Brigham Young University Science Bulletin, Biological Series

Volume 7 | Number 1 Article 1

5-1965 A comparative population study of small vertebrates in the uranium areas of the Upper Colorado River Basin of Wilmer W. Tanner

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Recommended Citation Tanner, Wilmer W. (1965) "A comparative population study of small vertebrates in the uranium areas of the Upper Colorado River Basin of Utah," Brigham Young University Science Bulletin, Biological Series: Vol. 7 : No. 1 , Article 1. Available at: https://scholarsarchive.byu.edu/byuscib/vol7/iss1/1

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JAN 2 8 1966 HARVARD UNIVERSITY BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN

A COMPARATIVE POPULATION STUDY OF SMALL VERTEBRATES IN THE URANIUM AREAS OF THE UPPER COLORADO RIVER BASIN OF UTAH

by

Wilmer W. Tanner

Biological Series — Volume VII, Number 1

MAY, 1965

BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN

A COMPARATIVE POPULATION STUDY OF SMALL VERTEBRATES IN THE URANIUM AREAS OF THE UPPER COLORADO RIVER BASIN OF UTAH

by

Wilmer W. Tanner

Biological Series — Volume VII, Number 1

MAY, 1965 tviuS. COMP. ZOOL LIBRARY JAW HARVARD UNIVERSITY

PREFACE

This paper constitutes a terminal report to the United States Atomic Energy Commission for contract AT(11-1)819. It provides not only a faunistic and floristic Study of areas having a high and low surface radiation, but also includes a con- sideration uf their general ecology. Major emphasis is placed on the variation nl the external characteristics of the small vertebrates of these areas with a statistical analysis of those characters examined in the lizards. The study was conducted in southeastern Utah (Tanner 1963, see map) from October 1959 through 1964, and was conducted as a test of the variation effects of natural surface radiation on the external characteristics of small vertebrates. TABLE OF CONTENTS

INTRODUCTION 1 PROCEDURE AND ACKNOWLEDGEMENT 3 GENERAL ECOLOGICAL BACKGROUND 6 Climate 6

GENERAL DESCRIPTION OF THE AREAS 8 Temple Mountain Mesa 8 Mamie Stover Area 10 Yellow Cat Area 10 Indian Creek Area 10

REVIEW OF THE OBJECTIVES 10 Objective 1 10 Cold-blooded Vertebrates 11 Warm-blooded Vertebrates 11 Plant Species and Vegetation Analysis 11

Objective 2 13 Home Range 15 Longevity 15 Growth 16

Objective 3 17 Objective 4 17

LIZARD SPECIES STUDIES 18 Uta stansburiana 18 Sceloporus undulatus elongatus 20 Cnemidophorus tigri? septentrionalis 23

MAMMALS 25 Ammospermophilus leucurus notium 26 Etttamias quadrivittatns hopiensis 26 Neotoma lepida sanrafaeli 26 Feromijscus maniculatus sonariensis 26 Peromyscus maniculatus nebrascensis 26 Peromyscus crinitus 27

STATISTICAL ANALYSES OF THE DATA 27 CONCLUSIONS 30 LITERATUDE CITED 31

LIST OF ILLUSTRATIONS

Figure Page

1. General view of the Temple Mountain study area, as viewed from north to south 2

2. General view of the Mamie Stover area, showing a small ravine which cuts through the overlying cap rock 4

3. Dorsal view of hands and feet of a Sceloporus undulatus elongatus showing the num- bers assigned to the digits 4

4. Six views of the toe marks as recorded on the rocks. All taken at Temple Mountain .... 5 \ diagramatlc sketch oi the Temple Mountain area showing the location of the home ranges ol sum,- Uta stansburiana. The legend provides the basic information. See first foldout

6 \ diagramatic sketch oi the Temple Mountain area, showing the location of the home ranges oi Sceloporus undulatus ehngatus. Tlic legend provides the basic information Sec second foldout

7. Number ol scale rows at midbody in six populations oi Uta stansburiana. The range of

variation is indicated by the horizontal line, the mean l>\ the vertical line, the rectangle encloses one standard error on each side of the mean and the shorter horizontal line

above the vertical line I mean) represents one standard deviation of the mean. The num- ber of scales (rows) is indicated both at the bottom and the top of the graph, with the V(>1 populations (areas | to the left of the range ( long horizontal line) and size of the population sample on the right margin of the range 19

8. Dorsal scales in Uta stansburiana counting from parietal to base of tail. See Fig. 7 for explanation of graph 19

9. Ventral scales in Uta stansburiana counting from the enlarged scales just posterior to

the gular fold (this is approximately at the interior square of the front legs) to the granule scales at the vent. See Fig. 7 for explanation of graph. 19

10. Total number of femoral pores on both femurs in Uta stansburiana. See Fig. 7 for ex- planation of graph. 20

11. Total number of supralabial scales on both lower lips in Uta stansburiana. See Fig. 7 for explanation of graph 20

12. Total number oi infralabia] scales on both lower lips in Uta Stansburiana. See Fig. 7 for explanation of graph. 20 13. Number of scale rows at midbody in Sceloporus undulatus ehngatus. See Fig. 7 for explanation of graph 21

14. Dorsal scale rows in Sceloporus undulatus ehngatus, counting from parietal to base of tail. See Fig. 7 for explanation of graph. 21 15. Ventral scales in Sceloporus undulatus ehngatus counting from the enlarged scales just posterior to the gular fold (this is approximately at the anterior square of the front legs) to the granule scales at the vent. See Fig. 7 for explanation of the graph. 21 16. Total number of femoral pores on both femurs in Sceloporus undulatus ehngatus. See Fig. 7 for explanation of the graph 22

17. Total number of supralabial scales on both upper lips in Sceloporus undulatus elon- gatus. See Fig. 7 for explanation of graph 22

18. Total number of infralabial scales on both lower lips in Sceloporus undulatus ehnga- tus. See Fig. 7 for explanation of graph 22

19. Number of scale rows at midbody in six populations of Cnemidophorus tigris septen- trionalis. See Fig. 7 for explanation of graph 23

20. Dorsal scales in Cnemidophorus tigris septentrionalis, counting from parietal to base of

tail. See Fig. 7 for explanation of graph. 23

21. Ventral scales in Cnemidophorus tigris septentrionalis, counting from the enlarged scales

just posterior to the gular fold (this is approximately at the anterior square of the front legs) to the granule scales at the vent. See Fig. 7 for explanation of the graph 24

22. Total number of femoral pores on both femurs in Cnemidophorus tigris septentrionalis. See Fig. 7 lor explanation of graph 24

2-3. Total number of supralabial scales on both upper lips in Cnemidophorus tigris septen- trionalis. See Fig. 7 for explanation of graph. 25

24. Total number of infralabials scales on both lower lips in Cnemidophorus tigris septen- trionalis. See Fig. 7 for explanation of graph 25 A COMPARATIVE POPULATION STUDY OF SMALL VERTEBRATES IN THE URANIUM AREAS OF THE UPPER COLORADO RIVER BASIN OF UTAH

INTRODUCTION

The discovery of uranium and vanadium The explorations since 1945, however, have ores in the area of Utah and been much more extensive and intensive, thus Colorado near the turn of the century has fin- providing a far greater knowledge of the size, ally led to a mining operation far beyond the quality, depth and extent of ore bodies through- dreams of those early prospectors who made out the entire plateau producing area of Utah their first ore shipments in 1902 (Sorensen: and the adjoining states of Colorado, New Mex- 282). In this early period of development of ico, Arizona and Wyoming. the area the ores were important because of the By the enormous numbers of claims through- radium that could be extracted. Although the out the area, the prospecting boom of the early amounts obtained were small and the process 1950's demonstrated that uranium is widespread of extraction was slow, tedious, and expensive, throughout the area. Although most of the es- radium was extracted from Utah ores until 1923, tablished claims did not become producing with the greatest production occurring during mines because the ore was mostly of a low the last ten years. During these few years the grade, the great extent of the natural ground Colorado Plateau area of Utah and Colorado level radiation particularly in "hot" areas was was the leading radium-producing area of the demonstrated. world. However, the processing of these ores Before leaving this aspect of the study it was marginal and the discovery of high-grade should be pointed out that three of the areas stu- ores in the Belgian Congo in 1923 made further died have been mined at various times for the operations too expensive. past sixty years and are still producing, although During the first fifty years of development not to any extent, from the same shallow de- the area actually passed through three rather posits used in the early days. A general descrip- distinct mineralogical stages which may be rep- tion of each area will serve as an orientation resented as follows: (a) Radium to 1923. A and as an introduction for necessary additional sidelight of these early explorations is the ship- details. (Also see Hayward et. al. 1958.) ping of carnotite ores from the The first field work on this contract was fossiliferous deposits to Pierre and Marie Curie, begun on June 1, 1960, with two graduate as- in Paris, France, in 1903 (Boutwell 1905:207). sistants and me; thus the present investigations At this same time the new yellow mineral was include four years of field work. It should be named Carnotite in honor of the French physi- indicated at the outset, however, that most of cist Adolph Carnot. During this period the re- the field work has been done and must be done covery of radium from the uranium ores placed during the summer months, primarily because this area in world prominence and produced we are dealing mostly with the cold-blooded more than 32 grams of radium in 1921. (b) Van- vertebrates which are not available in these adium, 1923 to 1944, and (c) uranium, 1944 to areas for more than about six months of the present (Sorensen, 1963). In contrast to the pre- year. ceding period, during the present period van- The role of isolation in this inland desert adium has been reclaimed as well as molyb- basin must have played its part in providing an denum even though uranium is now the primary opportunity for variation to occur. The fact that product. A history of the development of this so much variation does occur in most species of mining industry cannot be discussed further; this area suggests that perhaps other factors however, for those interested the following stu- may be operating. Few species occurring in the dies provide some of the historical facts and Upper Colorado Basin are of the same sub- cite additional references on this subject: Bout- species as those inhabiting the basins to the well 1905, Sorensen, 1963, Hilpert and Dasch west (Great Basin), to the southeast (Upper

