elevations, possibly because it does not hibernate and cannot 45 tolerate cool winter temperatures (Applegarth 1979). Since this area is near the northern limits of its range (Conant 1975), the cooler temperatures may restrict it to the lower elevations. Previous records include localities in the Patterson Hills and Guadalupe Canyon. The location recorded for speci men BCB #8843, 4.8 km north of Pine Springs, seems improbable. This would be in the extremely rugged higher elevations of the McKittrick Canyon drainage (Fig. 15). Crotaphytus collaris.—Bailey (1928) reported the collared lizard from near Carlsbad Caverns, and Mosaurer (.1932) found C. collaris to be common ". . .as they basked on isolated rocks along the road from Carlsbad to El Paso." Mecham (1955) found this lizard to be most abundant in the pinon-juniper association. I have also observed C. collaris in this plant community in the Guadalupe Mountains of New Mexico, but could find none there in the extreme southern mountains. C. collaris was found from 1340 to 1740 m (Fig. 16). This lizard was not abundant in the immediate vicinity of GMNP, although it seems widely distributed at lower elevations. Gambelia wislizeni.—The longnose leopard lizard has not been previously confirmed in GMNP. Mecham (1979) first collected this species from the mesquite dunes just southwest 46 < -; 1 ^ ^.^^ ^ ^'^ T i.1 ^^> / \, > ^ Fig. 15. Distributional records for Coleonyx brevis in the immediate vicinity of Guadalupe Moun tains National Park. Collection localities from this study or museum or literature records are indicated by circles. Triangles represent personal observations 47 "W ^^V 3 '^/ r • mi )•• 0 1 2 Fig. 16. Distributional records for Crotaphytus collaris in the immediate vicinity of Guadalupe Moun tains National Park. Collection localities from this study or museum or literature records are indicated by circles. Triangles represent personal observations Squares denote herpetofaunal remains. 48 of the park but suggested it did not occur in the Park. During this study G. wislizeni was found widely but uncom monly at low elevations in the southern and western portions of the park from 1105 to 1310 m (Fig. 17) (Table 2). This lizard was found only on soil or loose sand substrates in areas of open vegetation, unlike its congener C. collaris, which was found only once on soil (in a dense grassland) and never on loose sand. In the park reptile observation file, Tony Burgess recorded seeing two G. wislizeni near Lewis Well on 8 August 1973, probably the first within-park observation recorded. Where C. collaris uses large rocks for lookouts and cover, G. wislizeni uses desert shrubs for cover and shade as they lie in wait for prey (Stebbins 1966). These behav ioral and microhabitat differences along with serum protein and morphological dichotomies have been used to support separation of these two groups at the generic level by Montanucci et al. (1975). G. wislizeni as well as other lizards and small mammals burrow and use burrows at the base of the widely dispersed shrubs such as mesquite. In a loose sand substrate, the roots of these shrubs may provide a stable place for burrows, while the crown provides shade for thermoregulation and pro tection from aerial predators. 49 '''I^'^f^^ Fig. 17. Distributional records for Gambelia wislizeni in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indicated by circles. Triangles represent personal observations. 50 Bailey (1928) has a photograph of a "Tiger Lizard" (G. wislizeni) and calls it a conspicious lizard of the arid regions. This photograph may have been borrowed as the species is presently known in Eddy County only from the Pecos River east (Applegarth 1979). Another record of this species in the park file from west of the McKittrick Canyon Ranger Station is probably erroneous. Holbrookia texana.—The greater earless lizard is common in the vicinity of Guadalupe Mountains National Park from 1220 to 1700 m on rocky substrates (Fig. 18) (Table 2). Bailey (1928) mentions the "Texas bar-tailed lizards" (prob ably H. texana) from near Carlsbad Caverns and Mosaurer (1932) found this lizard in the southern Guadalupe Mountains on rocks and in canyons, as did Mecham (1955). The ranges of two subspecies, H. t. texana and H. t. scitula, meet in the region (Conant 1975). Peters (1951) suggested that the specimen collected by Mosaurer (1932) was an intergrade between the two subspecies. Observations made during this study concur for individuals from the higher ele vations of the east side of the mountains, but lizards seen or collected on the west side were brilliantly colored H. t. scitula. Although H. texana is considered a diurnal lizard, one small individual (snout-vent length 25 mm) was found active 51 mi Ic t 0 1 2 0 1 2 ki lomatar s Fig. 18. Distributional records for Holbrookia texana in the immediate vicinity of Guadalupe Moun tains National Park. Collection localities from this study or museum or literature records are indicated by circles. Triangles represent personal observations. 52 18 August 1979 on highway 62/180 near Guadalupe Canyon at 2300 hr after a shower (air temperature 15° C). The lizard's body was pressed to the pavement but it was difficult to determine if it was seeking the heat of the pavement or frightened by traffic. Holbrookia maculata.—Mecham (1979) first reported the lesser earless lizard from two specimens collected by Tony Burgess and Lloyd Logan west of Eclipse Well in the gypsum sand dunes (Fig. 19). However, three specimens had been collected earlier, two near Abies Well, and one in the gypsum sand duneS/by James F. Scudday (Appendix). H. maculata was searched for extensively throughout the area. Only one additional specimen was collected in the large unstabilized gypsum sand dunes west of Eclipse Well. A disjunct region of active gypsum sand dunes in the park 500 m southwest of Lewis Well was searched extensively and trapped with pitfall and driftfence traps, but no H. maculata were found there or in the quartz sand dunes in the park east of Eclipse and Abies Wells, near Scudday's collecting locality. These locations represent the eastern range periphery of the subspecies H. m. approximans. H. m. maculata is found east of the mountains. Conant (1975) reports a ". . . uniformly white or ash gray race . . . ." confined to the White Sands region of 53 ^ s--^^ 3 1 ki I om«(«r I Fig. 19. Distributional records for Holbrookia maculata in the immediate vicinity of Guadalupe Moun- tains National Park. Collection localities from this study or museum or literature records are indicated by circles. 54 New Mexico. Three specimens I examined from the gypsum sand dunes west of Eclipse Well were ash-white. The specimen collected there by Scudday (pers. coram.) had similar coloration. Sceloporus poinsetti.—One of the more ubiquitous and conspicous of the Guadalupe Mountain lizards, the crevice spiny lizard has been recorded there by early investigators (Bailey 1928, Mosaurer 1932, Mecham 1955, Gehlbach 1964). During this study the lizard was found from 1314 m to above 2440 m (Fig. 20) (Table 2). Seventy-eight percent of the S^. poinsetti were found on rocky substrates, but they were also found under rocks, under trash, and on the sides of Williams Ranch House, where as many as four adults could be seen basking at once. Although S^. poinsetti is wide ranging in the more rocky foothills and Patterson Hills, it was only found once in the south and west side lowlands of the park. This juvenile was seen in the creosote bush flats on a small rock pile that surrounded a U.S. Geological Survey section marker 0.8 km from the nearest rocky area. Sceloporus magister.—The desert spiny lizard is known by only five specimens from three collecting localities, two of which are shown in Fig. 21. Gehlbach (1964) and Mecham (1955) noted this lizard was to be expected in the Guadalupe Mountains; however, Mecham (1979) failed to include this species in his later species account for the area. 55 K i-'"A mtl* t 0 1 2 Fig. 20. Distributional records for Sceloporus poinsetti in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indi cated by circles. Triangles represent personal observations. Squares denote herpetofaunal remains. 56 Fig. 21. Distributional records for Sceloporus magister in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indi cated by circles. 57 £. magister occupies desert flatlands, but may also be arboreal (Parker and Pianka 1973) or saxicolous (Stebbins 1954). The collecting localities south and west of the Guadalupe Mountains are of the former habitat. One lizard believed to be this species was seen 6 m up a power-line pole immediately south of the park on the Ed Hammock Ranch, in the bolson plant community (Bunting 1978). However, it could not be caught for positive identification. S^. magister probably will be found in GMNP on the west side below 1220 m. A record from McKittrick Canyon Ranger Station in the Park observation file is probably erroneous. Sceloporus undulatus.—The eastern fence lizard is the most common and conspicuous reptile of the southern Guadalupe Mountains, and has been recorded there by numerous research ers (Bailey 1928, Mosaurer 1932, Mecham 1955, 1979, Gehlbach 1964). During this study £. undulatus was found from 1360 to 2590 m (Fig. 22) (Table 2). Although ubiquitous with respect to plant communities and supposedly elevation (Mecham 197 9), this lizard is absent from the creosote bush, mesquite, and fourwing saltbush communities at lower elevations. This is probably due to the lack of rocky substrates or favorable plants to run into for escape cover. This lizard was found below 1524 m only at a rocky wash in the east rim of the Patterson Hills. 58 i"i. If ^ > A # <^ »^/» I I { i^. mil«« 0 1 2 0 1 2 ki lomaiar « Fig. 22. Distributional records for Sceloporus undulatus in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indi cated by circles. Triangles represent personal observations. Squares denote herpetofaunal remains. 59 Urosaurus ornatus.—Mosaurer (1932) first reported the tree lizard from near Frijole and commented on its ecology as did Mecham (1955) and Applegarth (1979). During this study U. ornatus was found from 1524 to 244 0 m (Fig. 23) (Table 2). The overall distribution of U. ornatus in the vicinity of GMNP resembles that of S_. poinsetti and S. undulatus (Figs 20, 22 and 23). However, the tree lizard seems to be more restricted to boulder strewn washes and canyons. Uta stansburiana.—The side-blotched lizard is one of the common lizards of the desert lowlands to the south and west of the southern Guadalupe escarpment, found from 1105 to 1340 m (Fig. 24) (Table 2) (with the exception of a McKittrick Canyon record—see discussion below). One adult male (snout-vent length 47 mm) U. stansburiana (TTU R-9554) was collected 5 August 1979 ca. 1.6 km above the Pratt Lodge in South McKittrick Canyon (1535 m). This lizard was found active and seemingly healthy under a tree canopy of bigtooth maple and gambel oak on a leaf litter and grass covered terrace near the stream. This is not the usual habi tat of this species (see above. Tinkle 1967, Conant 1975, Applegarth 1979). Extensive searches for lizards have been conducted in McKittrick Canyon above Pratt Lodge in both 197 8 and 1979, but no other U. stansburiana were found. How one 60 w I Fig. 23. Distributional records for Urosaurus ornatus in the immediate vicinity of Guadalupe Moun tains National Park. Collection localities from this study or museum or literature records are indicated by circles. Triangles represent personal observations. Squares denote herpetofaunal remains. 61 Fig. 24. Distributional records for Uta stansburiana in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indi cated by circles. Trianges represent personal observations. 62 lizard got there is problematical, but it is doubtful if a population exists there. Phrynosoma cornutum.—Bailey (1928) reported two or three kinds of horned lizards from near Carlsbad Caverns. Only two, P. cornutum and P. modestum, are known from that park today (Gehlbach 1964). Mosaurer (1932) found P. cornutum up to 1829 m on the east base of the southern Guada lupe Mountains. During this study the Texas horned lizard was found from 1105 to over 1753 m (Fig. 25) (Table 2). Harry Steed (pers. comm.) of the National Park Service reported a Texas horned lizard from near the Dog Canyon Ranger Station (1917 m) in 1976. Phrynosoma douglassi.—Bailey (1905) collected the short- horned lizard at ". . . about 7000 feet [2134 mj altitude in the southern Guadalupe Mountains," but Mosaurer (1932) was unable to find this species. Mecham (1955) found two P. douglassi and considered them a relict species (Mecham 1979). Applegarth (1979) was unable to find this lizard in Eddy County, New Mexico, although there are two records for the county above 1600 m, and he considered them scarce from the Guadalupe Mountains of Texas and New Mexico. During this study over 50 £. douglassi were observed in the high country of GMNP (Fig. 26) (Table 2) above 18 00 m. As many as five of these lizards were seen per day on two 63 Fig. 25. Distributional records for Phrynosoma cornutum in the immediate vicinity of Guadalupe Mountains National Park. Collection localites from this study or museum or literature records are indi cated by circles. Triangles represent personal observations. 64 ' 'L^):F)^^^^ Fig. 26. Distributional records for Phrynosoma douglassi in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indi cated by circles. Triangles represent personal observations. Squares denote herpetofaunal remains. 