1964. The latter is particularly valuable because Rio Grande) or to the south (Little Colorado). of the large bibliography included in that study. In fact, some variation has been observed be-

1 )

I Si BfUGHAM YOUNC NIVERSm l) \. I Hi I I I i l\

populations occupying opposite sides ol inhabiting these areas Thus it was proposed

( iorado Hiver in Utah. thai several areas in the • ppei olorado Speciation, but more particularly subspecia- with a higher-than-average surface radiation be tion, has occurred regularly in the several val- studied and thai local populations of small ver-

leys extending northward from the southern tebrates lizards, and it possible the small mam- I deserts. Such areas as Death Valley and thos< mals) be compared for externa] morphological basins extending north, Lowei Colorado Basin variations. It was further deemed necessary to and the Virgin Valley in southwestern i tah, compare populations occurring in the .ocas ot adjoining Arizona and Nevada and S. California high surface radiation (25 to 100 mr./hr.) and the l pper Colorado Basin are examples. with populations in the nearby or adjacent areas

One example of subspeciation found in this with a lower surface radiation | less than 20). area is Arizona elegans with Candida in the These comparisons consist of a series ot char- Death Vallej .u

ed and others discovered or recognized only re- gross anatomy or histological comparisons is not cently, the Upper Colorado Basin lias been of included. Although physiological (including ser- special interest to us, leading directly to the ological ) and genetical ( chromosomal cytology request for research funds to further investigate studies may have been fruitful, the project was certain possible causes (Fig. 1). not designed to include internal cell or tissue

It was hypothesized, therefore, that perhaps variations. the numerous small areas of high surface radia- The question of what is an unusual or note- tion may be playing their parts by inducing var- worthy variation and when are such variations iation into the populations of small vertebrates significant in small populations has been a ma-

l-'n. l. General view ol the Temple Mountain studs area as viewed From north to south. Biological Series, Vol. 7, No. 1. May, 1965 jor challenge throughout the course of this in- radiation in some areas of the San Luis Valley vestigation. At the outset we postulated that of southern Colorado. Ecological studies now in there would be some variation and that these progress at Arco, Idaho, and Dugwav, Utah, are might take the form of, (a) an increase in the primarily concerned with controlled induced range of variation of a character within a single radiation on native animals. Those ecological population or, (b) a skewing of the mean to studies in the Atoll areas of the Pacific, at the the right or left of the means of other popula- Nevada test site, and other natural and labora- tions. tory areas represent environments where nuc- lear introduced radiation. In We realized at the outset that there was no reactions have certain way of determining the amount of radi- neither the ecological nor the many laboratory ation received by any one animal during a sea- and field-controlled studies have the experiment- ers dealt with the major problems considered son or its life, and were therefore obligated to determine the general radiation for the area. in this study. From this we hoped to determine the average During the first year four areas were select- dosage for the animals included in the areas ed, each of which is associated with an ore studied. Therefore, measured radiation listed in body and with an increase in surface radiation Table 12 represents the approximate dosage re- over adjoining areas. These were designated by the will ceived per hour. Because there is uneven radi- following names and appear in this ation, with some areas serving as pockets of high report as named here: Temple Mountain Mesa, radiation and others of low, only the approxi- Mamie Stover Incline, Yellow Cat area and In- area. first mate or general area average is considered. dian Creek The two are located 45 and 15 miles respectively southwest and west of Although there have been many and varied Green River, Emery Countv, Utah. The Yellow studies carried forward to determine the effects Cat area is approximately 15 miles southeast of of induced radiation on the inherited morpho- Thompson's, Grand Countv; and Indian Creek is logical characteristics of vertebrates, little is about 35 miles southwest of La Sal Junction, actually known about the effects of natural rad- San Juan County, Utah. In the latter area only iation on populations. A study by Daniel and the reptiles are included in the variation studies Blair (1959) has indicated an increase of soil (Fig. 2).

PROCEDURE AND ACKNOWLEDGEMENT

Several techniques were used in gathering being included in the home-range studies. In the field data and to collect the animal samples. each species the toe clip method was employed. The mammals were caught in live traps, frozen An illustration of how the toes were counted in a field dry ice cabinet and returned to the is seen in Figure 3, After the first twenty lizards University for mounting. Standard weights and (Ufa) were marked, two toes were clipped, measurements were taken, and the skins and such as 1 and 12 and the next series 2 and 15. skulls were prepared as is usual in museum col- Each lizard caught was marked, measured and lections. Previous studies (Brant 1962, Burt 1940, the sex determined. Care was taken to release

Calhoun and Casby 1958, Mohr and Stumpf each lizard in the exact area where it was caught 1964, and Tonaka and Teramura 1958), seem- so that established home ranges would be less ingly provide adequate home-range data to in- likely upset. dicate that the species inhabiting these areas Lizards were caught by hand rather than by did not need additional home-range study. How- traps. This proved successful particularly with ever, in the lizards I could not be sure; and Uta and Sceloporus but was difficult for Cnem- since the areas under consideration are relativ- idophorus. In order to determine lizards and the ely small, it became mandatory that the extent exact spot taken the toe marks were recorded of annual movement, at least, be determined for in the field book and at the same time painted

Sceloporus undulatus elongatus, I 'la standsbur- on a rock. Figure 4. Thus a toe clip of five on the iana stansburiana and Cnemidophorus tigris scp- front foot and of 17 on the hind foot was record- tentrionalis. These species were common to all ed as 5-17. The painted marks on the rocks areas studied. These were intensively studied proved relatively permanent, thus making the only at Temple Mountain, the other areas not measurement of distances between points of re- Huh. ham ^01 s(. UmvEBSm St IKSCE Blixetin

a, showing a small ravine which cuts through the overlying

capture easy. Second captures were recorded the figure (4-11X): and for Cnemidophorus the in the same way but with a line underscoring X was circled, 4-1 1® the figures. Vta were recorded as indicated; The methods of home-range study used in Sceloporus were recorded with an X following this project were in part dictated by the rough, rocky terrain but also from the fact that the pit- fall trap method was being used for similar studies by Jorgensen and Tanner (1963) at the Nevada Test Site. Although the surface funnel traps have been used successfullv bv Fitch (1951 and I960) in Kansas, the near absence of shade to protect captive lizards from the intense heat argued against their use.

In all other areas lizards were collected by using dust shot in .22 caliber pistols. This meth- od proved highly successful in providing a rel- atively large series of specimens from the various .mas However, in most areas almost the entire population was taken in order to provide the needed series. At Temple Mountain because of the home-range study only a few lizards were taken until the summer of 1963. By shooting, a lew specimens were damaged for one or two of the standard counts made, as reflected in the

I tc I Dorsal view <>! hands and feet of a Scelo- variable numbers used in the variation graphs, porta unduiattu elongattu showing the numbers assigned to the digits. but the increase in specimens made available by Biological Series, Vol. 7, No. 1, May, 1965

Fig. 4. Six views of the toe marks as recorded on rocks. All taken at Temple Mountain. BHIGHAM Vol m. I m\ I Msll 1 Si II N. I Hi I I h I IN the method more than made up foi the few areas. In our radiation measurements, these were

IllSSfS. avoided. Thus our data | Table 1- i in< hides only

Iii ordei tu better evaluate the total en\ iron those leadings taken m the natuial undisturbed incut a careful analysis ul the kinds ol pi. mis habitat ol ea< h area studied. percentage percentage ol species in three areas, I am indebted to the following foi then as- of plant covei and soil depths were recorded. sistance and for man) liclplul suggestions dur- Hen .is with the lizards the plant species ing the ionise ol the project. These Graduate were Inst determined al remple Mountain and students have been responsible tor much ol the vari- tin i parisons were made to determine field woik: \V Gerald Robinson, Robert E. Bul- ations between the areas. The plant canopy lock, Lynn Findlay, Elden Willis, fames Hop- cover was determined liv following a transecl kin, Clyde Pritchett, and John Childs. Their and placing a painted pint frame over the plants devotion to the project, often under adverse des along even meter ol the transect. While meas- ert conditions, is greatly appreciated. soil depths were uring the plant canopy, the de- The plants were collected by Veil Mlman steel penetrometer at regu termined by using a and Clyde Pritchett. Mr. Allman did the clas- along transect. lar intervals the sification, with final designations of a few used Various instruments have been to de- species being made by Dr. Stanley Welsh of the tect an increase in ground level radioactivity. In Brigham Young University, Botan) Department. were tins Study, measurements ol radioactivity The mammals were examined b) Dr. Stephen D. made in the field with a Tracerlab gamma scin- Durrant. To him I am grateful tor helpful sug- 2" iodide tillation detector, employing a sodium gestions concerning the variations occurring be- was by port- crystal, The equipment powered a tween populations in these species. able 110-volt generator with a voltage regulator. I am grateful also to Dr. A. Lester Allen of The minimum counting time for each station was the Brigham Young University, Department ol ten minutes. The equipment was standardized Zoology and Entomology for the use ol equip- against known background rates at Provo, Utah, ment and for the radiation measurements. Inline and alter use in the field. Supplemental Thanks is also extended to the field officials ol measurements were made with a portable beta- permitted gamma survey meter (geiger). the Union Carbide Company, who \s previously indicated, some mining opera- us complete freedom ol movement while at tions, either active or abandoned, occur in the Temple Mountain.

GENERAL ECOLOGICAL BACKGROUND

Climate \s is suggested b\ the low relative humidity

of areas, there is sporadic precipita- The general climatic conditions for those desert only tion. Actually the climate is relatively moderate parts ol the Upper Colorado Basin included in but with seasonal extremes. Temperatures range this studs are similar to those ot the Great Basin to above in to F. and below of western Utah, which are at about 4,500 to LOOT, summer in winter; summer rains may be of the cloud- 5,500 feet in elevation. burst variety, providing lor heavy flooding, and There are four weather stations in the gen- heavy wind storms occasionally produce intense eral area i ll.mksville. Green River, Thompson's, dust storms. Based on 30 or more years ol Wea- and Moab). These, except lor Thompson's, are ther Bureau records lor stations at Lmcrv and at lower elevations than the Study areas and are Green River, Emery County; Hanksville, Wayne therefore noi completely representative ol the County; Moab and Thompson's; Grand County, climate. Although we have 1 not been able to Utah, the averages listed in Table 1 have been carry forward a year-round check of climatic summarized, conditions at the stmK areas, some summer Mr. E. Arlo Richardsen, State Climatologisl maximums and minimums were taken at Temple at the Weather Bureau \irport Station. Salt Lake Mountain, Both temperatures and humidity were City, Utah, was kind enough to provide similar obtained from an Abheon indicator placed in a averages lor the studv areas. These he calculated juniper tree about three and one hall leel above bv using the data available troin the nearest the ground surface and approximately at the weather station and used the differences in tie middle ol the ana vation, slope and sun exposure to provide an es Biological Series, Vol. 7, No. 1, May, 1965

Table 1. Temperature and precipitation averages for five weather stations in Emery and Grand Counties, Utah for 30 or more years.

Jan. Feb. Mar. April May June July Aug. Sept. Oct. Nov. Dec. Annual

Emery Bricham V(/i si. JmvERsm St iknce Bulletin during tin- year. Thunder storms during July influence oi die Rock) Mountains to die east .mil August provide more rainfall than Foi an) rado. othei period. Both Moab and Thompson's re In summary, the climatic conditions in the port more precipitation than the station al < Ireen general area where this stud) was made are

River and those stations to the west. This may surprisingl) similar, particularly if the annual reflect the Colorado River air drainage or the averages an- considered as the basic criteria.

GENERAL DESCRIPTION OF THE AREAS

mediately east After having carefully examined the selected to the i one mile from study area) areas, I became apprehensive of the role played is the San Rafael Desert. Leaching of desert soils by plant cover, soil types and soil depths. Each proceeds very slowly because of the low rainfall: of these may be a determining ecologieal factor hence, surviving species must be able to tolerate in any natural system where conditions might the highly alkaline soils usually present in such produce an increase of variation in a gene pool. anas The two plant communities most wide-

It is not seemingly possible to rule out the prob- spread in the Great Basin Desert are those in able effects which may be produced by such which sagebrush Artemisia tridentata Nutt. and environmental factors in disjunct but similar shadscale Atriplex confertifolia (Torr. and areas. Therefore, in order to establish the extent Frem.) S. Watts, are the dominant plants Atri- of physical and biological factors within the plex is found in the more alkaline soils. areas, we determined in a general way the plant The area slopes from north to south, with a species in each area and their approximate per- low hill on the western edge and the high reef centage of surface cover. also determined We to the east, giving the hillsides of the study the soil depth and general composition. Although area essentially a southeastern exposure. A fairly there are some slight variations, both in kinds of large wash drains the area from north to south, plant species found and the depth of the soil with smaller washes draining into it from the in antic, these may represent fluctuations which slopes to tlu- west and north. No measurements would be expected to occur as a result of random of the precipitation were taken within the study sampling. area, but records at the city of Green River Temple Mountain Mesa. The Temple Moun- nearby show an average precipitation of 5.78 tain area consists of a small ridge extending for inches falling throughout the year but with the approximately one thousand yards north and greatest amounts during the months of July, south and varying in width from approximate!) August, and September. Precipitation, however, one hundred yards to slightly over two hundred is very erratic in such environments and signifi- yards near its middle. This formation is adjacent cant variations occur from year to year. to and just west of the eastern uplift of the sand- Besides the dominant Atriplex confertifolia stone of the . The western edge the hillsides are covered with a great variety ol of the Temple Mountain area consists of a ledge herbaceous and shrubby plants. Many annual of white limestone-conglomerate cap rock clip- and perennial grasses are found in scattered ping to the east and forming a natural barrier clumps with Oryzojjsis hymenoides (Roem. and on the western edge. The narrow northern Schult. ) being more abundant but with ephem- boundary lies along a fault, with the area to erals abundant on the slopes in the springtime. the north uplifted and sloping up to the base The latter germinate, grow rapidly and mature of Temple Mountain. in the span of a few short weeks. These include This study site lies at an elevation of about various species such as I.cpidimii densiflorum

5,500 feet. It, as well as the Mamie Stover area, Schred., l.cpidium montanum Nutt . Streptan- lies along the eastern edge of the San Rafael thus longirostiris S. Watts., Euphorbia fendleri Swell. Both fall within the influence of a desert Torr. and Gray, Oenothera eaespitosa Nutt., climate similar to that of the Great Basin Desert Oenothera mutiifuga Watson. Oenothera walk-

that is. they are both characterized by low eri (A. Nelson). Cilia gracilis (Dougl. ) Hook, rainfall and atmospheric humidity, hot sunny Descurainia sophis (L.) Webb, Lappula redow- days, a great daily range of temperature, inter ska (Hornem.) Greene, Plantago purshU Roam, mittent streams, high salt content of the soil and Shult, Lygodesmia exigua (Nutt.) Greene, and other desert environmental features. Im- and others. (See plant list.) )

Biolocical Series, Vol. 7, No. 1, May, 1965

In addition to the dominant Atriplex there cator of selenium is confirmed by many writers are a number of other shrubs. Some of these since it is found to occur almost exclusively on are Ephedra viridis Coville, Cercocarpus intri- selenium bearing soils. This plant is common catus S. Wats., Cowania stansbariana Torr., throughout the area. Amelanchier utahensis Gatt. Pomac, Rhus tri- Even more significant to this study is the lobata Nutt. Torr. and Gray, Rhus utahensis work of Helen Cannon (1952). She found that Godding, Artemisia tridentata Nutt., Chryso- uranium ores of the Colorado Plateau contained thamnus nauseosus (Pall.), C. viscidiflorus not only uranium but also considerable quanti- (Hook) Nutt., Tetradijmia glahrata A. Gray, and ties of vanadium and selenium, and that these an occasional Opuntia rhodantha Schumann, elements were absorbed in unusual amounts by Gesamtb. the roots of plants growing in these areas. She Many species of plants typical of areas of states: "A much higher content of uranium has higher rainfall are found adjacent to desert been reported by the Canada Department of washes. Found chiefly near the wash in the Mines ... in plants growing in the vicinity of study area are occasional small trees of the Pinon- pitchblend deposits. Plants growing on the as- Juniper Association which is so typical of the in- phaltic sandstone deposits at Temple Mountain, termountain region, Pinus edulis Engelm, and Utah, and on limestone deposits at Grants, New Juniperus utahensis (Engelm). Confined mainly Mexico, also absorb uranium readily. The max- to the wash are other species such as Amelan- imum values are considerably higher than those chier utahensis Koehne, Gatt. Pomac, Fraxinus found in plants associated with carnotite de- anomala Torr. S. Wats., Rhus trilobata Nutt. posits of the Colorado Plateau." She gives (Table Torr. and Gray, Rhus utahensis Godding, Astra- 3) the following analysis of the Asphaltic sand- galus spp., Cowania stansburiana Torr., Salsola stone, Shinarump conglomerate, and organic kali L., Cercocarpus intricatus S. Wats, and Sym- materials of the Temple Mountain area and for phoricarpos oreophilus A. Gray. the carnotite ore of Yellow Cat study site. Significant among the plants of the area are Thus we have a variable in the amounts of those that serve as indicators. R. F. Nelson minerals taken up by plants. When one considers (1957) reports that "in every case investigated, the availability of uranium in plants which are Beath, Eppson and Gilbert found Stanleya pin- used as food either directly or indirectly (eaten nata growing on seleniferous soils in 11 western directly as in the case of rodents or indirectlv states." The use of Stanleya pinnata as an indi- as in the case of lizards eating plant-feeding in-

Table 3. 1 Analyses of Plants Growing on Uranium-bearing Roeks and Soils of the Colorado Plateau 1949-50°