65 occasions. For the Texas Guadalupe Mountains, the believed scarcity of £. douglassi probably reflects a limited search ing effort in the relatively inaccessible high country. Recognizing its cryptic coloration and behavior, P. douglassi may be the most common lizard of the high country, along with S^. undulatus. Although most sightings (52%) of £. douglassi were made in the conifer forest, the lizards were never seen in the densely forested areas; they occurred almost as commonly in the more open pinon-juniper community. P. douglassi occurs north to Canada and south into Mexico at the higher elevations of the Sierra Madrae Occidental (Conant 1975) , and is aided by viviporous reproduction in living in these colder regimes (Dumas 1964) . Even so, broken forest with openings for light penetration may be necessary for thermoregulation, especially for the gravid female. Phrynosoma modestum.—Bailey (1905) reports the round- tail horned lizard from the "... west base of the Guadalupe Mountains." Mosaurer (1932) did not find this lizard while working near Frijole (1683 m), leading Applegarth (1979) to conclude the lizard was confined to below 1600 m, at least in Eddy County, New Mexico. During this study P. modestum was found from 1100 to 1845 m (Fig. 27) (Table 2). This lizard is a characteristic species of the Chihuahuan Desert (Morafka 1974), and is 66 Fig. 27. Distributional records for Phrynosoma modestum in the immediate vicinity of Guadalupe Moun tains National Park. Collection localities from this study or museum or literature records are indicated by circles. Trianges represent personal observations 67 distributed in the southern Guadalupe Mountains similar to P. cornutum (Figs. 25 and 27). £. modestum was a common road-kill, particularly on McKittrick Canyon entrance road. When threatened, this lizard rarely runs, but instead retracts the limbs and head, flattens the body, and remains motionless. I have seen them stepped on by hikers near Pine Springs. Cnemidophorus exsanguis.—-The Chihuahuan spotted whip- tail is the more common of two parthenogenic teiids of GMNP. During this study C. exsanguis was found from 1220 to 2012 m (Fig. 28) (Table 2). C. exsanguis ranges higher into the Guadalupe Mountains than the other whiptails, particularly in the canyons (i.e. West Dog, Dog, McKittrick, Pine, Guadalupe, and Bone as well as other unnamed west side canyons). This pattern of distri bution has been seen as the adaptability of a parthenogenic lizard to inhabit "weedy" or disturbed habitats (Wright and Lowe 1968), and the higher potential rate of population increase of the all female species compared with the bisexual species (Cuellar 1977). Although the distribution of the Chihuahuan spotted whip- tail overlaps slightly with C. inornatus, C. tigris, and more extensively with C. tesselatus, the species is absent from the desert flatlands at the west side of the park. 68 kT in•'T- — --r ^ A A -^^ J^l?/J :I mi )• • 0 1 2 Fig. 28. Distributional records for Cnemidophorus exsanguis in the immediate vicinity of Guadalupe Moun tains National Park. Collection localities from this study or museum or literature records are indicated by circles. Triangles represent personal observations. 69 The C. sexlineatus sackii reported by Mosaurer (1932) was probably C. exsanguis. Cnemidophorus inornatus.—The little striped whiptail was previously known from the vicinity of GMNP by two col lecting localities northeast of Pine Springs at ca. 1585 m (Appendix). During this study C. inornatus was found from 1220 to 1524 m on gravel-rock substrates, to the south and west of the mountains (Fig. 29) (Table 2). These lizards may be found more often on the east side of the mountains in the shrubby, grassy areas, their pre ferred habitat in southwestern Texas (Schall 1977) . Cnemidophorus tesselatus.—Bailey (1928) has a photo graph of the parthenogenic Colorado checkered whiptail, sup posedly from near Carlsbad Caverns, but does not give the location. During this study C. tesselatus was found from 1341 to 1738 m (Fig. 30) (Table 2). Other researchers have found rocky substrates (Mecham 1955, Wright and Lowe 1968, Applegarth 1979) and sparsely vegetated areas (Schall 1976) to be the preferred habitat of this species. The distribution of C. tesselatus resembles that of C. exsanguis (Figs. 28 and 30), but C. tesselatus was only found in two major canyons, Guadalupe and Bone. Cnemidophorus tigris .—Mecham (1979) reported the west ern whiptail from the mesquite dunes to the southwest of the park near the salt flats. Although several specimens have 70 Fig. 29. Distributional records for Cnemidophorus inornatus in the immediate vicinity of Guadalupe Moun tains National Park. Collection localities from this study or museum or literature records are indicated by circles. Triangles represent personal observations. 71 Fig. 30. Distributional records for Cnemidophorus tesselatus in the immediate vicinity of Guadalupe Moun tains National Park. Collection localities from this study or museum or literature records are indicated by circles. Triangles represent personal observations. 72 been collected from this area and west of the park, C. tigris is documented here from GMNP where it is common at the lower elevations to the south and west (Fig. 31) (Table 2) In southwestern Texas, Schall (1977) found C. tigris had the highest densities in sparsely vegetated, low eleva tion flatland desert. This description fits well the areas where C. tigris was found in the park, from 1100 to 1341 m. Conant (1975) maps C. t_. marmoratus in western Texas, while others (Hendricks 197 9, Vance 1978) separate this cluster into two subspecies, C^. t_. marmoratus to the west of the Guadalupe Mountains and C. t. reticuloriens to the east. However, Vance (pers. comm.) suggests that £. t^. reticuloriens is a synonym of C^. t,. marmoratus as the over lap in characters is great. Specimens I collected during this study could not be satisfactorily separated at the subspecific level. Eumeces obsoletus.—The Great Plains skink was first recorded from the southern Guadalupe Mountains by Bailey (1905) at 2073 m, and has been found to be common but secre tive by subsequent investigators. During this study E. obsoletus was found from 1586 to 2012 m (Fig. 32) (Table 2). In Kansas, E. obsoletus prefers grasslands with sunken rocks for shelter (Fitch 1955), but is restricted to signif icant topographic breaks in xeric regions (Hall 1976). Eumeces 73 ^%2r^- mi I* • 012 ki lomacar I Fig. 31. Distributional records for Cnemidophorus tigris in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indicated by circles Triangles represent personal observations. 74 Fig. 32. Distributional records for Eumeces obsoletus in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indi cated by circles. Triangles represent personal observations. Squares denote herpetofaunal remains. 75 obsoletus was found in the park mostly in riparian situa tions or in dry canyons. Applegarth (1979) suggested that E. obsoletus may not range above 18 00 m in the Guadalupe Mountains. In light of Bailey's (1905) record and observations made in this study, the upper elevational range seems to exceed 2000 m. Eumeces multivirgatus.—Bailey (1905) and Mosaurer (1932) recorded the many-lined skink from the southern Guadalupe Mountains. The E_. multivirgatus complex served as a source of confusion for early investigators due to two dis tinct pattern phases and ontogenetic pattern change (Mecham 1955) , but specimens from the Guadalupe Mountains aided Mecham (1957) in clarifying this taxonomic problem. During this study E. multivirgatus was found in the Bowl and McKittrick Canyon from 1555 to 2440 m (Fig. 33) (Table 2). All were on a leaf litter substrate, into which they quickly disappeared when approached. The many-lined skink seems to be particularly abundant on the hardwood forest stream terraces of McKittrick Canyon. Two found here, a juvenile (snout-vent length 3 2 ram) found 5 August 1979 and an adult (snout-vent length ca. 70 mra) photographed and released 21 April 1979, had bright orange (orange 6A7, [Konerup and Wanscher 19621) coloration on the supralabials and infralabials. The color extended poster iorly on the post labials nearly to the external ear opening 76 Fig. 33. Distributional records for Euraeces raultivirgatus in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museum or literature records are indi cated by circles. Triangles represent personal observations. Squares denote herpetofaunal remains. 77 and anteriorly completely to the tip of the snout, although the rostral was somewhat lighter. The orange color was more diffuse in the blue-tailed juvenile, and turned pale after a few days in 50 percent ethanol. This orange coloration has apparently not been pre viously recorded (Bailey 1937, Conant 1975, Mecham 1957, Smith 1946, Stebbins 1966, and Taylor 1935, 1936). Mecham (1957) reported pale labials on the specimens he examined from the Guadalupe Mountains. Specimens with the orange color that were preserved lost the color and the labials became pale, while pale labialed individuals collected retained that coloration when preserved. Thus examination of preserved specimens reveals nothing as to the labial color of the live animal. Twelve additional live E. raultivirgatusfro m GMNP were examined. All had pale labials. Leptothyphlops dulcis.—Mecham (1955) reported a single specimen of the Texas blind snake from beneath a rock near Pine Springs. Two more were found near the junction of North and South McKittrick Canyons, and one near the Ship of the Desert Ranger Station (Fig. 34) (Table 2). These collecting localities are between ca. 1524 and 1829 m. None was found during this study. Thamnophis cyrtopsis.—Mosaurer (1932) found the black- neck garter snake to be common in Dark Canyon, and collected 78 Fig. 34. Distributional records for Leptothyphlops dulcis in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or rauseum or literature records are indicated by circles. 79 one at the edge of the spring near Frijole (probably Man zanita Spring). Mecham (1955) also reported this species from the area. During this study T. cyrtopsis was observed from 1360 to 2353 m (Fig. 35) (Table 2). T. cyrtopsis was most common along the spring-fed McKittrick Canyon stream system, particularly in North McKittrick Canyon above the pools that contained fish. Above this line Rana berlandieri adults and tadpoles, a pre ferred food item of T. cyrtopsis (pers. observ.) appeared more numerous, thus possibly supporting a larger snake pop- pulation. Five of these snakes were seen 11 July 1978 in North McKittrick Canyon where it crosses into New Mexico, and three and four per day were regularly seen in this area at other times. T. cyrtopsis was also reported to me by park personnel frora the pools of Devils Den Canyon. The snake is frequently seen at Choza and Manzanita Springs, and has been observed in the past at nearby Smith Spring (Roger Reisch pers. comm.) but has not been seen there in the last few years, nor during this study. This species was also found far from permanent water on McKittrick Canyon entrance road, the Bowl Tank, Cox Tank, the tank behind Dog Canyon Ranger Station, and in a wash in the east rim of the xeric Patterson Hills. The last location is 4.8 km from Bone Spring, where the species has never been found but should be expected, and 12 km from Manzanita Spring, the 80 Fig. 35. Distributional records for Thamnophis cyrtopsis in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museura or literature records are indi cated by circles. Triangles represent personal observations. 81 closest permanent water from which the snake is documented. This small (total length 268 mm) individual might have washed down during a flood, but even Bone Spring, the only large west side spring, is not in the upper watershed in which the snake was found. The snake showed no signs of injury to in dicate transportation by a predator. Morafka (1974) found no more than three documented desert localities for T. cyrtopsis in the Chihuahuan Desert of the United States and Mexico. T. cyrtopsis were observed eating R. berlandieri adults and tadpoles in McKittrick Canyon, and S_. multiplicatus tad poles and tadpole shrimp (Triops longicaudatus) at Cox Tank. Harry Steed of the National Park Service also observed a large T. cyrtopsis eating these shrimp at Coyote Peak Tank on 9 September 1979. Diadophis punctatus.—Mecham (1955) reported the ring- neck snake from the southern Guadalupe Mountains, and sug gested that the two species thought to occur there, D. punctatus and D. regalis, were not valid species but rather clinal variations of D. punctatus (Mecham 1956). Gehlbach (1974) found evidence indicating that small D. p. arnyi and the large D. £. regalis forms may act as distinct species in the Guadalupe Mountains, with the former adapted to the more mesic hardwood forest, and the latter to the more 82 xeric coniferous forest. The two forms intergrade exten sively in other areas of contact (Mechara 1979). During this study, D. punctatus was found frora 1555 to 198 2 m (Fig. 36) (Table 2) . The snake is most commonly seen in Mckittrick Canyon, although nearly as many were seen near Pine Springs. Masticophis flagellum.—The coachwhip, common at lower elevations, is reported here from within the boundaries of GMNP where it has been found up to 198 2 ra (Fig. 37) (Table 2) This snake is a coramon road-kill at the lower elevations. On 24 April 197 9, a small M. flagellum was found under a four-wing saltbush eating an adult U. stansburiana, a known food item of this species in Winkler and Andrews Coun ties, Texas (McKinney and Ballinger 1966). Two coachwhips from near the Mckittrick Canyon entrance road showed the reddish coloration above and anteriorly, a condition common in specimens frora Trans-Pecos, Texas (Conant 1975). Masticophis taeniatus.