U Se PPM

Ash Soil Soil Soil Locality and Sample (per or Plant or Plant or Plant cent rock ash rock ash rock ash

Temple Mountain, Utah Asphaltic ss. ore, Shinarump conglomerate 760 1,500 0.8 Wood in ore 4,900 7,800 3.0 Juniperus monosperma tops 4.0 66 350 n.d. Juniperus monosperma tops 4.0 100 470 n.d. Oryzopsis hymenoides tops 5.5 20 260 10 Stanleya pinnata tops 10.0 37 120 190 Sta. 1 Yellow Cat area, Thompson's District, Utah S. 2 Sandstone near carnotite ore "°P. 8 Oryzopsis hymenoides—tops 7.3 30 70 Oryzopsis hymenoides—roots 35.2 40 1,600 S. 3 Surface soil 5 ft. above carnotite ore 180 S. 4 Shale layer in ore 290 23,000 P. 10 Ephedra viridis—tops 14.7 20 P. 12 Atriplex confertifolia—tops 31.3 50 P. 56 Artemisia spinescens—tops 18.4 70 Artemisia spinescens—roots 13.2 100

'Adapted from Cannon 1952, Tables 6 and 8. "F.S. Grimaldi, Ruth Kreher, Claude Huffman, and F. N. Ward, chief analysts. °° Selenium-indicator plants BlUCKAM YODNC UNIVEBSm SCKNCl Hi i i mis

the problem ol determining radiation at the surface and actually provides for tin- high- source becomes more complicated and its effects est ground-level radiation found in any of the more difficult to determine, areas under investigation. The basic nature of

riw s\/r 11! tins area (approximately thirty- the cap rock provides only for small pockets for

ac .1 vegetation development other than is five res I has piw ided sufficiently large pop- what ulation dI small vertebrates to make a variation found in the larger ravines. Thus, there is an ul iv study possible. This has been the area in which inns bareness in this area which is not ap- in we nave conducted .ill of our studies dealing parent the other areas. Transects were taken with home range in the lizards. in the more vegetated areas and therefore the figures in Table 5 do not reflect Approximately two miles west of Temple accurately the area as a but rather Mountain are two areas represented bv small whole the vegetated pocket areas where the animals drainage ravines, both showing a reduced radia- live. tion count, which have been used for the secur- Yellow Cat Area. This area is approximately ing of comparative specimens. Each of these fifty-five miles east of the San Rafael Reef and anas, designated as upper and lower ranges, fifteen miles Southeast of Thompsons, Grand represents approximately one half of the area County, Utah. We have examined the area lying at Temple Mountain, and has been intensely south of the first (east) junction and west of collected. In both of these areas the ore-bearing the road along the base of a ledge area in strata have been eroded away. They are, how- which is found pockets of uranium and vana- ever, at approximately the same elevation as the dium ores. This area is approximately one half Temple Mountain Mesa because of the San Ra- mile long, but not more than one hundred yards fael Swell uplift and the accompanying erosion. wide at any place. Because of its basic geological Even though there are some slight differences nature and based on our data gathered at Tem- in the soils, it appears to support the same gen- ple Mountain, it is considered that the small ver- eral type of plant cover and to represent approx- tebrates are rather restricted to this narrow belt. imately the same percentage of plant canopy- The general area is less variable in altitude, with cover when compared with the Temple Moun- fewer high ridges and deep vallevs. The eleva- tain Mesa Area. tion is perhaps a few hundred feet below that of the other two areas (according to Cannon, Mamie Stover Area. This area is located in 4,900 ft. , but does support approximately the the Northeastern edge of the San Rafael Swell. ) same plant and animal species. Much of the but lies just east of the San Rafael Reef. Overly- data gathered by Cannon (loc. cit.) indicates ing this area is a cap rock varying from five to that approximately the same plant species occur seven feet in thickness. Embodied in the cap rock here as in the San Rafael Swell areas. are a series of fossil trees. Much of the radioac- Indian tive ore is associated with these fossils, and in Creek Area. This is a small area at some instances rather high grade pockets have the bottom of a rather large canyon. Erosion has exposed the ore-bearing strata so that they been mined. Erosion has cut a series of channels are exposed on each side of the small creek for through the cap rock. Other areas are partly a short distance. The greatest contrast here is eroded. This, plus the fact that small cracks and in the greater abundance of vegetation, particu- crevices have developed in the cap rock, has pro- larly along the stream. The apparent increase in vided for a high surface gamma radiation in this plant cover is seemingly reflected bv the more area (Table 12). In some areas where erosion dense lizard populations, particularly in Cnemi- has exposed the fossilferous material the ore is dophorus tigris.

REVIEW OF THE OBJECTIVES

In order to be more explicit, each of the above. Most of this will be included as a part tour original objectives will be summarized with of the first objective. as much detail as the data will permit. The ob- Objective 1. Determine kinds of populations jectives as originally listed have required some modifications. Perhaps the most significant oi of vertebrates occurring in these natural radia- these is the inclusion of the hasie plant ecology tion areas as compared with nearby nonirradi- and of soil depths for the major areas listed ated areas. (The word iwnirradiatcd should be )

jioLociOAL Series, Vol. 7, No. 1, May, 1965

changed to read low surface radiation, since all so complete at Temple Mountain that no small areas show some surface radiation. mammals were seen or trapped during the sum- mer of 1961 and onlv a few chipmunks were A. Cold-Blooded Vertebrates. Three species seen in 1962 and 1963. of lizards are relatively common throughout

the area ( Uta stansburiana subsp., Sceloporus C. Plant Species and Vegetation Analysis. undulatus elongatus, and Cnemidophorus tigris The vegetation was analyzed in all areas being septentrionalis). Other reptiles taken in limited studied in order to determine the dominant plant numbers are these: a few specimens of Crota- species within each area. phtftus colhnis, Crotaphytus wislizeni, Urosaurus The canopy-cover was determined by follow- ornata, Masticophus taeniatus and Crotalus vir- ing a transect and placing a pointed plot frame idis concolor. Amphibians have been found at over the plants along the transect every meter. Mamie Stover and at Yellow Cat. In both in- The purpose of using this method was to obtain stances only a few individuals of Bufo punctatus a two-dimensional evaluation of the influence and Bufo woodhousi were taken, and they are each plant taxon exerts over the other compon- not therefore being included in the project ents of its ecosystem. All of the individuals of study. one taxon that fall within the plot frame are con-

sidered as a unit. An imaginary line is drawn B. Warni-Blooded Vertebrates. Seven species around the leaf tips and projected onto the of mammals were found to be most abundant ground. The taxon is then placed into a cover in the study areas. Phylogenetically they are as class according to the percentage of the plot follows: Ammospermophilus leucurus, Eutamias frame the imaginary polygon covers. The re- quadrivittatus, Peromyscus crinitus, Peromyscus maining taxa within the plot frame are then maniculatus, Peromyscus truii and Neotoma le- pida. Other species taken but not in sufficient numbers to justify variation studies are Dipo- domijs ordii, Neotoma cineria, Lepus sp. and Table 4. Lists of mammal species taken at three of the study areas. Sylvilagus sp. Signs of large carnivores of both

canids and felids were observed in all areas. Species No. of Specimens A more detailed comparative studv of the TEMPLE MOUNTAIN MESA, EMERY COUNTY populations of each species will be made later; however, the skin and skull material available Ammospermophilus leucurus 18 Eutamias quadrivittatus 13 for study is listed in Table 4 for each area stu- Di)>odomys ordii 1 died. This material is now available at the Peromyscus crinitus 4 Brigham Young University Museum. Peromyscus maniculatus 20 Those listed from Temple Mountain Mesa P. truii 4 Neotoma lepida 8 were taken during the late summer and fall of 1960. They appeared to be abundant, and we were therefore selective, taking only a small sample of adults. For reasons not fully under- MAMIE STOVER. INCLINE, EMERY COUNTY stood by us, all small mammals disappeared dur- Ammospermophilus leucurus 2 Eutamias ing the winter of 1960-61. In June of 1961 we quadrivittatus 5 Peromyscus crinitus 6 did not trap a single mammal. An investigation Peromyscus truii 1 of the wood rat (Neotoma) nests revealed no Neotoma lepida 15 live inhabitants, but did provide several skele- tons. It is our guess that we experienced a local 29 crash cycle which eliminated the Temple Moun- YELLOW CAT MININC DISTRICT tain populations. We did not notice a reduction GRAND COUNTY in populations at other areas, although there Ammospermophilus leucurus was not a large population of any mammal spe- cies at Mamie Stover. Also of importance was the apparently com- plete elimination of all small mammals in the nearby upper and lower ranges. There are there- fore no comparative mammals from these areas. The geographical extent of this "crash" was ,not determined. It did not affect those popula- tions at Mamie Stover or at Yellow Cat, but was 12 Hltll. HAM ><>! M. I.MIHIMII S( l| M | BcLLETIN

iMni 5. Five dominant plants In the Temple Mountain, Mamie Stovet and Yellow Cat Areas.

Temple Mountain )

Biological Series, Vol. 7, No. 1, May, 1965

Lepidum montanum Nutt. the soil. The depth is measured to the nearest

Malcolmia africana ( L. ) R. Br. decimeter. Physaria australis (Payson) Rollins The relative stoniness was greatest at the Stanleya pinnata ( Pursh ) Britton Mamie Stover area; the average soil depth is 1.1 Streptanthella longirostris S. Wats. Mountain the average Euphorbiaceae. decimeters. At Temple Euphorbia fendleri T. and G. soil depth, 1.2 decimeters, was slightly greater Gnetaceae. than at Mamie Stover. The soil depth at Yellow Ephedra viridis Coville Cat was much greater than that in the two pre- Gramineae. vious areas. Here the soil depth was 2.9 deci- Agropijron inerme (Schribn. and Smith) Rydb. Agropyron smithii Rydb. meters. The minimum, maximum, mode and Aristida longiseta Steud. Syn. average soil depths are listed in Table 7.

Bouteloua gracilis (H.B.K. ) Lag. Bromus tectorum L. Tarle 7. Soil Depth Data for Temple Mountain, Hilaria jamesii ( Ton. ) Benth Mamie Stover and Yellow Cat Areas. Measurements are Oryzopsis hymenoides (Roem. and Schult.) Decimeters. Smith in Sitankm hystrix ( Nutt) J. G. Sporobolus crypUindrus ( Torr. ) A. Gray Temple Mt. Mamie Stover Yellow Cat Stipa columbiana Macoun. Stipa comata Trin and Rupr. Min. 0. dm. Min. 0. dm. Min. 0. dm. Stipa speciosa Trin. and Rupr. Max. 5. dm. Max 5. dm. Max. 9. dm. Tridens pulchellus (H.B.K.) Hitch. Mode 1. dm. Mode 0. dm. Mode 3. dm. Hydrophyllaceae. Ave. 1.2 dm. Ave. 1.1 dm. Ave. 2.9 dm. Phacelia crenulata Torr. Leguminosae. Objective 2. Determine the approximate Astragalus flavus ( Nutt. ) T. and G. Dougl., Hook Astragalus lentiginosus palans ( Jones. home range, seasonal distribution and migration Astragalus praelongus Sheld. of individuals in these areas. Loasaceae. In the area here designated as Temple Mentzelia albicaulis Doug. toe marked over four-hund- Malaceae. Mountain, we have Amelanchier utahensis Koehne, Gatt. Pomac red lizards. During the summer of 1960 (June 1 Malvaceae. to September 1) marked ones were also recap-

Sphaeralcea coccinea ( Pursh ) Britton tured in the course of capturing for marking. Nyctaginacese. However, most recaptures were made during Albronia elliptica A. Nels. dur- Allionia linearis (Pursh) the summer of 1961, with some being made Onagraceae. ing 1962 and 1963. Figures 5 and 6 provide the Oenothera caespitosa Nutt. approximate shape and size and indicates home Oenothera multijugas Wats ranges of a few Ufa and Sceloporus of the Tem- Oenothera walkeri (A. Nels.) Raven area. Oleaceae. ple Mountain Fraxinus anomala (Torr.) S. Wats. Tables 8, 9, and 10 provide data from which Pinaceae. the home ranges are determined. Other useful Engelm. Pinus edulis data (to be discussed below) such as size, rate Plantaginaceae. of growth and longevity are also indicated in Plantago purshii Roem. and Schult. Polemoniaceae. these data.

Gilia gracilis ( Dougl. ) Hook. Many Uta and Sceloporus were captured two Polygalaceae. or more times during the same summer and Polygala subspinosa S. Wats. found to have remained within a few feet or Polygonaceae. original point of capture. The data Eriogonum bicolor Jones yards of the Eriogonum fasciculatum Benth. gathered during the summers of 1960 and 1961 Eriogonum hookeri S. Wats. were generally confirmed by the report of Tinkle Eriogonum inflatum Torr. et. al. published in early 1962. Because of the Eriogonum pusillum T. and G. similarity between our data and that of Tinkle Rosaceae. plotted on Cercocarpus intricatus S. Wats. only the larger seasonal ranges were Cowania stansburiana Torr. the map. However, by 1962 it became evident Scrophulariaceae. that Uta of the Temple Mountain area were Penstemon utahensis /Eastw. longer lived than those reported by Tinkle; thus the home range studies were continued. What primary The penetrometer is a pointed steeel rod, seemed pertinent and obviously our about one-half inch in diameter and ten deci- concern was to determine if these lizards actu- meters long. A short pipe welded on one end ally did remain in a limited area during their

makes it possible to push the penetrometer into lifetime. Bricham Yoi m. i \i\ i. us 11 , s< u st h Bulletin

Table 8. Via ttatuburiana mbsp Two I Biological Series, Vol. 7, No. 1, May, 1965 15

Table 10. Sceloporus undulatus elongatus, Two and Three Catches Voi si. 16 Bhicham UmvEBsm Science Hi i 1 r 1 is

may in some individuals include .it least two Tins requires additional growth after hiberna- and perhaps three breeding seasons and include tion and provides for a mixed population of

.it least three to four years <>f lift-. adults and juveniles during the spring breeding In l'J'il the opportunity came to make simi- season. It is not until June that the population lar test-, at the Nevada Test Site, near Mercury, has reached a Dearly uniform adult status. lunty, Nevada. Under the supervision of Our observations indicate that mating oc- Allred and the direction of Clive D. D. M. Jor- < ins in April and May, much too early for juv- gensen pints were established for the purpose eniles to have reached sexual maturity. Our lim- of testing home range, growth and longevity. ited data in this area indicate that Uta do not The latter was demonstrated to be at least three mate until the second spring and may then live years for Uta in the populations of southern Nev- for at least one additional reproductive cycle; ada (Tanner and Jorgensen, 1963). perhaps some individuals live for a total of The three locations where these studies have three. been made (western Texas, southern Nevada An example is found in Table 1, Specimen and southeastern Utah) all vary as to elevation, No. 10, a male toe-clipped on June 9, 1960, and yearly temperature ranges, yearly and perhaps measuring 43.3 mm., recaptured on September also seasonal precipitation and other basic eco- 2, 1962, nearly 27 months later, at which time logical factors. However, they are all in desert it measured 49 mm., a growth of 5.7 mm. This areas with sparse vegetation and much exposed lizard was a young adult, hatched in late July surface of either sand or rocky surface areas. or August of 1959 and having been extant for at A general consideration of the elevation may least 37 months when last seen in September of have the greatest effects on the longevity, since 1962. According to our field notes this lizard a decrease to less than 2,000 ft. elevation in- was active and in apparent good health when creases the general range of temperatures; al- last seen. though this may, and undoubtedly does, make 3. Growth. A summary of the field data dealing for higher temperature during the summers in with growth and age in Uta is presented in Table Texas, it also extends the activity range for Uta 6. A comparison of these general data to those to ten or more months and for mild winters of Tanner and Jorgensen (1963) for the Nevada some activity throughout the year. The Temple test site are surprisingly similar and reveal that Mountain area is approximately 5.500 feet in ele- there is little ecological difference between the vation and several hundred miles north of either population in east central Utah and southern the Texas or southern Nevada study areas. At Nevada. this elevation and north latitude Uta are forced to hibernate for at least three and usually for Uta are the most abundant lizard in the areas four or five months. Some activity may occur studied. Although no attempt was made to de- in March and October; however, in general only termine density of any species population, there six months (April through September) provide appeared to be little difference in general abun- climatic conditions suited to full lizard activity. dance between the Temple Mountain, Utah and The effect of such climatic conditions in Utah Nevada Test Site populations of Uta. is to extend the reproductive cycle into the sum- There is undoubtedly some competition be- mer rather than the spring months. Eggs are tween the three species here considered. Scel- laid in late June and July. Most hatchlings ap- oporus and Cneniidophorus are both larger than pear in August. Thus only two, or possibly '2 Al Uta and may feed on larger insects, thus re- months, remain for growth before hibernation. ducing their competition with Uta. Scelopoms

Table 11. Size and Approximate Growth Hates for Uta stansburiana Summarized from Data Compiled over a Period of Four Years at Temple Mountain, Emery County, Utah.

Snout-Vent Date Age Group Length

I it. |nl\ and August Hatchlings is they emerge from eggs 21-25 mm. August Hatchlings approximate!) one month old 25-30 mm. Middle to Late September [uveniles six. weeks to two months old 31-35 mm. March and April [uveniles emerging from hibernation 36-39 mm. Ma) SubadultS reaching adult size !>nt not showing the nuptial color 38-41 mm. [une Subadults or adults 42-45 mm. July Adults 44-47 mm. 2nd Year Adults 45-51 mm. —

Biological Series, Vol. 7, No. 1, May, 1965 17 inhabits the rocky sidehills and ledges, whereas Objective No. 4.—The fourth objective was Cnemidapharus is in the depressions and along to determine the degree of external morpholog- gullies where there are fewer rocks and open ical variation existing in populations inhabiting spaces of sand and eroded soil. Thus the size of areas of natural radiation and to compare these food items taken and the ecological niche in- with each other and with populations occurring habited by these species reduces the competi- in nearby areas having a lower surface radiation. tion for food and space. The data presented in the following graphs Because these three species live together have been taken from those external character- at least Uta is found throughout the ranges of istics for which a more exact statistical analysis the larger species— it was suspected that the could be made. Therefore, scale counts in lizards larger species may prey upon the smaller Uta. are used for the graphs and a discussion of color