—Mosaurer (1932) first collected the striped whipsnake near Frijole. During this study, M. taeniatus was found frora 1128 m in quartz sand dunes, to 2380-m in the Bowl (Fig. 38) (Table 2). This wide habitat range makes M. taeniatus one of the more ubiquitous of the southern Guadalupe Mountain snakes. 83 mi *• s 0 1 2 Fig. 36. Distributional records for Diadophis punctatus in the immediate vicinity of Guadalupe Mountains National Park. Collection localities frora this study or rauseura or literature records are indi cated by circles. Triangles represent personal observations. Squares denote herpetofaunal reraains. 84 Fig. 37. Distributional records for Masticophis flagellura in the iramediate vicinity of Guadalupe Moun- tains National Park. Collection localities from this study or museum or literature records are indicated by circles. Triangles represent personal observations. 85 Fig. 38. Distributional records for Masticophis taeniatus in the iraraediate vicinity of Guadalupe Moun tains National Park. Collection localities frora this study or museum or literature records are indicated by circles. Triangles represent personal observations. 86 The striped whipsnake was most common in the mountain shrub habitats on escarpment sides and bajadas, where it quickly ascended into small trees and shrubs when approached. Salvadora graharaiae.—The mountain patchnose snake was first reported from near Frijole (Mosaurer 1932), and during this study it was found from 1524 to 2058 m (Fig. 39) (Table 2). This snake apparently inhabits the mid and upper eleva tion rocky areas, and may be restricted from the lower ele vations by the lack of suitable rocky areas. Mecham (197 9) reports the S_. hexalepis {=§_. deserticola) in the collection of Carlsbad Caverns National Park (data of Gehlbach), but did not examine the specimen. My examina tion reveals that the specimen is a S^. deserticola based on two scales between the posterior chin shields and the lateral line on the fourth scale row (Conant 1975). However no data tag was attached to the specimen, and it was in a jar with three other §. grahamiae that were labeled S^. hexalepis. Four specimens of S_. graharaiae collected in 1978 and 197 9 show variability in the number of scales between the posterior chin shields (two or three in various arrangements), but never only one, as suggested by Conant (1975). The lateral line on these specimens is on the third scale row. Considering the confusion surrounding the Carlsbad Caverns National Park specimen, and because the presumed overlapping ranges of S. grahamiae and S^. deserticola meet 87 Fia 39. Distributional records for Salvadora arahamiae in the immediate vicinity of Guadalupe ZllTAnl National Park. Collection localities from this study or museum or literature records are indi cated by circles. Triangles represent personal observations 88 near the southern Guadalupe Mountains, these discriminating characters or the validity of the two species may need to be re-evaluated. Morafka (1974) discusses additional problems with this complex; he considers S^. deserticola a subspecies of S_. hexalepis. Elaphe guttata.—Mosaurer (1932) recorded the corn snake frora "... the garden of Mr. Smith at Frijole." During this study three E_. guttata were found, two at Pine Springs on highway 62/18 0 (172 0 m) and the other near highway 62/18 0 on McKittrick Canyon entrance road (1524 m) (Fig. 40) (Table 2). The snakes were found frora 2 000 to 023 0 hr during rainy or overcast nights. This snake has also been found in McKittrick Canyon (Appendix). Elaphe subocularis.—The Trans-Pecos rat snake reaches its northeastern range lirait in the Guadalupe Mountains. Six E. subocularis were found near GMNP during this study. Three were road-kills; the others were found on highway 62/ 180 on warra overcast nights from 2200 to 2355 hr, from 1150 to 1524 m (Fig. 41) (Table 2). E. subocularis has been recorded in the park only by osteological material from Upper Sloth Cave (Logan and Black 1979). This snake will undoubtedly be found in the park at lower elevations in habitats similar to those near Carlsbad Caverns headquarters, where the snake is quite coramon. 89 Fig. 40. Distributional records for Elaphe guttata in the immediate vicinity of Guadalupe Moun tains National Park. Collection localities from this study or museum or literature records are indicated by circles. Triangles represent personal observations Squares denote herpetofaunal remains. 90 cr^ Tj '^ Fig. 41. Distributional records for Elaphe subocularis in the iraraediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or rauseum or literature records are indi cated by circles. Triangles represent personal observations. Squares denote herpetofaunal remains. 91 Arizona elegans.—The glossy snake is reported here from within the boundaries of GMNP for the first time, although it has been found previously at lower elevations to the southwest of the park on highway 62/180, and in osteological material from Upper Sloth Cave (Logan and Black 1979) and Pratt Cave (Gehlbach and Holman 1974) (Fig. 42). One road- kill from 0.8 km southwest of Pine Springs, highway 62/18 0 (1707 ra),wa s exarained, and it docuraents the species frora the park. Others found during this study were at lower elevations (Fig. 42) (Table 2). All these records were either road-kills or found late at night on the highways. Four road-kills on 3 km of high way 62/18 0 near the Culberson-Hudspeth County line were found 19 May 1979, after a cool clear night that followed two rainy days. The snakes were probably thermoregulating on the warm asphalt when they were run over. Pituophis melanoleucus.—Bailey (1905) reported the gopher snake from the head of Dog Canyon and near Carlsbad Caverns (Bailey 1928) . Mosaurer (1932) found P. melanoleucus at approximately 243 9 ra in the coniferous forest and at the east base of the southern Guadalupe Mountains. Mecham (1955) observed this snake frora the lowland raesquite dunes up to the pinon-juniper association. During this study, P. raelanoleucuswa s found at nearly all elevations in all communities (Fig. 43) (Table 2), making 92 L._ Fig. 42. Distributional records for Arizona elegans in the immediate vicinity of Guadalupe Moun- tains National Park. Collection localities frora this study or museum or literature records are indicated by circles. Triangles represent personal observations Squares denote herpetofaunal remains. 93 u.— Fig. 43. Distributional records for Pituophis raelanoleucus in the iramediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or rauseura or literature records are indi cated by circles. Triangles represent personal observations. 94 it one of the most ubiquitous snakes of the area. While other snakes have nearly the same altitude range in the area (e.g. T. cyrtopsis, M. taeniatus, Crotalus raolossus), only the gopher snake appears coramon throughout this diverse range. Lampropeltus getulus.—The common kingsnake is known from the vicinity of GMNP by two specimens collected south west of Pine Springs (Appendix) (Fig. 44) (Table 2), and, three precaudal vertebrae from Pratt Cave near the mouth of McKittrick Canyon (Gehlbach and Holman 1974). This snake will probably be found in the park at lower elevations. Lampropeltus mexicana.—The rare and secretive gray- banded kingsnake is documented from the Guadalupe Mountains, where it reaches the limits of its northern range, by two records (Gehlbach and McCoy 1965, Worthington and Arvizo 1974). Worthington and Arvizo (1974) report the locality of the latter specimen (UTEP #542) frora the "... east side of Guadalupe Peak, Culberson County, Texas." The snake was indeed collected east of Guadalupe Peak, but more correctly on the southeast side of Hunter Peak (2134 ra) (Fig. 45) (Table 2), where it was found under a rock in June,1967, by Steve West (pers. coram.) Extensive road driving and searching in the two areas where the snake was previously found failed to produce 95 r c •% ;--) J > W4< mi !•• 0 1 Fig 44 Distributional records for Lampropeltis getulus in the immediate vicinity of Guadalupe Moun- tains National Park. Collection l^^^l^^^^fJ^^^^^^^^^ study ©r museura or literature records are indicated by circles. Squares denote herpetofaunal reraains. 96 "H Fiq 45. Distributional records for Lampropeltis mexicana*in the immediate vicinity of Guadalupe Moun- tains National Park. Collection localities from this study or museum or literature records are indicated by circles. 97 another L. mexicana. However, for this secretive species, especially at its range periphery, this rarity is expected. Rhinocheilus lecontei.—The longnose snake is docu mented from the immediate vicinity of Guadalupe Mountains Park for the first time by three records south of the park (Fig. 46)CTable 2). The individuals were collected in August of 1978 and 1979 at 1158 to 1237 ra on asphalt roads at night. R. lecontei has been found in herpetofaunal reraains from Pratt Cave (Gehlbach and Holman 1974). This snake has been found commonly to the east near the Rattle snake Springs, Whites City, New Mexico area (pers. observ.). R. lecontei undoubtedly occurs in the park at least at the lower elevations of the west side. Sonora semiannulata.—The ground snake is known frora the vicinity of GMNP by four speciraens frora McKittrick Canyon (Fig. 4 7) (Table 2). No £. semiannulata were found during this study, but they may occur widely in the park at the lower elevations. The ranges of £. semiannulata and £. episcopa, previously considered separate species, met in the GMNP area. Frost and Van Devender (1979) have suggested that S^. episcopa be placed in the synonymy of S. semiannulata and that no subspecies be recognized. Gyalopion canum.—The western hooknose snake is docu mented by two specimens from west of the Guadalupe Mountains (Appendix) . A third Ficimia cana (=Gyalopion canum) was 98 Fig. 46. Distributional records for Rhinocheilus lecontei in the immediate vicinity of Guadalupe Moun- tains National Park. Collection localities from this study or museura or literature records are indicated by circles. Squares denote herpetofaunal remains. 99 Fig. 47. Distributional records for Sonora semiannulata in the immediate vicinity of Guadalupe Mountains National Park. Collection localities frora this study or museum or literature records are indi cated by circles. 100 found on highway 62/180 169 km east of El Paso, Texas (ca. 8.9 km southwest of Pine Springs (Fig. 48) (Table 2), but the specimen escaped (Chrapliwy and Ward 1963) . Mecham (1955) found this nocturnal snake of the lowlands in the Guadalupe Mountains in the pinon-juniper zone, prob ably its upper limit (Conant 1975). G. canum is to be expected in the park, more commonly at lower elevations. Hypsiglena torquata.—Three road-kills of the night snake were found during this study near GMNP from the low lands to 1524 ra (Fig. 49) (Table 2). Mechara (1955) found H. torquata in the pinon-juniper zone near Queen, New Mexico. H. torquata has been recorded frora within the boundaries of the park only by vertebrae frora Pratt Cave (Gehlbach and Holraan 1974) , but undoubtedly occurs there at lower elevations. Tantilla nigriceps.—The plains blackhead snake is reported frora the iramediate vicinity of GMNP for the first time. A road-kill specimen 9.7 km southwest of Pine Springs on highway 62/180 (Fig. 50) (Table 2) was collected 13 July 1979, and together with two specimens frora near Salt Flat, Texas (Appendix), represent the known speciraens frora the area. T. nigriceps will undoubtedly be found at lower elevations within the boundaries of GMNP. 101 ii Fig. 48. Distributional records for Gyalopion canum in the iraraediate vicinity of Guadalupe Mountains National Park. Collection localities frora this study or rauseum or literature records are indicated by circles. 102 Fig. 49. Distributional records for Hypsiglena torquata in the iraraediate vicinity of Guadalupe Moun tains National Park. Collection localities frora this study or museum or literature records are indicated by circles. Squares denote herpetofaunal remains. 103 Fiq 50. Distributional records for Tantilla nigriceps in the immediate vicinity of Guadalupe Mountains National Park. Collection l^^^^^^^f.^d?- this study or rauseura or literature records are mdi cated by circles. 104 Tantilla atriceps.—No additional Mexican blackhead snakes were found during this study, but speciraens in five collections (Appendix) docuraent the species frora the area (Fig. 51) (Table 2), to ca. 1524 m. Crotalus atrox.—Mosaurer (1932) found the western diamondback rattlesnake in Dark Canyon and at the foot of the southern part of the Guadalupe Mountain range. Mechara (1955) found C. atrox to be coraraon at lower elevations. Dur ing this study C. atrox was found frora 1105 to 183 0 m (Fig. 52) (Table 2) . This crepuscular snake was common on area roads and was also abroad during the day in shaded areas or on overcast days. C. atrox apparently does not invade the mountains proper in the park, but is replaced there by C. molossus (Figs. 52 and 55) . Crotalus scutulatus.—Two specimens of the Mojave rattle snakes are known from the iraraediate vicinity of G^ITJP (Fig. 53) (Table 2), where the species reaches the liraits of its north eastern range (Conant 1975). One speciraen was taken by Tony Burgess and Lloyd Logan in the gypsura sand dunes west of Eclipse Well; the second was found on highway 62/18 0 by Tom Van Devender 18 km southwest of Pine Springs. Others have been found toward Dell City, Texas, to the west, and Van Horn, Texas, to the south (Appendix). C. scutulatus probably occurs in the park on the west side at lower elevations. 