To determine this a thin metal sheet fence of and color pattern is provided for the lizards and about 2Ji feet high was set up in the Temple small mammals. Mountain area, and a known number of individ- Measurements were taken for all species ex- uals of each species was placed inside. The amined. These data are, however, dependent on fence surrounded several small brush and an such variables as age and available food and open sandy area. Rocks were placed in a loose thus were not included as graphed data. An pile for added shelter. For nearly a month the analysis of growth and size is provided for Uta lizards were fed insects gathered from an alfalfa and Cnemidophorus in the Nevada Test Site field some miles away. The insects varied in study by Tanner and Jorgensen (1963). These size and kind and were readily taken by the data correspond closely to that observed in this Uta and Sceloporus. Cnemid&phorus was shy study. Although growth and size were not ma- at first and one refused to eat. By controlled jor considerations, some information has been feeding we could be assured that the larger liz- included in previous discussions or will be in- ards were hungry, yet neither of the larger liz- cluded in subsequent discussions under each ards preyed on the Uta. species. Objective No. 3—The third objective was to Before considering the variations in lizards, establish the amount of radiation in the various it should be emphasized that in the areas stu- areas under consideration. died only a few snakes (four Crotalus and two The ground surface radiation count of gam- Masticaphis) were seen during the four years ma and beta rays was made at various times of field work. Thus adequate data for such a since June 1, 1960. Tests were made at Temple study are available only for the lizards. Mountain, three mLles west of Temple Mountain, An examination of reptiles in general and and at Mamie Stover during August 1960. In Sauria in particular shows that these as a group September 1961 tests were made at Yellow Cat have scale patterns which are used as standard and in May 1962 Indian Creek was tested and characteristics in determining their classifica- rechecks were made at the other areas. In order tion. I have used six of these scale counts in that all radiation measurements would be figur- gathering the variation data: dorsal scales, from ed the same, all tests were made by Dr. A. Les- parietal to base of tail; ventral scales, beginning ter Allen and me. Table 12 provides the avail- with enlarged scales at or near anterior margins able data. of front legs to the last enlarged scales anterior

Table 12. Radiological Survey (mr/hrs). Units: X background (11 mr/hr, at Provo, Utah)

Test Area Location Bun. I i \M YOUNC I'm. I km II SCIENCI Hi I I Ills tn Miit, scale rows around midbody; total fe- TABLE 13. Ana Code Numbers fur Uta stans- buriana* moral pores; supralabials, counting each side separately; and infralabials, counting each side A-01 Temple Mountain Mesa separately. These characteristics are graphed \-o_: Lowei remple Mountain Range tin . .M li spri irs and tin each population. A-03 Upper Temple Mountain H.inge

\ oi SI The Uppei Colorado River Basin "I I tah Mamie \-

I ta an- coded as ol. 02, 03, etc. Table 13 will provide The six graphs prepared for each species | the code information for all graphs, species and stansburiana, Sceloporus undulatus and Cnemi- areas. dophorus tigris) will be briefly discussed as Each graph provides the Following informa- each species is considered. A general considera- tion: the area, siieh as A.-01; the range ol vari- tion of variation, as it pertains to the lizards and ation, indicated In- the long horizontal line, and small mammals, will then summarize the results the mean by the vertical line above the range. of the project.

1 T/AKD SPECIES STUDIED

Uta stansburiana stansburiana Baird and Cirard lor the six external morphological characteristics used in the analyses of ("/<;. Each one will be The Northern side-blotched lizard inhabits briefly reviewed. As indicated in Table 12. the most biotic communities in the Upper Colorado area coded as A-01, A-Of. A.-05 and V.-06 were River basin From the low river valleys up to the the stuck areas with a higher surface radiation. funiper-Pinyon pine Forests. In this general area The other two were adjoining A-01 and had a Uta is the most abundant of the lizard species lower surface radiation. Hereafter only the code and is more commonly seen in the brushy habi- numbers will be referred to rather than a re- tat than on the ledges or rocky talus slopes. peating ol localities. Our Field records indicate that of the three lizards studied Via is the most active. That is. in- Results: 1. The scale rows as analyzed in dividuals leave hibernation earlier in the spring Figure 7 do not provide significant differences and seek it later in the tall. During the summer in the variations between the populations stu 0~> hi months ol activity, Uta is the Inst tn le.ive died. Are, is \ 01 and \ are the most diver- shelter ill the moinuig and the last to seek it at gent, yet in both of these populations the stan- evening. The availability of Uta lor study, not dard deviation ol their means is overlapping only its abundance in the study aieas but also one another. its wide range ol activity, has permitted me to Equally significant is the lack ol variation use larger series in all characteristics analyzed. between area \-0l and the adjoining areas ol Figures 7, 8, 9, 10, 11 and 12 provide the data \ 02 and V-03. Biological Series, Vol. 7, No. 1, May, 1965 19

I I I I I I I I I I I I I I I I I I I I I l I I I I I I I I I | I I | I ' I I I M

I i i' i iI ' i i i i i i i i i i i i i I I I I I I I ' iI iI I I Ii ' ' Mi il I i I n I| | | |

A-OI 315

A-02 251 A-03

A -03 jtL 261 — A-04

A-04 . -T±L 130 A-05

A-05 280 A " 06

A-06 JZtL

' » ' I l I I ,1m 80 90 100 110

Fig. 7. Number of scale rows at midbody in six populations of Uta stansburiana . The range of variation is indicated by the horizontal line, the mean by the vertical line, the rectangle encloses one standard error on each side of the mean and the shorter horizontal line above the vertical line ( mean ) represents one standard deviation of the mean. The number of scales ( rows ) is indicated both at the bottom and the top of the graph, with the populations (areas A-01 ) to the left of the range (long horizontal line) and size of the population sample on the right margin of the range.

2. The number of scales in the dorsal series (Fig. 8) is noticeably more variable than the number of scale rows (Fig. 7). A skewing to the right of the populations A-04, -05 and -06 in this character is obvious. Again the areas A-01, -02 and -03 are similar. The greatest variation between these populations occurs between A-02 and A-05. In these there is a difference of 6 scales between the means; and although the standard deviations overlap, this is less than 50 percent.

3. The numbers of ventral scales as graphed in Figure 9 show little variation.

4. Figure 10, representing the total number of femoral pores, provides a different pattern of variation. In the first four populations there is a decrease in the mean from A-01 to A-04. The means of A-05 and A-06 are noticeably skewed to the left, yet as graphed the means vary only approximately three pores and the standard dev- iations of all populations overlap.

5. Variation of the supralabials in these pop- ulations is nil. The variation between the means is less than one scale ( Fig. 11 ). I i

20 BlUCHAM Yi'l NC IMsikmii Si itNCE BULLETIN

1 ' "i r "i 1 r ' ' 71 ' r — —— A-OI- _r±i_ 290

A-02- 177

A-03 _L±L 192

A-04 98

A-05 I—I— .255

A-06 -33

21 22 23 24 25 26 27 28 29 30 31 32 33 34 35

Fie. 10. Total number of femoral pores on both femurs in Uta stansburianu . See Fig. 7 for explanation of graph.

1 1 1 1 1 1 1 "I 1 i r

, ttl A-OI , 296 A-OI

A-02 rh 183 A-02 rh _m. A-03 _r±L 98 A-04 271

37 A-05

A-06

Fie. 11. Total number of supralabial scales on both lower lips in Uta statwburiana. See Fig. 7 for explana- tion of graph. 10

Sceloporus undulatus elongatus Stejneger

The Northern Plateau Lizard inhabits the rocky boulder-strewn sidehills and the ledgy out-

croppings within its range. It is found in north- western New Mexico, northeastern Arizona, western Colorado and all ol the Upper Colorado Basin oi Utah extending as far north in the Green River Valley as southern Wyoming. Foi this species the same general procedures

will be followed .is foi ' to. The graphed data foi Sceloporus are found in Figures 13 to 18, Biological Sebies, Vol. 7, No. 1, May, 1965 21

' l I I l l I ' l | l l l ' ' l I I I l I I I I I I 1 I I I I | | I I I | I I | I |

B-OI — r—

BRICHAM Vm MC I Milium S< it s< i Hi i i i iin

1 B-OI I— I 40

B-02 r-t-i 54

B-03 47

B-04 i— 13

B-05 c=t 75

B-06 cb 53

30 35 40 45

FlC. 16. Total number of femoral pores on both femurs in Sceloporus undulatus elongatus. See Fig. 7 for explanation of graph.

the almost complete overlapping of the stan- 5. There is little variation in the supralab- dard deviations. ials. The population means are all between scales 8 and 9, and the standard deviations are also basically in this same range. Noticeably dis- 1 1 1 1 tinct from ah other graphed data for this species

1 I 1 1 1 1 1 1 i I

Biolocical Series, Vol. 7, No. 1, May, 1965

greatest. This emphasizes the regular occurrence i i i I I I I I I I I II i i i i i I I I I I I I I I I of 8 scales in all populations.

6. In the infralabials there is a greater range C-OI 24 of variation, 6 scales, but much less skewness. Populations B-01, B-02 and B-06 are skewed to C-02 ri— 25 the left, whereas the others are slightly skewed to the right. The variation between the popula- tion means is less than 1 scale, and the standard C-03 48 deviations of all populations overlap.