105 Fig. 51. Distributional records for Tantilla atriceps in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or museura or literature records are indi cated by circles. 106 Fig. 52. Distributional records for Crotalus atrox in the iraraediate vicinity of Guadalupe Mountains National Park. Collection localities frora this study or rauseura or literature records are incidated by circles. Triangles represent personal observations. 107 Fig 53. Distributional records for Crotalus scutulatus in the iraraediate vicinity ^^ Guadalupe Mountains National Park. Collection localities frora this study or rauseurao r literature records are indicated by circles. 108 With these recent records, the range maps of Gloyd (1940), Klauber (1972), and Stebbins (1966) for C. scutulatus are not as precisely correct for the species west of GMNP as Conant (1975). Jacob (1977), however, found the range raap of Stebbins (.1966) to be more correct for extreme southwestern New Mexico. The Guadalupe and Delaware Mountains to the south, may serve as barriers to the eastern dispersal of C. scutulatus. Although an inhabitant of the teraperature raesquite grass lands, this species has been found up to 2 07 3 ra in the Chiricahua Mountains of Arizona (Klauber 1972). Crotalus viridis.—The western rattlesnake is reported here frora the park for the first tirae and from additional locations in the southern Guadalupe Mountains (Appendix). National Park Service employee Harry Steed, stationed in Upper Dog Canyon, captured a large (total length 965 ram) male C. viridis 6 July 197 9 and reported seeing several others around the ranger station (1917 m) (Fig. 54) (Table 2). Other C. viridis were found down to 1110 ra. One road- kill individual with head and rattles recently reraoved was found at 2134 ra near Queen, New Mexico, in the pinon-juniper comraunity. Although the western rattlesnake is considered a snake of the grassy plains (Klauber 1972), this record is not surprising in light of high elevation records for other mountain ranges. 109 Fig. 54. Distributional records for Crotalus viridis in the iraraediate vicinity of Guadalupe Moun tains National Park. Collection localities frora this study or rauseura or literature records are indi cated by circles. Triangles represent personal observations. 110 crotalus raolossus.-Bailey (1905) lists the blacktail rattlesnake as the "... coraraon rattlesnake of the Guada lupe Mountains," where he found the snake in the gulches, high up on the range, as did Mosaurer (1932). During this study, C. raolossus was found from 1110 to 2450 m (Fig. 55) (Table 2). This wide range of habitats makes C. raolossus, 1^^® £• melanoleucus, one of the raost ubiquitous snakes of the area, but unlike the gopher snake, it seems to be more abundant at elevations above 14 00 m. Below 18 00 ra the blacktail rattlesnake overlaps exten sively with C. atrox. A rancher living west of the Patterson Hills, near the east edge of the Salt lake reported the "green rattler" on his ranch after heavy suraraer rains, indi cating that the saxicolous C^. molossus may have been coming down out of the rocky Patterson Hills with increased precip itation. However, during the summer of 1979, I examined several C^. molossus, C^. atrox, and C. viridis that were all killed on this ranch during July and August when there were no exceedingly heavy rains, although this is the season of highest precipitation. Crotalus lepidus.—Schwartz and Babis (1949) first reported the rock rattlesnake frora the Guadalupe Mountains, frora a specimen collected by Louis Kincaid (probably Noel Kincaid, foreman on the J. C. Hunter Ranch) on the "Man- zanital Ranch" owned by Wallace Pratt at 1524 ra (undoubtedly 109 mil* « 0 1 2 Fig. 54. Distributional records for Crotalus viridis in the iraraediate vicinity of Guadalupe Moun tains National Park. Collection localities frora this study or rauseum or literature records are indi cated by circles. Triangles represent personal observations. 110 Crotalus molossus.—Bailey (1905) lists the blacktail rattlesnake as the "... common rattlesnake of the Guada lupe Mountains," where he found the snake in the gulches, high up on the range, as did Mosaurer (1932). During this study, C. raolossus was found frora 1110 to 2450 m (Fig. 55) (Table 2). This wide range of habitats makes C. raolossus, like P. raelanoleucus, one of the raost ubiquitous snakes of the area, but unlike the gopher snake, it seeras to be raore abundant at elevations above 14 00 ra. Below 18 00 ra the blacktail rattlesnake overlaps exten sively with C. atrox. A rancher living west of the Patterson Hills, near the east edge of the Salt lake reported the "green rattler" on his ranch after heavy suraraer rains, indi cating that the saxicolous C^. raolossus raay have been coming down out of the rocky Patterson Hills with increased precip itation. However, during the suraraer of 1979, I exarained several C. raolossus, C^. atrox, and C. viridis that were all killed on this ranch during July and August when there were no exceedingly heavy rains, although this is the season of highest precipitation. Crotalus lepidus.—Schwartz and Babis (194 9) first reported the rock rattlesnake frora the Guadalupe Mountains, frora a speciraen collected by Louis Kincaid (probably Noel Kincaid, foreraan on the J. C. Hunter Ranch) on the "Man- zanital Ranch" owned by Wallace Pratt at 1524 m (undoubtedly Ill mil* « 0 1 2 0 1 2 kilomacar« Fig. 55. Distributional records for Crotalus molossus in the immediate vicinity of Guadalupe Mountains National Park. Collection localities from this study or rauseura or literature records are indi cated by circles. Triangles represent personal obsen/'ations. 112 McKittrick Canyon). Several additional specimens, all from GMNP, represent the species from the area (Appendix). During this study, C. lepidus was found down to 1616 ra (Fig. 56) (Table 2). Mecham (1955, 1979) reports an indi vidual at 2500 ra (probably coniferous forest). The Guada lupe Mountains are the northeastern range periphery of this species (Conant 1975). Mechara (1955) found the characters used to discrirainate between the two subspecies that raeet in the area interraediate, and Gehlbach (1964) states that C. _1. lepidus and C. _1. klauberi intergrade in the area. The individuals I exarained also appear to be intergrades between the two races. University of Wisconsin geologist Kent Kirby reported seeing several sraall rattlesnakes in upper Shuraard Canyon. His careful description fits C^. lepidus perfectly. Two of the snakes he observed at the entrance to a small shelter cave, but when I visited the shelter with him they were not found. Other possible species.—Hylactophryne augusti is known from three records east of GMNP. Koster (194 6) collected a juvenile 19.3 km northwest of Carlsbad, New Mexico, Seifert (1978) reported a speciraen frora Pyote, Texas, and tape recordings were made of a calling frog south southwest of Carlsbad Caverns (Applegarth 197 9). This species may be found in the park, most likely on a grassy bench adjacent 113 /^: ^^ >r\ ^ ^-. V < - r' f- Fig. 56. Distributional records for Crotalus lepidus in the iramediate vicinity of Guadalupe Moun- tains National Park. Collection localities from this study or rauseura or literature records are indicated by circles. Triangles represent personal observations. 114 to cliffs (Applegarth 1979) calling frora a depression or rodent hole on a rainy spring night. Acris crepitans occurs in drainages east of the moun tains including the Pecos River (Stebbins 1951) and Dark Canyon (Mechara 1955) . Once considered a widespread relict of pluvial periods in the Chihuahuan Desert (Milstead 1960), this species no longer exists in the lower reaches of the Pecos River, except at a small refugiura near Balmorhea, due to raan'sdegradatio n of the river (Scudday 1974). A. crepitans probably does not occur in the disjunct water pools of GMNP. Bufo woodhousei is found east of the park near the Pecos River (Applegarth 197 9) and raapped as occurring west of the Park (Conant 1975) . It is doubtful that B. woodhousei occurs in the park, as it prefers streara and river bottomlands, although its southern range is characterized by sraall, appar ently disjunct populations (Axtell 1963). Hyla arenicolor was indicated on a raap by Stebbins (1951) to occur in or near the Guadalupe Mountains, but this is alraost certainly in error (Mechara 197 9). This species probably does not occur in the park even though sorae of the springs look like favorable habitat. Rana blairi occurs east of the Guadalupe Mountains in open grassland habitats. Applegarth (1979) discusses Eddy County material, including a record just south of Eddy 115 County in Texas frora Delaware Creek, where John S. Frost found R. blairi sympatric with R. berlandieri. R. blairi apparently does not occur in the park. Rana catesbiana was originally absent from Trans-Pecos Texas (Raun and Gehlbach 1972), but has been widely intro duced. The species does not presently occur in GI4NP. Five aquatic turtles, Chelydra serpentina, Chryserays picta, Chryserays scripta, Chryserays concinna, and Trionyx spiniferus occur in the Pecos and Black River (Degenhardt and Christiansen 1974) but do not enter the park. A nuraber of whiptail lizards frora the area have been previously confused with Cneraidophorus gularis (Gehlbach 1964) but are now considered to be C. exsanguis, deraon- strating the confusion that has surrounded Cneraidophorus taxonoray for years. C. gularis is found to the east in Eddy County, New Mexico, where Applegarth (1979) suggests it reaches the western liraits of its range. It prefers the wetter floodplains or watercourses, habitats similar to those found by Schall (1977) for the species in southern Trans- Pecos Texas, although he failed to find this species in southern Culberson County (pers. coram.). Some aquatic and serai-aquatic snakes that have been recorded in drainages to the east of the Guadalupe Mountains (Mechara 1979), but that probably do not occur in the park. 116 are Nerodia erythrogaster, Tharanophis proximus, and T. marcianus. Heterdon nasicus is found near Carlsbad, New Mexico, and invades the mountains at the lower elevations such as Dark and Walnut Canyons. H. nasicus may occur at the lower elevations of GMNP. Lampropeltis triangulum occurs south of the Guadalupe Mountains but has not been recorded near the park. Coluber constrictor is known from isolated populations in the south west (Conant 1975) , the closest being in the Davis Mountains (Glidwell 1974) . Sistrurus catenatus is also found in the region, but has not been recorded from the Guadalupe Mountains. Two additional snakes probably occur in the southern Guadalupe Mountains. Opheodrys vernalis was suspected on the basis of a rancher report (Mecham 1955) and a sight record by Tony Burgess (Mecham 1979), and is documented by three precaudal vertebrae (TTU P-83 67) from Upper Sloth Cave (Logan and Black 197 9) . A live specimen reraains to be collected. 0. vernalis should be looked for at interraediate and upper elevations. Five precaudal vertebrae (TTU P-8374) from the same cave document Trimorphodon biscutatus frora the Guadalupe Mountains for the first tirae (Logan and Black 197 9). This is the only record of this rare snake frora the area and is 117 the northeastern limit of its known range (Gehlbach 1971). T. biscutatus should be expected in the desert areas at lower elevations. The status of several species of amphibians and rep tiles, particularly snakes, that occur in the vicinity of GMNP remains in question. It is often difficult to find these secretive nocturnal animals. This problem is cora- pounded in the park because of the difficulty in traveling in the remote areas, especially at night. If the surfaced road planned by the National Park Service (National Park Service 1973) is built along the western base of the moun tains, it may provide an avenue for increased knowledge of some of these questionable species. The road would provide an excellent place to search frora at night by road driving, and paradoxically would lead to an increased incidence of road kills in this area. The effect of years of ranching on sorae species cannot be overlooked. In the early 1900's landowners grazed large nurabers of livestock in the Guadalupe Mountains of Texas. Much of the land was severely overgrazed for sorae period of tirae prior to the park's establishraent in 1972 (Krysl 1979). Degenhardt (1977) reported changes in vegetation and saurofauna on study areas after 10 years or protection frora grazing. He found an increase in plant density and diver sity and suggested that lizard density increased with 118 increasing vegetation to a raaxiraura and then declined. On plots Degenhardt established in Big Bend National Park in 1966, vegetative growth increased and H. texana becarae rauch less abundant, while the effect on cneraidophorines was not as great. Milstead (1977) discussed population dynaraics of three species of whiptail lizards (Cneraidophorus spp.) on Black Gap Wildlife jyianagement Area, Brewster County, Texas, over a 2 0-year period. He found interpretation of popula tion fluctuations difficult, but noted increased rainfall and concomitant vegetative changes were plausible explanations. Mechara (1979) noted considerable differences between herpetofaunas on the eastern and western sides of the Guada lupe Mountains at lower elevations. Riparian eleraents are limited to the eastern side; bolsons predominate to the west. This difference in eastern and western lizard faunas is not clear, however. For example, sand and dune situations irarae- diately to the west of the raountains support such lizards as G. wislizeni, H. raaculata, £. raagister, U. stansburiana, and C. tigris. Similar habitats are also found east of the mountains near the Pecos River where the same species are present (Applegarth 197 9). Some snakes raay also follow this pattern, but raore documentation is needed. The most obvious factor relating the distribution of these lizards appears to be substrate. Applegarth (1979) 119 broadly categorizes the soils of Eddy County, New Mexico, into limestone rock and rocky soils, loamy soils, and dune and sandy soils. This classification can be generally applied to the southern Guadalupe Mountains. The mountains and the bajadas emanating from them have rocky soils, while the loaray, and sandy and dunal soils are found to the west. The herpetofaunal coraposition on the loaray soils and sandy to dunal soils is nearly interchangeable, but consider ably different than the group of species that occurs in the rocky areas of the raountains and foothills. Only a few species seera to frequent both, and a predorainance of the reptile species (ca. 57%) of the area are limited to the rocky habitats of the mountains. Analysis of Study Plots Species Present Lizards were observed on study plots in all plant com munities except the conifer forest. No lizards were found on four other plots, grassland Oil and 013, mountain shrub 052, and pinon-juniper 092. Creosotebush plots had the most species, followed by fourwing saltbush and desert shrub plots (Table 3). Five species were seen only once. C. brevis was found under a rock on desert shrub 163. H. texana was also found on this plot. S. poinsetti was seen on creosotebush 111 on a rock pile. P. cornutum was observed on fourwing saltbush 132. A juvenile P. douglassi was observed on pinon-juniper 091. 120 TABLE 3.—Lizard species observed in plant comraunities X cn Xi D A-> U 0u) ja r-i J3 Xi 04 cn CH P TJ cn •H 0 tn U ^3 Xi x: (TaJ C c TS OJ OJ CP cn cn (T3 •cH: •an 0 -p 4J C rH (d uCD I 0 0 -H •H J-) N cn w +J M-l C 5 cn P 15 ^ -p rH W c •H 0 T3 0 rr iM 0) (T3 (0 13 C c ^ (U cn a cn u(« V U 0 0 •H (tJ }^ 0) 0 Gambelia wislizeni X X X X 4 Holbrookia texana X 1 Sceloporus poinsetti X 1 Sceloporus undulatus X X X 3 Urosaurus ornatus X X X 3 Uta stansburiana X X X X 4 1 Phyrnosoma cornutum X Phrynosoma douqlassi X 1 2 Phrynosoma modestum X X X 2 Cnemidophorus exsanguis X Cnemidophorus inornatus X 1 Cnemidophorus tesselatus X 1 X X X X 4 Cnemidophorus tiqris Eumeces multivirqatus X 1 7 3 5 5 3 Totals 1 2 0 1 3 121 Although no lizards were found on seven study plots, this does not raean none was present in the general area. Plots were placed in areas of uniform topography and vegetation, while lizards are often found only in topo-- graphic or vegetative breaks. For example, S. poinsetti, S. undulatus, U. ornatus, P. douglassi, and E. multivirqatus occur in the coniferous forest of the Bowl. But in this area, S_, poinsetti was found on exposed rock outcrops; S. undulatus was found on rock outcrops, dead logs, and old water tanks in exposed areas; U. ornatus was found on boulders in rocky areas; and P. douglassi and E. multi virgatus were found along trails or in open areas. No indi viduals of these species were found on plots in the more dense and uniform coniferous forest. Several species occurred near the grassland plots, but seldom used areas of open grassland, away from open spaces for running and rocks and shrubs for cover. On the pinon-juniper plot where no lizards were found, ground cover was sparse. The plot was near Cox Tank and many mule deer (Odocoileus hemionus) and elk CCervis elaphus) traveled across the plot enroute to water. Nearby, S_. undulatus was coraraon on the cairns used to raark hiking trails. At the higher elevations, raore lizards, particularly S. undulatus and P. douglassi, were coraraonly noted along cleared hiking trails than in vegetated areas nearby. On several 122 occasions sections of trails were walked and the lizards counted. Then similar sections of an untraveled area nearby were examined. In each case many more lizards were seen along the trail. Several factors may be suggested to account for this difference in lizard abundance. Lizards along the trail could be easier to see and hear. Hiiing the trail is also quieter and lizards may be approached more closely. Thus the higher count could be a reflection of differential observability. The discrepancies in counts, however, would not seem to be totally explained by this bias. Another plausible explanation is that the trail openings may provide an excellent place for feeding for a sit-and-wait strategist such as S_. undulatus. Lizards could dart into the opening of the trail to capture insects and then return quickly to cover. Also, in the cooler raoun tains, the openings provided by the cleared trails in areas of dense vegetation may provide warmer substrates for therraoregulation. Environraental Factors Analysis was conducted to determine if the environmen tal factors of air and substrate temperatures, relative humidity, wind speed, date, and time since sunrise could be used to predict lizard abundance. No factor or group of factors were found that accounted for a significant araount 123 of the variation in abundance, when the analysis was con ducted by species, the correlations reraained low. In species seen often, the changes in the environraental factors reraained poorly correlated with changes in abundance (Table 4). Cer tain trends were apparent, however. Fewer lizards were seen during high winds and raore lizards were active during the hotter portions of the day in spring and the cooler or cloudy portions in suraraer. Simi lar seasonal shifts in daily activity patterns have been noted in other lizard species (Mayhew 1964, Pianka 1969). Relative humidity seeraed to have little effect. The difficulty in predicting lizard abundance based on environmental factors such as temperature, time and weather conditions is not surprising. Lizards are active at optimal temperatures that vary with the species. But incident radi ation, substrate temperatures, and behavioral therraoregula tion raay be raore important than air in determining body temperature (Bogert 1949, Pianka 1970, Schall 1977). Daily activity patterns are also complex. Cowgell and Underwood (1979) found a distinct behaviorally regulated daily pattern in body temperatures of Sceloporus occidentalis independent of light-dark regimes. Whiptail (Cnemidophorus spp.) activ ity patterns have been described as biraodal by Milstead (1957a, b), Echternacht (1967), and Medica (1967), while more C. tigris were seen in the morning than in late afternoon 124 Q) cn n 0^ 00 ^ iH (U •H (^ ^ 00 00 CN 0) o !M vo fH cn r- rH C H LD rH r-i CN •eH •cH 0 • • • • t en to m o O o o O u 1 1 1 1 fO N •H H c o ^ CN cn r» T3 r^ 0^ cn in ro G +J -H r^ CO 00 m r> (T3 tC rH CN LO ro Pianka (197 0) found that C. tigris abundance was corre lated to rainfall patterns of the preceding 5 years in rauch of the species' geographic range. He suggested that increased rainfall led to an increased food supply and raore lizards. Scudday and Dixon (1973) and Mitchell (1979) have related diet shifts in cneraidophorines to changes in rainfall patterns. Rainfall in the southern Guadalupe Mountains has been measured at stations near Salt Flat, Texas since 1952, and Carlsbad Caverns, New Mexico since 1931. Data for portions of 1977 through 197 9 have also been gathered in the park near Frijole (Fish and Dvoracek 1980). Precipitation for the years 1973-1977 averaged 215 mm at Salt Flat while the long-term mean was 217 mra. Data for 1978 were not available. The long-term mean at Carlsbad Caverns was 35 9 ram with the precipitation from 1974 to 1978 averaging 453 ram. The higher 5-year mean is due largely to a period of heavy rainfall from 21-26 September 1979 when 319 mm fell at Frijole. The 5- year period from 1973-1977 had a mean precipitation of 403 mm at Carlsbad Caverns, more nearly that of the long-terra aver age. Although Fish and Dvoracek (1980) have argued caution 126 in extrapolating exact precipitation in the park from mea surements taken at outside stations, the general pattern for the 5 to 6 years previous to 197 9 appears nearly average when compared to the long-term data, except for the flood of September 1978, and 1979 did not deviate largely frora the pattern. Lizard abundance, then, at least for C. tigris, based on its correlation with previous rainfall, would be expected to be nearly average to slightly higher, depending on the effect of the high 1978 precipitation on the food supply and the speed with which lizard populations would reflect this change. Effect of Operator on Count The lizards seen per round were recorded to deterraine the effect of the operator walking through the study plot on the behavior of the plot residents. When all plots were analyzed as a group, raore lizards were seen during round one, the first and second walk through the plot (Fig. 2), than during rounds two or three (p<0.05). When the corarau- nity plots were analyzed separately, however, only the creo sote bush and fourwing saltbush coramunites showed this trend, with more lizards seen on rounds one and two than on round three (p<0.05) . One obvious problem that might arise frora walking rounds through the plot is that the same individuals might be seen 127 again and again on consecutive passes, tending to give a count biased too high. Degenhardt (1966) noted this, but suggested the error might have a high degree of uniformity with the same or different operators. Providing lizards were distributed randoraly over the plot, a large nuraber of lizards seen on the early rounds raight indicate that raost residents were being seen and flushed with the early inva sion of the operator, leading to a bias of a lower count. More lizards found during the later rounds on the other hand, raight suggest that residents were remaining on the study plot and being seen more than once. The data collected during this study indicate a trend toward seeing raore lizards on the early rounds through the plot. As I counted a plot, lizards nearly always ran to the cover of a plant or burrow not to be seen again during the count. Thus, the problem of counting an individual raore than once was probably rainor. Coraraunity Coraposition Pianka (1967) found the nuraber of syntopic lizards increased frora four to ten along a latitudinal transect from Idaho to southern Arizona. Predatory lizards, snakes, and birds also increased from north to south. A sirailar pattern appears on the Guadalupe Mountain study plots frora high ele vation to low elevation. 128 In Fig. 57 the plant coraraunities are arranged frora rela tively raore mesic to more xeric. This arrangement also cor responds roughly to higher altitude to lower altitude. Determining the order of some of the coraraunities was subjec tive, but there is still a general trend of raore lizards, both in nuraber of species and particularly in abundance, frora the raore raesic to the raore xeric plant coraraunities. The desert shrub, creosotebush, and fourwing saltbush plots had the highest nuraber of lizard species, or highest species diversity (Pianka 1967), while mesquite and fourwing saltbush plots had the greatest lizard abundance (p<0.05). Of the individual plots, fourwing saltbush 131, 132, and mesquite 122 had the greatest lizard abundance (p<0.05), but of the three, only plot 131 had more lizards than mesquite 121, 123, and cresotebush 111 (p<0.05). The most common predatory bird of the lowlands was the loggerhead shrike (Lanius ludovicianus). Shrikes were observed on all low elevation study plots, and were coraraonly observed preying on lizards. Roadrunners (Geococcyx californianus) were uncoramon. The nuraber of snake species, most of them potential predators on lizards, also increased at the lower elevations. The predatory lizard G. wislizeni was present on the flatland plots. Pianka (1970) suggested the increase in predatory lizards, snakes, and birds led to greater predation pressure 129 7 . 