Cnemidophorus tigris septentrionalis Burger C-04 53 The northern whiptail is the largest and fastest of the common lizards found in the gen- C-05 _T±L 48 eral study area. It is more commonly found along the washes but forages along ledges and among the larger shrubs and juniper trees. This C-06 r-M 18 species does not climb on the ledges as is com- monly done by Sceloporus and occasionally by Uta, but remains on the ground. It may find I , shelter under ledges or large rocks, but is seem- 80 90 100 ingly more at home in rodent holes under the Fig. 19. Number of scale rows at midbody in six branches of a shrub or a hole under or along populations of Cnemidophorus tigris septentrionalis. See the side of a rock. Fig. 7 for explanation of graph. As in the case of Sceloporus, the whiptails are not as abundant as Uta and were practically is the extreme skewness to the right of the means eliminated from the study areas as we secured and standard deviations for all populations. In the series of specimens. those populations with the greatest range of Those graphs with the data for CnemidopJio- variation (4 scales, 8 to 11) the skewness is rus are found in Figures 19 to 24. The same

' i i i i i i i i ' ' i l I I l i i i i I I i ' i I ' I i ' I i i ' ' ' I i | | | |

C-OI 23

C-02 J=t 26

C-03 I I I 48

I C-04 — — 52

C-05 I I I 48

C-06 18

U-L i I I i I i I I I I i I I I I I I I I I I I I I 150 160 170 180 190

Fig. 20. Dorsal scales in Cnemidophorus tigris septentrionalis, counting from parietal to base of tail. See Fig. 7 for explanation of graph. i i

S( i J I Hun. ham Vounc Untvebsiti knce Bulletin

characteristics presented in the sequence as I same I I I I I I I | I T I I I I | I | I I I I | I I | in the two preceding species will be given fur this spi c-oi —cb 24 1. There is little variation in the range of the stale rows. The greatest range of 22 rows is found in population C-04 and least of 14 rows c-02 —cb- 26 in < 0] and C-06. An extreme of 4 rows (be- i !-03 and C-04) occurs in the popula- tion means; however, the standard deviations for all populations are not significantly variable. C-03 — lfa_ 50 2. There is little variation in number of stales in the dorsal rows. In this characteristic c-04 ih- 54 the i rtremes oi the range of variation is 41 scales, and although this extreme is not rea< hed

bv all populations, the minimums is 27 scales.

It is noteworthy that the means for these cb popula- c-05 — 49 tions vary less than 3 scales.

3. The number of scales in the ventral rows of these populations are remarkably uniform and J±L 18 C-06 show little variation. An extreme variation in the range of only 12 scales and of only approx-

imately 2 scales in the population means is evi-

i i . i l i i i i i i ' dence of the sameness of these populations in 30 40 this character.

I i.. Ventral tigris 21. scales in Cnemidophorus 4. The range of variation in the total num- septentrUmaUs, counting from the enlarged scales just ber of pores is from 34 to 49, or 15 pores. The posterioi to the gular fold (this is approximate!) .it the two population extremes are between C-03, with . interior square of the front legs) to the granule scales

.it the sent See Fig. 7 for explanation of the graph. 34 to 48, and C-05, with 40 to 49. Population

C-01 i—i— 24

C-02 23

C-03 i—t— 42

C-04 r-1-1 54

C-05 m 49

C-06 i 18

35 40 45 50

in. iz Total number >! Femoral pores on both femurs in Cnemidophorus tigrlt septenMonaUs. See Fig. 7 lor explanation "t graph — L I

25 Biological Series, Vol. 7, No. 1, May, 1965

1 significant, not perhaps in this case to establish I — an important and significant population differ- ence, but to indicate the separateness of popula- C-OI r±3 23 tions in a given small area. Populations C-02 and C-03 occur in adjoining small tributary washes which are separated by a small ridge. C-02 26 The washes join about half a mile below the designated areas. Yet these populations in spite of their close approximations do not apparently -dn. 49 interbreed. This is an added evidence of close c-03 territorialism in this species.

6. The infralabials show no more variation in the population means than occurred in the C-04 • 1— 52 supralabials even though the total range is twice as great (6 scales). Figures 23 and 24 show a very similar pattern of variation but with more C.05 55 48 overlapping of the standard deviations in the infralabials.

C-06 r-H . 18

1 T i 1

C-OI rm 10 II 12 13

number of supralabial scales on both Fig. 23. Total C-02 i—t-i 26 upper lips in Cnemidophorus tigris septentrionalis . See Fig. 7 for explanation of graph. C-03 50 means vary only 2.5 pores, and the standard deviations overlap widely. C-04 5. Perhaps the most significant variations observed in this species occur in the supralabials. C-05 47 Although one might expect the extremes of var- iation to occur between those populations in- habiting the areas with high surface radiation, in C-06 this instance at least those two areas with lowest radiation (C-02 and C-03) have not only the greatest variation in range but also the greatest mean variation. In a character such as this, with a total range Fig. 24. Total number of infralabials scales on both of variation equaling only three scales, a popu- lower lips in Cnemidophorus tigris septentrionalis. See lation means which varies 1 scale is seemingly Fig. 7 for explanation of graph.

MAMMALS

Five species of small mammals were found mens now available at the Brigham Young Uni- in each of the study areas. Several other species versity Museum. Species with at least four adult were taken in limited numbers in some of the specimens from an area have been carefully areas, thus not providing enough specimens for compared as to their pelage and color variations. adequate comparative purposes. Table 4 lists In making a preliminary comparison of the the species taken at each area and the speci- allopatric species, several variations are notice- Biucham Voi mc University s< ^se i Bulletin able The squirrels provide a variation in the according to our present understanding, they color ot the pelage, Those- taken at Temple belong in .1 single subspei ies However, it is very Mi tain show greatei variation within the sei (ili\i(nis that tin- Temple Mountain-Mamie Sto- us and arc- not like the series from eithei Yellow vei serii s n present a single population, whereas

Cat or those taken thirty miles to (he north along those from Yellow < at represent .1 second dis- the s.m Rafael Reel in the Mamie Stover tinct population. The latter r\liil>its .1 distinct

',, is not as we had expected, In lac t pelage pattern in which the four stripes stand other series. Neotoma lepida sanrafaeli and out sharpl) in contrast to the reddish ground PeromySCUS CrinitUS doutti, show similar popula- color. In contrast those taken at Temple Moun- tion variations. Based on the nearness ol the tain and Mamie Stover show less distinctness,

Temple Mountain and the Mamie StOVl 1 with both the stripes and the reddish ground and the complete lack ol barriers, one would color appearing washed out or laded. This is re- expect greater similarities between these popu- flected also in the tail and particularly in the lations than between those occurring across the ventral-lateral region, which in some of the Em- Green River in the Yellow Cat district. How- ery County population lias laded to a yellowish ever, only in the chipmunk and white footed linll rather than retaining the reddish ground deer mouse do we Find a close similarity between color. In the chipmunks a distinct pelage vari- the two San Rafael populations. In the mammals, ation occurs between those populations sampled

as is evident in the reptiles, relatively minor in the San Rafael populations and the Yellow barriers seemingly have served to provide for Cat population. There is seemingly a different population and subspecies variations, or other gene pool than that occurring in the Yellow Cat environmental factors operating in conjunction population. with the barriers are responsible for the pelage Neotoma lepida sanrafaeli variations. The common packrats provide a similar pre- sumably anomalous in which {mmospermophUus leucurus notum phenomenon the pelage color and the basic variations in the The antelope ground squirrel occurring in Mamie Stover population are duplicated by the the three areas in which were studied mammals series from Yellow Cat, whereas the population represents two subspecies, notutu in Emery from Temple Mountain is relatively uniform in and Grand County. The pop- County pennipes in color ,md distinct when compared with the ulations indicate very definite pelage varia- a other populations. The pattern of the Temple tion, particularly between the Yellow Cat and Mountain series has a greater amount of yellow, the Mountain populations. In the Yel- Temple particularly laterally, but blending into the low Cat population and dorsal-lateral line stripes brownish-grey dorsal pelage and giving a more are usually straight or with only a slight curve, yellowish, almost buff, color. In A', lepida. as in whereas in the Temple Mountain series the A. leucurus discussed above, a similar popula- stripes are noticeablv curved, often providing tion variation relation exists; that is, the Yellow for two undulations. The basic pelage color in ( at Mamie Stover series seem to represent or ap- tin Temple Mountain is lighter, with population proximate the same variations within each popu- a more buff-reddish color. Furthermore, there is lation, considerably more variation in the series, ex- tending from dark brownish-gray to a light red- Peromyscus maniculatus sonariensis dish buff. The lack ol variation between indi- The deer mouse from the Temple Mountain viduals of the Yellow ( !at population is most area represents a population with considerable sinking when compared with the Temple pelage variation, ranging from slate to gray to Mountain series. The Mamie Stover area is rep- yellowish-buff, with several gradations in be-

resented by only two specimens, both of which tween these extremes. The range of variation is are very similar to those taken at Yellow Cat. striking, particularly when contrasted with such 'They do not exhibit the basic pelage or pattern uniform color patterns as are found in Scotoma characteristic ol the 'Temple Mountain series in lepida and Eutamias. The remarkable factor in spite ol the that veiv fad they inhabit a simi this species is its absence from the trapping areas lar habitat. at Mamie Stover.