6 _ cn QJ •H u o cn u 03 N •H 03 0 &H 2 - 1 - 0 10 06 09 05 01 16 11 12 13 Plant Community Cocie Fig. 57. Number o f lizard species, and the total number of lizards per hectare seen in the ten plant coraraunities. The coraraunities are arranged frora relatively raore raesi c to raore xeric (left to right). Plant coramunit ies are as follows: 01- grassland, 05-raountain shrub, 06-conifer, 09-pinon- juniper, 10-hardwood, 1 1-creosotebush, 12-raesquite, 13-fourwing saltbush, 1 6-desert shrub, and 17-quartz sand. 130 on southern lizards, while cliraatic factors imposed the greater pressure on the northern lizards. Abundance Variation in lizard abundance between months within plant communities was great. However, on plots in commu nities where many lizards were seen (i.e. creosotebush, raes quite, and fourwing saltbush), the trend was of raore lizards during the late suraraer raonths (Fig. 58). Lizard abundance increased (p<0.05) frora April to Septeraber in creosotebush, raesquite, and fourwing saltbush coraraunities. On the fourwing saltbush and mesquite plots, abundance estimates dropped during July, but then reached high levels in August and September. The July decrease in abundance can be attributed largely to the decrease in C. tigris in July, August, and Septeraber. U. stansburiana abundance also decreased soraewhat in July. The increases of August and Septeraber are due largely to the large nuraber of U. stansburiana juveniles and adults observed. Plots were not counted in late Septeraber or October, but visits to the areas in the latter part of October indi cated that the nuraber of active lizards had decreased substantially. C. tigris had the highest density (20/ha) of all lizards in creosotebush during July (Table 5). U. stansburiana also 131 12 ~"~ creosotebush mesquite f— f 10 /,. •••«•• fourwing saltbush '. o u // (d 4J overall mean !' u 8 OJ x: u 6 cn na u 03 N •H 4 / /^ ^ ^ 2 April May June July August Sept, Month Fig. 58. Monthly change in lizard abundance in creosotebush, raesquite, fourwing saltbush, and the raean for all plant coraraunites. (23 in ptrcs zq.j-Bnb CN 132 cn qnjqs q.j9sap "^ T^* "^ c 0 00 qsnqq.x-BS BuT/AJcnoj CN CN in u >1 •H c^ in cn 9q.Tnbs3ui CN t CD c un m U 0) fd Q in r- cn 4J qsnq3q.oso3j:o CN CN X) U fd cn • » CN CN OJ s x: cn pooAvpjE-eq m u in Q) a 00 c» u qnjqs UT^:4.unoui (U T^ Xi (U e Ti 3 ;3 puBXSS'ejB rH c 0 c; X cd X) U +J qsnqq-X^s Bumjcnoj CN Oi g •H •H (U cn X cn CN fd £1 0) aq.Tnbs9ni OJ g 0 Q *«^ 4-1 qsnq9q.oso9J0 rsj >sD CN o Q) 0 I CN 0 G •H C CO fd X pooAvpjreq 00 ^ cn 03 •^3 (d c ^ S :3 qnjn:[S UTPq.unoai 00 00 XI cn fd (U •H PU"BXSS"BJ6 ^ TJ o u cu fd a N cn •H H 0) JC cn M-l +J m cn 3 0 •H 3 4J j:5 3 +J 03 cn cn u cn CT> 03 H cn 3 0 •H 3 6 d C cn •H 1^ r-l 3 03 cn X c (U •iH u T> tsl 3 +J C CD OJ •H +J 4-> •H c c: H t3 03 03 tJ > H •H r-l c •H 0 cn cn cn cn •H cn 3 U g 3 3 3 3 4-> 1 0 3 SH V4 rH 1 1 ? cn Xi 03 0 0 0 0 3 • 3 cn cn g x: x: ^ x: LO 03 u 3 c 0 ft ft ft ft e r-l 0 ^ 03 cn 0 0 0 0 cn TU l •7TU l 3 H 3 4J 0 TTU l J t3 t3 w ft •iH Degenhardt (1977) discusses additional attributes of the modified-transect method. He found estimates derived using it to be comparable to findings of other researchers using different methods. The method has been used alraost exclu sively on flatland desert lizard populations. Its effective ness for estiraating abundance of species occurring in raore heavily vegetated habitats is not clearly understood, but values derived using it undoubtedly represent rainimal esti- mates there also. It is difficult to make interspecific comparisons of abundance as equal proportions of the merabers of different species raay not be active or equally observable. 135 Therefore, coraparisons were not attempted between different species. Relatively few density estimates are available for rep tile populations. It has only been within the last two decades that attempts to estimate reptilian densities have been made, and those reported pertain largely to lizards. Tinkle (1967) sxiraraarized sorae early studies in his extensive deraographic work on U. stansburiana. Tanner and Jorgensen (1963) found raaxirauradensitie s of a Nevada population of C. tigris of 53.8/ha. Milstead (1965) found 181 C. tigris/ha. Degenhardt (1966, 1977), using an index sirailar to the one used in this study and working in the Big Bend of Texas, found raaxirauraC . tigris densities of 10.4 and 12.6/ha, and suggested that Milstead's estimates were unrealistic. Turner et al. (1969) suggested that spring densities of C. tigris greater than 4 9.4/ha. would be remark able. They found spring densities from around 7.4-19.8/ha. in southern Nevada. The raean density estimates (Table 5) of this study fall in the lower range of these estimates. Tanner and Jorgensen (1963) estimated the raaxiraura den sity of U. stansburiana on a Nevada test site at 81.3/ha. U. stansburiana density estiraates were also established by Tinkle (1967) for populations in Winkler County, Texas, and Mesa County, Colorado. The Colorado population had 4 2 adults/ha. in 1965 and 24.7/ha in 1966. The Texas population 136 had a resident population of 34.6/ha. Because Tinkle caught each of his lizards, he was able to deterraine accurate age class distributions. He found populations of adults and juveniles on two plots over the 3-year period of 1961 to 1963 to be roughly 111-232/ha. The density estimates of this study for U. stansburiana are considerably lower (Table 5) than those found above. Density estimates are also available for £. undulatus. Tinkle and Ballinger (1972) recorded mean densities for adults and yearlings in Schleicher County, Texas, frora 1969 to 197 0 at 1.98/ha. Juveniles were not included. Ferner (1976) found 25-37 resident adults/ha in Boulder County, Colorado. Relative density estimates of this study (Table 5), including juveniles and adults, are intermediate to the above, but closer to those estimated for the Texas population by Tinkle and Ballinger (1972). Age Ratios Two species, U. stansburiana and C. tigris, had signif icant (p<0.05) increases in the number of juveniles seen in July, August, and September (Fig. 59). More juveniles were also seen of U. ornatus and C. tesselatus Cp<0.25), and qual itatively, other species as well in late suraraer. U. stans^ buriana adult abundance reraained relatively high throughout 137 60 h 50 - Uta stansburiana 40 - 30 c ^ 20 cn -S 10 u 03 2 N •rH MH 0 60 U 3 50 Cnemidophorus tigris 40 30 20 10 April May June July August Sept, Juveniles adults Fig. 59. Monthly changes in the relative pro portion of juveniles to adults of Cneraidophorus tigris and Uta stansburiana. 138 the summer, except in July, when the number of juveniles increased. The adults of C. tigris nearly disappeared frora the plots after a high of 5 9 in June, with a steady decrease to only one in Septeraber (Fig. 59). The increase in the ratio of juveniles to adults in the population is an obvious consequence of suraraer reproduction. The araount of this increase varies with the reproductive strategy of the species in a given area (Tinkle et al. 1970). Tinkle (1967) noted increased juvenile densities in mid to late summer in U. stansburiana. Tanner and Jorgensen (1963) also recorded increased juveniles of U. stansburiana in mid and late suraraer, while juvenile increases in C. tigris popu lations were raost dramatic in August and Septeraber. Juveniles seen in the spring may be attributed to the previous year' s young. Tinkle et al. (1962) found U. stansburiana populations in Texas to have essentially 100% annual turnover. Minimal annual survival in C. tigris populations have been recorded at 59% (Tanner and Jorgensen 1963) and 54 to 60% (Turner et al. 1969) . The disappearance of adult C. tigris is difficult to explain. Food is relatively abundant due to late suraraer rains and teraperatures do not change significantly. Tanner and Jorgensen (1963) reported inactivity by adults in August and Septeraber in Nevada. Pianka (1970) found C. tigris of the Mojave and Sonoran deserts active all suraraer, while more 139 northern populations were active in May and June and aes- tivated during July and August. Tanner and Jorgensen (1963) demonstrated lower natural mortality in aestivating than in nonaestivating individuals, probably due to freedom frora above ground predation (Pianka 1970). It appears that with the breeding and egg-laying periods completed earlier in the suraraer, the less active and nongrowing adults raay quickly build up fat reserves and reduce the need for continued activ ity with exposure to predation. The adults of other teiids, C. exsanguis, C. inornatus, and C. tesselatus, were also fewer in the late suraraer. In contrast, adults of the iguanid lizards G. wislizeni, £. undulatus, U. ornatus, U. stans buriana, and £. raodesturareraaine d active until later suraraer. Cover Certain plants are iraportant to lizards. They provide escape refuges, feeding stations, egg laying sites, and pro tection from intense insolation (Tinkle et al. 1962). The plants that lizards ran to when flushed, or cover plants, may not be the factor selected for by the lizard seeking cover. It may be using a burrow or rock under the plant, or the plant may simply be the closest cover available. How ever, lizards in GMNP almost always ran to a plant rather than an opening, and the plant was usually a shrub (Table 6). Exceptions to this were E. multivirgatus and U. ornatus. The 140 skink always crawled into leaf litter or dead vegetation, although in two cases this litter was at the base of trees. U. ornatus usually flushed frora perches at the top of bould ers to the sides of these boulders away frora the observer, although shrubs and trees were also used. Tinkle et al. tl962) found raesquite to be the raost iraportant plant used by U. stansburiana, partly because it afforded suitable sites for woodrat (Neotoraa raicropus)nest s which in turn provided shelters for the lizards. Few woodrat nests were observed in the honey raesquite plants on study plots during this study, but raesquite reraained an iraportant cover plant for U. stans buriana and C^. tigris. The diversity of plants selected raay be a reflection of the diversity of plants available to the aniraal. Thus the high cover index for S_. undulatus (Table 6) raay be a reflec tion of the high plant diversity of the raountain shrub, hardwood, and desert shrub coraraunities in which the species was found. Lizards occurring in the sarae coraraunities, how ever, would have the opportunity to run to similar cover plants. U. stansburiana and C. tigris occurred on the sarae study plots, but U. stansburiana used a greater diversity of cover than C. tigris (Table 6). S. undulatus and U. ornatus occurred on study plots in the sarae three coraraunities, and S. undulatus used considerably raore types of cover. Fewer U. 141 TABLE 6.—Lizards observed on plots raore than once, the cover they flushed to, and the diversity index for cover No Times % Species Cover Used Use Index (H) Gambelia wislizeni p_. incana 1 14 1.5498 L. tridentata 2 29 P. glandulosa 2 29 Y. elata 1 14 other 1 14 Sceloporus undulatus J. deppeana 2 8 2.5132 N. micrantha 4 17 0. engelmannii 1 4 D. leiophyllum 3 13 P. incanum 1 4 P. ponderosa 1 4 Quercus spp. 1 4 dead vegetation 5 21 opening 3 13 under rock 1 4 other 2 8 1 13 0.9795 Urosaurus ornatus £. deppeana N. micrantha 3 38 opening 4 50 4 2.076 P_. incana 6 Uta stansburiana 26 A. canescens 45 <1 B_. breviseta 1 4 C_. dioicus 7 5 E_. torreyana 8 12 7 L. tridentata 2 1 L. montanum 1 <1 Opuntia spp. 55 32 p. cjlandulosa 1 <1 giganteus 's. 7 4 X. sarothrae 17 10 7. elata 4 2 forb 4 2 opening 1 <1 other 142 TABLE 6—Continued No Times % Species Cover Used Use Index (H) Phrynosoma modestum L. tridentata 1 50 0.6932 A. canescens 1 50 Cnemidophorus exsanguis Quercus spp. 1 33 1.0976 R. aromatica 1 33 0. imbricata 1 33 Cnemidophorus inornatus L. tridentata 1 25 1.0397 P. glandulosa 2 50 X. sarothrae 1 25 Cnemidophorus tesselatus L. tridentata 6 67 0.8487 P. glandulosa 2 22 opening 1 11 1.5358 Cnemidophorus tigris P. incana 10 6 A. canescens 40 24 E. torreyana 3 2 L. tridentata 21 13 P. glandulosa 77 46 Y. elata 3 2 opening 9 5 other 3 2 33 1.3297 Eumeces multivirgatus J. deppeana 2 M. pauciflora 1 17 dead vegetation 1 17 leaf litter 2 33 143 ornatus were observed, however, and the diversity of plants used would be expected to increase up to a point as the nura ber of sightings increased. C. tigris ran extensively to the larger shrubs, partic ularly raesquite, while U. stansburiana used forbs and grasses as well as shrubs. Vitt and Congdon (1978) found that U. stansburiana used known escape routes, raoving only a short distance to the safety of a burrow while C. tigris used a flight response as a predator escape strategy. The observed flight patterns of U. stansburiana and C. tigris differed considerably. The uta would quickly run a short distance (x=1.84 ra) when first flushed, and then stop and remain motionless, usually at the periphery of a shrub, but often near grasses or forbs away from the larger shrubs. On sub sequent flushes, the lizard would run to a large shrub and/or enter a burrow. C. tigris were raore wary. When initially flushed, lizards ran further (x=7.0 ra) to large shrubs, where they would stop briefly before entering a burrow. The raost wary lizards, in terras of how close they could be approached before flushing, were C. tigris (x=5.7 ra) and G. wislizeni (x=4.6 ra). The overlap of cover used was calculated for sympatric species pairs. The value for C. tigris and U. stansburiana was high (0.94). Both species ran to honey mesquite, four wing saltbush, and creosotebush extensively (Table 6). The 144 C. tigris and G. wislizeni overlap was high intermediate CO.73) while overlap between G. wislizeni and U. stansburiana was low intermediate (0.49). S. undulatus and U. ornatus had an intermediate value (0.64), as did the C. tesselatus, C. inornatus pair (0.64). Ectoparasites The ectoparasites collected in 1978 from one species of amphibian and eleven species of reptiles were all chiggers (Order Acarina) of four species (Table 7). Most chiggers were found in the lateral throat folds of lizards. On E. multivirgatus, chiggers were in the groin and/or axilla region or between the digits. Teiid lizards had chiggers under the ventral scales, as did T. cyrtopsis. Chiggers were occasionally present in a lizard's post-femoral pocket, exter nal ear opening, or skin folds near the eye. The endodermal Hanneraania hylae occurred on the ventrura of R. berlandieri, most coraraonly in the pelvic region. Chiggers were found on lizards frora late April through Septeraber. Infestations varied frora one to over 100 chiggers per lizard, but raost involved less than ten chiggers. Eutrombicula alfreddugesi is a coraraon pest chigger of the area and occurs widely on reptiles (Loorais and Crossley 1963). This species was found on reptiles to the highest elevations. E. belkini was found on lizards frora Sraith 145 Spring and the lower west side of the park. Acoraatacarus arizonensis were taken frora a P. raodestura and two H. texana collected near Frijole Ranch, Signal Peak housing area, and Williaras Ranch House, respectively. Parasitized R. berlandieri adults were taken frora Choza Spring and South McKittrick Canyon. All chiggers are deposited in the Medical Zoology Col lection of The Museura of Texas Tech University. TABLE 7.—Chiggers found on amphibians and reptiles from Guadalupe Mountains National Park Hosts Chiggers Rana berlandieri Hannemania hylae Holbrookia texana Acomatacarus arizonensis Eutrombicula alfreddugesi Sceloporus poinsetti Eutrombicula alfreddugesi Sceloporus undulatus Eutrombicula alfreddugesi Urosaurus ornatus Eutrombicula alfreddugesi Phrynosoma douglassi Eutrombicula alfreddugesi Phrynosoma modestum Acomatacarus arizonensis Cnemidophomis exsanguis Eutrombicula alfreddugesi Cnemidophorus tesselatus Eutrombicula belkini Cnemidophorus tigris Eutrombicula belkini Eumeces multivirgatus Eutrombicula alfreddugesi Eutrombicula belkini Thamnophis cyrtopsis Eutrombicula alfreddugesi 146 Summary A study of the herpetofauna of Guadalupe Mountains National Park was conducted from March to October, 1978 and 1979. Seven species of amphibians and 35 species of reptiles were documented frora within the park boundaries. Eleven of these species were docuraented frora within the park for the first tirae. Eleven additional species, two araphibians and nine reptiles, were found near the park and undoubtedly occur within its boundaries. Five species were determined that possibly occur in the iraraediate area. Thirty-five 0.25- and 0.50-ha study plots were estab lished in grassland, raountain shrub, conifer, pinon-juniper, hardwood, cresostebush, raesquite, fourwing saltbush, desert shrub, and quartz sand coraraunities. Ten species of lizards were seen raore than once on 27 of the plots. More lizard species and individuals were found in the creosotebush, mesquite, fourwing saltbush, and quartz sand comraunities. Lizard abundance increased from April to Septeraber, with a general decrease in July. Density estiraates were raade for each species in each coraraunity. The raost abundant species were Uta stansburiana and Cneraidophorus tigris in raesquite and cresostebush coraraunities, respectively. 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Sorae recent changes in the herpeto fauna of the northern Chihuahuan Desert PP;^5^13-522. S t'e'sy^po^Lrof'he'BiolScal^esi^^ ChihuahuIS^Desert Region, United States and Mexico Nat. Park Serv. Trans, and Proc. Ser. No. 3. 658 pp. 155 scudday J. F. and J. R. Dixon. 1973. Diet and feeding behavior of teiid lizards frora Trans-Pecos, Texas. Southwest. Nat. 18:279-289. xt^a;,. Seifert, W._ 1978. Geographical distribution: Hylactophrye augusti latrans (barking frog). Herp. Rev. 9:61. Shannon, C. E. 1949. The raatheraaticaltheor y of coraraunica- tion. In Shannon and Weaver (eds.). The raatheraatical theory of coraraunication. University of Illinois Press, Urbana. 125 pp. Smith, H. M. 1946. Handbook of lizards. Corastock Publ. Co., Ithica. 557 pp. Snedecor, G. W. and W. G. Cochran. 1967. Statistical Methods. Iowa State Univ. Press, Araes. 5 93 pp. Stebbins, R. C. 1951. Amphibians of western North America. Univ. Cal. Press, Berkeley. 539 pp. . 1954. Araphibians and reptiles of western North Araerica. McGraw-Hill Book Co., New York. 523 pp. . 1966. A field guide to western reptiles and araphibians. Houghton Mifflin Co., Boston. 279 pp. Tanner, W. W. and C. D. Jorgensen. 1963. Reptiles of the Nevada test site. Brighara Young Univ. Sci. Bull. Biol. Ser. 3:1-31. Taylor, E. H. 1935. A new species of the genus Euraeces frora New Mexico. Univ. Kans. Sci. Bull. 22:219-223. . 1936. A taxonoraic study of the cosraopolitan scTncoid lizards of the genus Euraeces, with an account of the distribution and relationships of the species. Univ. Kans. Sci. Bull. 23:1-643. Tinkle, D. W. 1967. The life and deraography of the side- blotched lizard, Uta stansburiana. Misc. Publ. Mus. Zool. Univ. Michigan 132. 182 pp. Tinkle, E. W. , D. McGregor, and S. Dana. 1962. Home range ecology of Uta stansburiana steinegeri. Ecology 43: 223-229. 156 Tinkle, D W. and R. E. Ballinger. 1972. Sceloporus demographdemoqraphy; of af V^lizard^ ^'. "^Ecolog" intraspecifiy 53:570-584c comparativ. e Tinkle, D. W. , H. M. Wilbur, and S. G. Tilley. 1970 Evolutionary strategies in lizard reproduction.* Evolution 24:55-74. '^'"'"''^'/y l'\^' ^k ^^^i^^' J- R- Lannora, Jr., and G. A. Hoddenbach. 1969. A deraographic analysis of fenced populations of the whiptail lizard, Cneraidophorus tigris, m southern Nevada. Southwest. Nat. 14:18 9- 202. Vance, T. 1978. A field key to the whiptail lizards (genus Cneraidophorus) , Part 1: the whiptails of the United States. Bull, of the Maryland Herp. Soc. 14:1-9. Vitt, L. J. and J. D. Congdon. 1978. Body shape, reproduc tive effort, and relative clutch raass in lizards: resolution of a paradox. Ara. Nat. 112:595-608. Worthington, R. D., and E. R. Arvizo. 1974. Western records of the Davis Mountains kingsnake, Larapropeltis raexicana alterna, in Texas. Southwest. Nat. 19:329-340. Wright, J. W. and C. H. Lowe. 1968. Weeds, polyploids, parthenogenesis, and the geographical and ecological distribution of all-feraale species of Cneraidophorus. Copeia 1968:128-138. Zweifel, R. G. 1968. Reproductive biology of anurans of the arid southwest, with eraphasis on adaption of erabryos to temperature. Araer. Mus. Nat. Hist. Bull. 140:1-64. APPENDIX: LIST OF SPECIMENS 157 158 The following is a list of specimens collected in or near Guadalupe Mountains National Park, Texas. Fourteen collections are represented from the 4 0 United States institutions queried. Species arrangement follows Conant (1975) . Distances are given in kilometers via the shortest highway when a highway is present; other distances are air line kilometers unless otherwise stated. Collection data on specimens collected during this study are available frora the herpetology section of the Museum of Texas Tech University. Institutional abbreviations follow Duellman et al. (1978). Other abbreviations include BCB (Bryce C. Brown, personal collection), CCNP (Carlsbad Caverns National Park), JSA (John S. Applegarth, personal collection), SM (Strecker Museum, Baylor University), SRSU (Sul Ross State University), TTU (Texas Tech University), and UTEP (University of Texas at El Paso). Location terminology and distances follow 7 .5 minute U.S. Geological Survey topographical maps (1973) or locally accepted names. Two of the following locations may be con fused. Specimens listed from Bear Canyon and McKittrick or Bear Canyon Ranger Station with the exception of TTU nurabers, refer to Laraar Canyon and the Ship of the Desert Ranger Sta tion, respectively, just southwest of the mouth of McKittrick canyon, or ca. 9.6 km northeast of Pine Springs. TTU speci- is refer to Bear Canyon 1.6 km north of Pine Springs. men; 159 Ambystoma tigrinum Culberson County: Guadalupe Mtns. (USNM 46188); Bowl Tank, (TTU R-9663-67, R-9669-70, R-9744-46). Soldier Curve, 32 km SW Pine Springs (TTU R-9668) . Scaphiopus couchi Culberson County: 3.2 km E Hudspeth County line CTNHC 20178-82; 85 km N Van Horn (UMMZ 126289); 2.4 km E McKittrick Ranger Station, Parker's Tank (USNM 147869, UMNZ 123553, CCNP 2641); 19 km NE Pine Springs (UMMZ 126289) ; 9.6 km SW Pine Springs (TTU R-9626) ; 48 km N Van Horn (TTU R-9631). Hudspeth Coiinty: 12.8 km S Dell City (TTU R-9620-21) ; 16 km S Dell City (TTU R-9622-23) ; 21 km S 4.8 km E Dell City CTTU R-9624-25) ; 21 km S Dell City (TTU R-9627-28) ; 12.8 km W Hwy. 54, on Hwy. 62/180 CTTU R- 9629-30); Cottonwood Tank, E base of Patterson Hills (TTU R-9738-40) . Scaphiopus bombifrons Culberson County: 56 km N Van Horn (TCWC 396) . Hudspeth County: White Sands, 21 km E Dell City (SRSU 994-5); 16 km S Dell City (UMMZ 124082). Scaphiopus multiplicatus Culberson County: 9.6-14.5 km SW Pine Springs CTNHC 20190-94); 3.2 km E Hudspeth County line, Hw. 62/180 (TNHC 20183); 3.2 km SE McKittrick Ranger Station, Bear Canyon (USNM 147868); 2.4 km E McKittrick Ranger Station, Parker's Tank (UMMZ 123554, CCNP 2639); 84 km N Van Horn (TCWC 40366) ; 85 km N Van Norn (UMMZ 126290) ; 19 km NE Pine Springs 160 (UMMZ 123557); 4.8 km SW Pine Springs (TTU R-9642) ; 21 km N Van Horn (TTU R-9645-48) ; 45 km N Van Horn (TTU R-9649) ; Bowl Tank CTTU R-9650- 53); McKittrick Ent. Rd. , 0.8 km NW Hwy. 62/180 CTTU R-9654) ; Ship of Desert Ranger Station CTTU R-9655); Bowl Tank CTTU R-9716, R-9720) ; Cox Tank CTTU R-9717-18, R-9723-24); Coyote Peak Tank CTTU R-9725); Indian Meadow Tank, Dog Canyon (TTU R-9719). Hudspeth County: 12.8 km S Dell City (TTU R-9641) ; 1.6 km S Dell City (TTU R-9643); Culberson-Hudspeth County line, Hwy. 62/180 (TTU R-9644); Salt Lake, Ed Hammock Ranch (TTU R-9721-22) . Bufo cognatus Culberson County: Near Signal Peak, Hwy. 180 CSRSU 254); 81 km N Van Horn (SRSU 257) ; 58 km N Van Horn (TCWC 40163) ; 85 km N Van Horn (UMMZ 126286) ; 48 km N Van Horn (TTU R-9615) ; 45 km N Van Horn (TTU R-9616) ; 32 km N Van Horn (TTU R-9617) ; 13 km N Van Horn (TTU-R-9618); 9.7 km SW Pine Springs (TTU R-9619) . Hudspeth County: 21 km E Dell City (SRSU 996); 16 km S Dell City (UMMZ 124081) ; 21 km S 4.8 km E Dell City (TTU R-9611) ;3.2kmE3.2kmS Dell City (TTU R-9612-14) Bufo speciosus Culberson County: 84-km N Van Horn (TCWC 40252); 85 km N Van Horn (UMMZ 126285); McKittrick Ent. Rd. 0.4 km NW Hwy. 62/180 CTTU R-9672) ; McKittrick Ent. Rd. , 0.8 km NW Hwy. 62/180 (TTU R-9673) ; Hwy. 1108 55 km NW Orla (TTU R-9674). Hudspeth County: 16 km S Dell City (TTU R-9671) . 161 Bufo punctatus Culberson County: 11-15 km SW Pine Springs (TNHC 20199-203); Mouth Bear Canyon, Ranger Station CUMMZ 121852); 2.4 km E McKittrick Ranger Station, Parker's Tank CUMMZ 123555); Bear Canyon 3.2 km SE McKittrick Ranger Station (CCNP 2638); McKittrick Canyon, mouth (UMMZ 122975); 85 km N Van Horn (UMMZ 126287); Marcus Cabin, West Dog Canyon (TTU R-8221) ; 1.6 km E Culberson-Hudspeth County line, Hwy. 62/180 CTTU R-9636) ; McKittrick Ent. Rd. parking lot CTTU R-9637) ; 12.8 km NE Jet McKittrick Ent. Rd., Hwy. 62/180 (TTU R-9638) ; 1.6 km SW Pine Srpings (TTU R-9639) ; Jet. McKittrick Ent. Rd.-Hwy. 62/180 (TTU R-9640); Lower Guadalupe Spring (TTU R-9727-28, R-9731); North McKittrick Canyon (TTU R-9730); 0.4 km W Nipple Hill CTTU R-9732-33) ; Bone Spring CTTU R-9734, R-9736- 37); South Fork Shumard Canyon CTTU R-9735) . Hudspeth County: 21 km S 4.8 km E Dell City CTTU R-9635) . Bufo debilis Culberson County: 2.4 km E McKittrick Ranger Station, Parker's Tank (UMMZ 123556); Bear Canyon, Ranger Station (CCNP 2640); 85 km N Van Horn (UMMZ 126288) ; 45 km N Van Horn (TTU R-9632) ; 6.4 km N Van Horn (TTU R-9633) ; 63 km NW Orla, Hwy. 1108 (TTU R-9634) . Hudspeth county: 4.8 km W Culberson-Hudspeth County line CTNHC 20163-77). Rana berlandieri Culberson County: 11.n 3-5 kVTTm, SqWw Pine i^prxuySprings (TNHC 20206) ; Manzanita fr-,u,rjt'7 i9iq-?3 123551 BCB 8828); spring (TNHC 16307); McKittrick Canyon (UMMZ 121933, 162 Bear Canyon, near mouth (CCNP 2657); Pool in Bell Canyon, Hwy. 62/180 bridge CUNM 23595-96); South McKittrick Canyon Cmj R-9278, R-9657, R-9660, R-9662, R-9741-42); North McKittrick Canyon CTTU R-9659); Middle Mckittrick Canyon CTTU R-9743); Choza Spring (TTU R-9656, R-9658, R-9661); Frijole CUMMZ 70125). Hudspeth County: 4.8 W Culberson-Hudspeth County line, S Hwy. 62/180 (TNHC 17766-76). Kinosternon flavescens Culberson County: Pine Springs, Hwy. 62/180 (TTU R-9699) . Terrapene ornata Culberson County: 1.6 km W Pine Springs (TNHC 17954-55); 9.7 km SW Pine Springs (TTU R-9695) ; 3.2 km NE Pine Springs CTTU R-9696) ; McKittrick Ent. Rd. , 0-4 km NW Hwy. 62/180 (TTU R-9698) . Hudspeth County: 21 km E Dell City (SRSU 4340); 800 m SW Eclipse Well (TTU R-9697) Coleonyx brevis Culberson County: Guadalupe Canyon, 8 km SW Pine Springs CUMMZ 125300); 4.8 km N Pine Springs (BCB 8843). Hudspeth County: Culberson-Hudspeth County line, Patterson Hills (SRSU 4864); 4.8 km W Culberson-Hudspeth County line, S Hwy. 62/180 (TNHC 17953). Crotaphytus collaris Culberson County: Hw. 62/180 rest stop, below, S of El Capitan (SRSU 3269); 1.6 km NW Pine Springs (TNHC 16301-02); 3.2 km NE Nickel 163 (KU 72131-32); 1.6 km SE McKittrick Ranger Station (USNM 147871); 11.3 km SW Pine Springs, 3.2 km W Hwy. 62/180 CTTU R-9520) . Hudspeth County: 21 km E Dell City CSRSU 4274). Gambelia wislizeni Culberson County: 103 km NE Van Horn (TCWC 39921) . Hudspeth County: Between Abies and Eclipse Wells, 23 km E Dell City (SRSU 4282-83); 4.8 km W Culberson-Hudspeth County line, S Hwy. 62/180 (TNHC 17951); 500 m SE Lewis Well (TTU R-9516) ; Lewis Well (TTU R-9517) ; Gypsum Dunes W Eclipse Well CTTU R-9518) . Holbrookia texana Culberson County: 1.6 km NW Pine Springs CTNHC 16292); 2.4 km NE Pine Springs (TNHC 16308) ; Guadalupe Pass (AMNH 71039); Near McKittrick Ranger Station (USNM 147872); Frijole (UMMZ 125303); Bear Canyon, McKittrick Ranger Station (CCNP 2651; UMMZ 123540); 13.7 km S Jet. Hwy. 62/180-Hwy. 54 (JSA 3219-20); Patterson Hills, East Rim CTTU R- 9521); 1.6 km WSW Williams Ranch House CTTU R-9523) ; Guadalupe Pass, 8.05 km SW Pine Springs CTTU R-9524); Guadalupe Canyon (UMMZ 70079). Hudspeth County: Patterson Hills (SRSU 4289). Holbrookia maculata Hudspeth county: Abies Well Area (SRSU 4311-13); Ne^ Lynch's Tank, ca. 12.9 tan E Dell City (SRSU 4813); White Sand Dunes, 21 loa E Dell City (SRSU 4814, TTU R-7419-20) . 1.2 tan W Eclipse Well, Gypsu. Sand Dunes (TTU R-9525). 164 Sceloporus poinsetti Culberson County: Hw. 62/180 rest stop, below El Capitan (SRSU 3648); Bowl (TNHC 16288, 20184-85); 1.6 km NW Pine Springs (TNHC 16303-05); Mouth Pine Springs Canyon (TNHC 16309) ; 8-16 km NE Pine Springs CTNHC 20212); Guadalupe Mtns., Signal Peak CKU 15055-57); McKittrick Canyon (KU 62892); 3.2 km NE Nickel (KU 72215-18); Mouth McKittrick Canyon (TCWC 378-79, 535, CCNP 2650, TTU R-9482, UMMZ 123538); Pratt Lodge, McKittrick Canyon (UMMZ 121753); North McKittrick Canyon CUMMZ 122966, TTU R-9481) ; 4.8 km N Pine Springs (BCB 8698); Pool in Bell Canyon, downstream from Hwy. 62/180 bridge (JSA 2336); Marcus Cabin, West Dog Canyon (TTU R-8220) ; 31 km E Dell City (TTU R-8751, 8756-57); McKittrick Ridge (TTU R-9479); Williams Ranch House CTTU R-9480); Choza Spring (TTU R-9483) ; 3.2 km NW Pine Springs (TTU R-9484) ; Frijole (UMMZ 70084-85). Hudspeth County: Patterson Hills (SRSU 4277). Sceloporus magister Culberson County: Hwy. 54, 13.7 kia S Hwy. 62/180 (JSA 3218); Hwy. 54, 2.3 tatlS . Hwy. 62/180 (JSA 3216-17). Hudspeth county: Near Abies and Eclipse Wells (SRSU 4293-94). Sceloporus undulatus Culberson County: McKittrick Canyon (SRSU 469, 472-74, 476-77, TCWC 26129-30, UMMZ 121741-42, TTU R-9485) ; Guadalupe Mtns. (SRSU 761); H-. 