Eutamias quadrivittatus hopiensis Peromyscus maniculatus nebrascensis

The chipmunks ot the area oiler a better The maniculatus population at Yellow Cat example for comparative variation studies since represents the subspecies nebrascensis. This sub- Biological Sehies, Vol. 7, No. 1, May, 1965 27

species provides a number of interesting varia- and Peromyscus truei, but not in sufficient num- tions ranging from a dull slate gray to a rich bers to provide for reliable variation data. buffv brown. Specimens representing the ex- A study in variation based on pelage pattern tremes in pelage color are adults, and both and color is neither satisfactory nor satisfying. were taken within a day of each other in 1962. The lack of small mammals at Mamie Stover Even though we cannot compare direetlv the and the crash of the population at Temple maiiicuhitus because of the different subspecies, Mountain limited the numbers and this greatly the variation within the population is obvious affected the securing of adequate data for the and marked. mammals. Also the apparent similarities between some species in the Yellow Cat and Mamie Sto- Peromyscus crinitus ver populations would require much more samp- The crinitus deer mouse occurring in the ling of intermediate populations to permit any areas trapped consists of two subspecies, doutti conclusions. in the Emery County areas and auripectus at Although several obvious variations have been Yellow Cat. The latter is a dark brownish-gray found between these mammal populations, pelage with some buffv colors, principally on indicating a definite difference in the gene pool of the dorsal and lateral parts the body and of several species, the cause of these variations snout regions, whereas the subspecies doutti is is not at all obvious. One is not, on the basis of much lighter, with yellowish-buff pelage nearly the data available, able to suggest that the var- uniform over the body and head. The series iations occurring in these populations is not due from Temple Mountain is noticeably lighter than to normal individual variation induced by those is the one from Mamie Stover, although it is ob- (actors in nature responsible for induction of vious that these two populations are actually variation in most populations and maintained in not at as great a variance with each other as the limited are the individual members of Peromyscus population by geographical or eco- maniculatus sonariensis from Temple Mountain. logical isolation. Data now available from the study areas, particularly the limited amount from The Kangaroo rats, Dipodomys ordii, occur in the San Rafael populations, indicate that most the Temple Mountain and Yellow Cat areas, but species are variable and that these variations represent two subspecies. Only a few specimens represent, as in the squirrel A. leucurus and the are available for comparison, and these show a packrat N. lepida, a noticeable distinctly lighter pelage pattern for the Temple and pronounced Mountain material corresponding very closely variation. Unfortunately these variables cannot to a few specimens taken at Green Water Spring be subjected to statistical analyses, and there in San Juan County. A few other species are is therefore no satisfactory test for these exter- present in the areas, such as Neotoma cinerea nal characteristics.

STATISTICAL ANALYSES OF THE DATA

In attempting to test the effects of surface In spite of all such attempts and the relative radiation on the external characteristics of small nearness of the study areas, some physical and

vertebrates, I have investigated the general eco- perhaps biological differences, even though

logical factors in the immediate environment of small, still exist. The statistical tests indicate the lizards and mammals and have measured that these differences did exist in the areas tested the approximate soil mantle which covers the (see Table 14). The extent to which these physi- rocks bearing the radiation minerals. Insofar as cal and biological differences created or influ- possible the study areas selected were as simi- enced the variations cannot be separated. The

lar in elevation, general climatic conditions, predominance of the similarities, however, is type and percent of plant cover, animal types considered to be of great importance and to pro- and geological formations as one could obtain vide as much ecological uniformity as can be

and at the same time include the areas of high expected in such a study program. It is there-

surface radiation. An attempt was also made to fore assumed that if the data indicate substan- keep the study areas on approximately the same tial variations in some of the characteristics stu- southeast slope. died, radiation may be, in part at least, respon- Brich m v<» m. i mvBtum Senates Bulletin

I i mi II The analysis of variance (tests humus I Ix-twirii high and low exposure groups and areas within exposure. Biological Series, Vol. 7, No. 1, May, 1965 29

Table 15. An analysis of the variation in the means of the area group (high and low exposure), with a listing of the F values for each characteristic.

Scale Rows Dorsals Ventrals Pores S. Labials I. Labials Source D.F. M.S. F M.S. F M.S. F M.S. F M.S. F. M.S. F

H.E. 3 776.673 1.55 1577.245 9.37 209.887 30.41 62.428 62.18 1.361 1.73 20.133 2.01 L.E. 1 166.577 168.264 6.903 1.004 .787 10.035

H.E. L.E. Id BrICHAM Vh m. UNIVERSm SciENCl Bill. KM IN

<>r i I A in h IT The results <>l il»- approximate test, testing whether not the lack of b variance wa greateeatet In high exposure >» low exposure

Si ale Rows Biological Series, Vol. 7, No. 1, May, 1965 31

Literature Cited

Bellis, Edward D. 1964. A Summer Six-lined Race- logical and Mineralogical Survey and the Utah Population in Water and Power Board. Prepared at the request runner ( Cnemidophorus sexlineatus ) South Carolina, Herpetologica, 20(1):9-16. of Senator Frank E. Moss. U. S. Government Printing Office, Washington, 1964:1-275. Boutwell, M. 1905. Vanadium and Uranium in J. Jorcensen, C. D. and W. Tanner. 1963. The Southeastern Utah, United States Geological Survey W. application of the density probability function to Bulletin, 260:202-210. determine the home ranges of Uta s. stanshuriana 1962. of the movements Brant, Daniel H. Measures and Cnemidophorus t. tigris. Herpetologica, 19(2): and population densities of small rodents. Univ. 105-115. in Zool., 62:105-184. Calif. Publ. Mohr, C. O. and W. A. Stumpf. 1964. Relation of Burt, W. H. 1940. Territorial behavior and popula- tick and chigger infestations to home areas of Cali- tions of some small mammals in southern Michigan. fornia meadow mice. Jour. Med. Ent., 1:73-77. Misc. Publ. Mus. Zool., Univ. Michigan, 45:1-58. Nelson, R. F. 1957. A preliminary investigation of Calhoun, John B. and James N. Casby. 1958. Cal- the role of selenium in the growth of S. pinnata. culations of home range and density of small mam- B.Y.U. Master's Thesis—unpublished. mals. U. S. Public Health Monograph, 55:1-24. Sorensen, Don. 1963. Wonder mineral: Utah's Ura- nium. Utah History Quarterly, 31(3) :280-290. Cannon, Helen. 1952. The effect of uranium-van- adium deposits on the vegetation of the Colorado Stebbins. Robert C. 1954. Amphibians and reptiles Plateau, American Journal of Science, 250:735-770. of western North America. McGraw Hill Book Co., Inc., New York. 537 pp. Daniel and Blair. 1959. Background radiation and Tananka, R. H., Suciyama and S. Teramura. 1958. endemic faunal range in the San Luis valley of A clumped distribution of trombiculid mites viewed Southern Colorado. Great Basin Naturalist, 19(1): from the true home ranges of host roles. Japanese 37-42. Jour. Sanitary Zoology, 9:28-32. Durrant, Stephen D. 1952. Mammals of Utah. Uni- Tanner, Wilmer W. 1963. Natural Radiation Ef- versity of Kansas Publ., Mus. Nat. Hist., 6:1-549. fects on Vertebrate Animals inhabiting the Uran- Fitch, H. S. 1951. A simplified type of funnel trap ium Areas of Southeastern Utah, pp. 325-326. In: for reptiles. Herpetologica, 7:77-80. Radioecology, Proceedings of the First National . 1960. Autecology of the Copperhead. Uni- Symposium on Radioecology held at Colorado State versity of Kansas Publ.. 13(4):85-288. University, Fort Collins, Colorado, September 10- Harrison, Bertrand F., Stanley L. Welsh, Glen 15, 1961. Vincent Schultz and Alfred W. Klement, Moore. 1964. Plants of Arches National Monu- editors. Reinhold Publ. Corp., New York and The ment. Brigham Young University Sci. Bull., Biol American Institute of Biol. Sci., Washington, D. C, Series, 5(l):5-23. 746 pp. Hayward, C, Lynn, D Elden Beck, and Wii.meh W. Tanner. Wilmer W. and Clive D. Jorcensen. 1963. Tanner. 1958. Zoology of the Upper Colorado Reptiles of the Nevada Test Site. Brigham Young River Basin. The biotic communities. Brigham University Sci. Bull. 3(3): 1-31. Young University Sci. Bull. Biol. Series, 1(3): 1-74. Tinkle, Donald W., Don McGregor, and Dana Sum- Hilpert, L. S. and M. D. Dasch. 1964. Mineral and ner. 1962. Home Range Ecology of Uta stans- water resources of Utah, Report of the U. S. Geo- huriana stejnegeri. Ecology, 43(2) :223-29. Temple Mountain. UTA STANSBURIANA Z^ ROADS

//fl„ EDGE OF WASHES

7^" LEDGES

^§&»~ UTA HOME RANGES

,--— DISTANCE BETWEEN TWO CATCHES

5 A dia graln , c of the Temple »«ne „ *<** Mountain area, showing the location of the home ranges of MotL Xamlmrmna., i; 8 ™ The legend provides the basic information. SCELOPORUS UNDULATUS ELONGATUS

" ROADS „„„ EDGE OF WASHES

T^jf LEDGES

pggg» SCELOPORUS HOME RANGES — DISTANCE BETWEEN TWO CATCHES

K A diagramatic sketch of the Temple Mountain area, showing the location of the home Sr™porw„; unduktm ebngatus. The legend provides the basic information.