62/180 rest stop, below El Capitan (SRSU 3319, 3331); 1-6 ^ NW Pine springs (TNHC 16298-99); 3.2 ^ NW Pine Springs (TNHC 20229-30); 3.2 165 km NE Nickel (KU 72704-08); 3.2 km E Nickel Creek CTCWC 25889); Mouth Bear Canyon CUMMZ 121743); South McKittrick Canyon CUMMZ 121744, TTU R-9493) ; North McKittrick Canyon (UMMZ 122968); Bear Canyon CCCNP 2652); 31 km E Dell City CTTU R-8750, R-8752, R-8754, R-8758, R-8761-62) ; Bowl CTTU R-9486); Upper Bear Canyon. (TTU R-9487) ; Dog Canyon CTTU R-9488) ; Bone Canyon, above Spring CTTU R-9489-90) ; Marcus Cabin, West Dog Canyon (TTU R-9491) ; 0.8 km S Lost Peak (TTU R-9492) ; Choza Spring (TTU R-9494) ; 0.8 km NE Cox Tank, West Dog Canyon (TTU R-9495) ; Guada lupe Canyon (UMMZ 70089-90). Urosaurus ornatus Culberson County: 3.2 km NNW Pine Springs CTNHC 20219-21, 20223-28); Bowl (TNHC 16286-87); Frijole CKU 17843-44,UMMZ 70097); Near Signal Peak CKU 15525-26); McKittrick Canyon (USNM 147873, TCWC 26144, UUMZ 122964-65, TTU R-9566); North McKittrick Canyon (UUMZ 121728, TTU R- 9563, R-9565, R-9567); South McKittrick Canyon CUMMZ 121729-30); 31 km E Dell City (TTU R-8753, R-8759-60) ; Bone Canyon, near Spring (TTU R- 9555-57); Dog Canyon, above Ranger Station (TTU R-9558-60) ; 1.6 km W Pine Springs (TTU R-9561); Shumard Canyon (TTU R-9562) ; Guadalupe Canyon CTTU R-9564); Guadalupe Canyon CUMMZ 70098). Hudspeth County: Patterson Hills (SRSU 4315). Uta stansburiana Culberson County: 91 km N Van Horn CTCWC 40042); Hwy. 54, 2.3 km S Hwy. 62/180 (JSA 3213-15); South McKittrick Canyon, 2.4 km above Pratt Lodge (TTU R-9554). 166 Hudspeth County: Between Abies and Eclipse Wells (SRSU 4297); 2.4 km NE Gypsum Sand Dunes (SRSU 4299-4300); 4.8 km W Culberson-Hudspeth county line, Hwy. 62/180 CTTIHC 17738, 11765, 20207-11); 4.8 km W Salt Flat, Hwy. 62/180 CTCWC 36845-46); 0.8 km W Jet. Hwys. 62/180 and 54 CJSA 2337, 2358); 4.8 km SW Eclipse Well CTTU R-9545-46) ; 2.4 km SE Eclipse Well CTTU R-9547) ; 1.6 km E Eclipse Well CTTU R-9548) ; 3.2 km SW Eclipse Well CTTU R-9549-50) ; 0.3 km W Eclipse Well CTTU R-9551) ; Gypsxam Sand Dunes, W Eclipse Well (TTU R-9552) ; Gypsum Sand Dunes, SW Lewis Well (TTU R-9553). Phrynosoma cornutum Culberson County: Pine Springs (TCWC 380); 1.6 km N McKittrick Ranger Station (UMMZ 125309); 9.7 km SSW Pine Springs CTTU R-9534) ; McKittrick Canyon Ent. Rd. (TTU R-9535-36, R-9538); Frijole (UMMZ 70139). Hudspeth County: Between Dell City and Guadalupe Mtns. (SRSU 4281); NE Corner Gypsum Sand Dunes, 21 km E Dell City CSRSU 4284); 2.4 km E Jet. Rd. 1576 and Hwy. 62/180 (JSA 3409); 0.8 km S Eclipse Well (TTU R-9537). Phrynosoma douglassi Culberson County: Bowl (TNHC 16285, 20186, TTU R-9539-40,. BCB 11573); Guadalupe Mountains (USNM 32984); Marcus Cabin, West Dog Canyon (TTU R-8219) ; Near Pine Top Mtn. (TTU R-9541) ; West Dog Canyon CTTU R-9542) ; Lost Peak (TTU R-9543) ; West Rim, Dog Canyon (TTU R-9544) . 167 Phrynosoma modestum Culberson County: 3.2 km NE Nickel (KU 72194-96); West Dog Canyon CKU 61494); valley W Guadalupe Peak CUSNM 32999); Mouth McKittrick Canyon CUMMZ 121707, 123544); Patterson Hills, East Base of West Riiu CTTU R-9526); Near Ship of Desert Ranger Station CTTU R-9527) ; Williams Ranch House CTTU R-9528) ; 9.7 km SSW Pine Springs CTTU R-9531) ; Pine Springs CTTU R-9532). Hudspeth County: 2.4 km S Lewis Well (TTU R-9529) . Cnemidophorus exsanguis Culberson County: McKittrick Canyon (SRSU 468, 3641, KU 62891, TCWC 25665-70, UMMZ 121772-75, 122963, TTU R-9505-06) ; 3.2 km NE Nickel CKU 72242-48); Mouth McKittrick Canyon (TCWC 536, TTU R-9514); 9.7 km NE Pine Springs (UMMZ 125323, 123524); Bear Canyon, near Green McCombs (CCNP 2648); Guadalupe Mtns. (SRSU 3642); Bone Spring, Bone Canyon (TTU R-9503) ; Upper Dog Canyon CTTU R-9504) ; 1.6 km NW Pine Springs (TTU R-9507); Patterson Hills (TTU R-9508, R-9515) ; Near Williams Ranch House (TTU R-9509); South Mckittrick Canyon, 2.4 km above Pratt Lodge (TTU R-9510) ; 0.8 km NW Pine Springs CTTU R-9511) ; Guadalupe Canyon CTTU R-9512) ; 3.2 km NNW Pine Springs CTNHC 20222); Frijole (UMMZ 70074). Hudspeth County: Culberson-Hudspeth County line, Patterson Hills (SRSU 4239). Cnemidophorus inornatus Culberson County: 11-15 tan SW Pine Springs (TNHC 20204); 3.2 tar. NE Nickel (KU 72259-81); 9.7 tan NE Pine Springs (UMMZ 125319); Bear Canyon 168 9.7 km NE Pine Springs (CCNP 2647^. TT I ^ y K^^NF 2647); 31 km E Dell City CTTU R-8755); 0.4 km S Williams Ranch House ^TTTT P Qt;Dc\ -, ^ House CTTU R-9586); 3.2 W Williams Ranch House (TTU R-9587). Hudspeth county: Hwy. 62/180, 4 tan W Jet. with Rd. 1437 (UTEP 1882, 1564); Hwy. 62/180, 35 tan E Jet. with Rd. 2317 (UTEP 1880-81); 4 tan WNW Williams Ranch House CTTU R-9585) . Cnemidophorus tesselatus Culberson County: Hwy. 62/180, rest stop, below El Capitan (SRSU 3310); 11-15 km SW Pine Springs (TNHC 20205); 1.6 km NNW Pine Springs (TNHC 16310); 3.2 km NE Nickel (KU 72308-24); Bear Canyon CUSNM 147870, UMMZ 123523); 9.7 km NE Pine Springs (UMMZ 125324); Bear Canyon, McKittrick Ranger Station (CCNP 2649); Patterson Hills, East Rim (TTU R-9496) ; 1.6 km W Williams Ranch House (TTU R-9497) ; 100 m N Williams Ranch House (TTU R-9498) ; Wash below Indian Cave CTTU R-9499) ; 3.2 km WNW Williams Ranch House (TTU R-9500); Guadalupe Canyon, Hwy. 62/180 (TTU R-9501). Hudspeth County: Culberson-Hudspeth County line, Hwy. 62/180 (SRSU 3344); Culberson-Hudspeth line, Patterson Hills CSRSU 4238, 4273); North end of West Rim of Patterson Hills (TTU R-9502) . Cnemidophorus tigris Culberson County: 91 km N Van Horn (TCWC 39666-67); 81 km N Van Horn CTCWC 39665); 1.6 km N Hwy. 62/180-Hwy. 54 Jet. CTTU R-9579) ; South base of East Rim of Patterson Hills (TTU R-9582) . Hudspeth County: Between Abies and Eclipse Wells (SRSU 4235, 4251-52, 169 4262-63); Guadalupe Mtns. Nafl Par)c boundary, E Abies Well (SRSU 4236, 4272); 2.4 tan NE Gypsu. Sand Dunes (SRSU 4264-65); 21 tan SW Pine swings (UMMZ 125321); 4 tan W Jet. Hwys. 62/180 and 54 (UTEP 1543-47); 4.8 tan NW Williams Ranch House CTTU R-q5fift ^Q^ n ^ -, tiiu K yb68-69) ; Guadalupe Mtns. Nafl Park boundary, SW Eclipse Well (TTU R-9S7n ii\ o ^ i r, ^ . • i- '=-Lj. ^liu K-yb/o-77); 2.4 km E Eclipse V7ell CTTU R-9578) 3.2 km S Lewis Well CTTU R-9580) ; 1.6 km S Lewis Well (TTU R- 9581); 3.2 km SSW Lewis Well (TTU R-9583) ; 400 m SW Lewis Well (TTU R-9584). Eumeces obsoletus Culberson County: 1.6 km NW Pine Spring (TNHC 16296); Guadalupe Mtns. (USNM 32829); North McKittrick Canyon (UMMZ 121720, 122971); Pratt Lodge, McKittrick Canyon (UMMZ 123525-26, CCNP 2654); Hunter Picnic area. South McKittrick Canyon CTTU R-9588) ; 2.4 km NNE Pine Springs (TTU R-9589); Frijole CUMMZ 70101). Eumeces multivirgatus Culberson County: McKittrick Canyon CSRSU 465-66, 475, UMMZ 121716-17, 123530-31, 123533, 125314); South McKittrick Canyon (CCNP 2653, TTU R-9591-92, R-9594); Narrows, South McKittrick Canyon CTTU R-9590) ; North McKittrick Canyon (UMMZ 122969-70); Bowl (TCWC 457) ; Head of Bear Canyon (UMMZ 123532, SM 6144); Bear Canyon, 9.7 km NE Pine Springs (UMMZ 123534); Smith Spring (TTU R-9593) ; Bowl, 3.2 km NNW Pine Springs (TNHC 16289-91); Bowl, 4.0 km NW Pine Springs (TNHC 17993-180::, 20214, SM 6286); Frijole (UMMZ 70516-18). 170 Leptotyphlops dulcis Culberson County: 1.6 tan NW Pine Springs (TNHC 16300); Jet. area, McKittrick Canyon (UMMZ 124078, 125326); McKittrieJ. Ranger Station, Bear Canyon (SM 6287). Thamnophis cyrtopsis Culberson County: McKittrick Canyon (SRSU 1869, USNM 147880, TCWC 26159, 381, 523, UMMZ 123499, TTTJ R-9595, R-9597) ; South McKittrick Canyon (UMMZ 121808, CCNP 2642); North McKittrick Canyon (TTU R-9600) ; 1.6 km NW Pine Springs (TNHC 16295); 3.2 km NE Nickel CAMNH 73015); Guadalupe Mtns. Nafl Park CAMNH 110457); Bear Canyon, 9.7 km NE Pine Springs (UMMZ 123500); Choza Spring (TTU R-9596) ; Cox Tank, West Dog Canyon (TTU R-9598); Little Sandy Wash, East Rim, Patterson Hills (TTU R-9601). Diadophis punctatus Culberson County: McKittrick Canyon (TCWC 26170, UMMZ 122948, 122955, 123481-85); South McKittrick Canyon (UMMZ 121825, CCNP 2644); Jet. area, McKittrick Canyon (UMMZ 124553, 125351-52); Mouth McKittrick Canyon (TTU R-9609) ; 6.4 km S Signal Peak (KU 72753); 4 km NE Pine Springs (TCWC 40051); 0.8 km above Pratt Lodge, South McKittrick Canyon (TTU R-9607) ; 0.8 km NW Pine Springs (TTU R-9608) ; 2.4 km NNE Pine Springs (TTU R-9610). Masticophis flagellum Culberson County: 16 km SW Pine Springs (USNM 147877); 4.8 km S Pine Springs (UMMZ 122939) ; 22 km NE Pine Springs (TTU R-9677) ; McKittrick Ent. Rd., 3.2 km NW Hwy. 62/180 (TTU R-9678) . 171 Masticophis taeniatus Culberson County: 3.2 km NE Nickel (KU 72759-60); McKittrick Canyon (USNM 147878, UMMZ 123469); Bear Canyon, McKittrick Ranger Station CUMMZ 123514); 8 km NE Pine Springs (CCNP 2643); 10.5 km ENE Pine Springs, Hwy. 62/180 (TTU R-9675); Frijole CUMMZ 70134). Salvadora grahamiae Culberson County: 3.2 km SE McKittrick Canyon (KU 62889); McKittrick Canyon (USNM 147879); Frijole (UMMZ 123475, 70138); Bear Canyon, McKittrick Ranger Station (UMMZ 124555, CCNP 2646); Mouth Shirttail Canyon (TTU R-9603) ; Marcus Cabin, West Dog Canyon (TTU R-9604) ; Bone Springs (TTU R-9606). Elaphe guttata Culberson County: Near Pratt Lodge, McKittrick Canyon (USNM 147876); 3.2 km N Pine Springs (UMMZ 123502); Pine Springs (TTU R-9689-90) ; Frijole (UMMZ 70497). Elaphe subocularis Culberson County: 11.3 tan SW Pine Springs, Hwy. 62/180 (TTU R-9693-94) . Arizona elegans Culberson County: 9.7 km SSW Pine Springs (USNM 147874); 16 tan SW Pine springs (UMMZ 123505); 15.3 km SW Pine Springs (UMMZ 125328); 12.9 tan SW Pine Springs (UMMZ 123507); 11.3 tan SW Pine Springs (TTU R- 9691); 8.8 tan SSW Pine Springs (TTU R-9692) . Hudspeth county: 29 tan SW Pine Springs (UMMZ 123503). 172 Pituophis melanoleucus Culberson County: North McKittrick Canyon (UMMZ 122982); 8 km S Pine springs (UMMZ 123501); Bear Canyon, main house (UMMZ 124028); 8 : NE Pine Springs CUMMZ 125339); 9.7 km SW Pine Springs (UMMZ 125340); 24 km SW Pine Springs (UMMZ 125341); Pine Springs CTTU R-9602) ; Frijole CUMMZ 70135). Lampropeltus getulus Culberson County: 16 km SSW Pine Springs CUMMZ 124079); 11.3 km SW Pine Springs CUMMZ 125337). Lampropeltus mexicana Culberson County: Ease Side Guadalupe Peak, Guadalupe Mtns. (UTEP 542); 4.8 km S Pine Springs .(UMMZ 123494). Rhinocheilus lecontei Culberson County: Jet. Hwys. 62/180 and 54 (TTU R-9681) ; 26 km SW Pine Springs, Hwy. 62/180 (TTU R-9682) ; 3.2 km S. Hwy. 62/180 on Hwy. 54 (TTU R-9683). Sonora semiannulata Culberson County: North McKittrick Canyon (UMMZ 122944); South McKittrick Canyon (UMMZ 122945); McKittrick Canyon, Jet. area (UMMZ 124080, 125345). Gyalopion canum Hudspeth county: 119 tan E El Paso (UMMZ 123487) ; 18 tan S Dell City (TTU R-9684). Hypsiglena torguata Culberson County: 68 tan N Van Horn (TCWC 40059); 9.7 tan SW Pine springs (TTU R-9685) ; 9.7 tan NE Pine Springs (TTU R-9686) . Tantilla nigriceps Culberson County: 9.7 tan SW Pine Springs (TTU R-9687). Hudspeth County: 40 tan SW Pine Springs (UMMZ 123493); 4.8 tan W Salt Flat (TTU R-9688). Tantilla atriceps Culberson County: Guadalupe Mtns. (SRSU 2018); 3.2 km NE Nickel (KU 72769-71); McKittrick Canyon (TCWC 25980, UMMZ 122946, 123491-92); McKittrick Canyon, Jet. area (UMMZ 124554); 3.2 km E Nickel Creek (TCWC 25904); Bear: Canyon, 1.6 km SE Ranger Station (UMMZ 123490); Bear Canyon, Green McCombs (CCNP 2645); Guadalupe Canyon, 8 km SW Pine Springs (UMMZ 125347). Crotalus atrox Culberson County: Signal Peak (KU 14170); 16 km SW Pine Springs (UMMZ 123464); Bear Canyon, mouth (UMMZ 123465); 2 km SW Pine Springs (JSA 2359); McKittrick Canyon Ent. Rd. (TTU R-9707-08) ; 11 km SW Pine Springs (TTU R-9711) . 13 km SW Pine Springs (UMMZ 122942) ; Bear Canyon, McKittrick Canyon Ranger Station (CCNP 2656). Hudspeth County: 24 km SW Pine Springs (USNM 147899). 174 Crotalus scutulatus Culberson County: 52 km N Van Horn (TTU R-8008) ; 58 km N Van Horn (UMMZ 91468) ; 18 km SW Pine Springs (SRSU 4662) . Hudspeth County: 5.2 km W Salt Flat (SRSU 4663); 24 km S Dell City (UMMZ 123466); 21 km S. 0.4 km E Dell City (TTU R-9704) ; 2.4 km E Salt Flat (TTU R-9710) . Crotalus viridis Culberson County: 42 km N Van Horn (UNM 22342); 8 km SW Pine Springs (TTU R-9701); Dog Canyon, Ranger Station (TTU R-9702). Hudspeth County: White Gypsum Sands, 21 km E Dell City (SRSU 4329); 400 m SW Eclipse Well, Gypsum Dunes (TTU R-9700) . Crotalus molossus Culberson County: Guadalupe Pass, Guadalupe Mts. CAMNH 70990); Bowl CTCWC 383); J. C. Hunter Ranch (TCWC 10290); McKittrick Canyon (BCB 8890-91); South McKittrick Canyon (UMMZ 121913, 122981); Pratt Lodge, McKittrick Canyon (UMMZ 122631); Bear Canyon, 9.7 km NE Pine Springs (UMMZ 125357); Dog Canyon, Ranger Station (TTU R-9706) ; 12.9 km SW Pine Springs (TTU R-9705); Near Pine Springs CTNHC 20148). Crotalus lepidus Culberson County: West Side Bartlett Peak CUTEP 2544, 2548); NNE side Guadalupe Peak (TNHC 20145); Bowl(TNHC 20147); Near Pine Springs (TNHC 20148); Pratt Lodge, McKittrick Canyon (UMMZ 122630); South McKittrick Canyon (UMMZ 121837); North McKittrick Canyon (TTU R-9703); McKittrick Canyon (BCB 8835).