Title: Comparative Ecology of Pinyon Mice and Deer Mice in Mesa Verde National Park, Colorado
Author: Charles L. Douglas
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Most recently updated: January 8, 2021
Language: English
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University of Kansas Publications
Museum of Natural History
Volume 18, No. 5, pp. 421-504
August 20, 1969
Comparative Ecology of Pinyon Mice
and Deer Mice in
Mesa Verde National Park, Colorado
BY
CHARLES L. DOUGLAS
University of Kansas
Lawrence
1969
University of Kansas Publications, Museum of Natural History
Editors of this number: Frank B. Cross, Philip S. Humphrey,
J. Knox Jones, Jr.
Volume 18, No. 5, pp. 421-504
Published August 20, 1969
University of Kansas
Lawrence, Kansas
PRINTED BY
ROBERT R. (BOB) SANDERS, STATE PRINTER
TOPEKA, KANSAS
1969
32-6879
Centuries ago in southwestern Colorado the prehistoric Pueblo inhabitants of the Mesa Verde region expressed their interest in mammals by painting silhouettes of them on pottery and on the walls of kivas. Pottery occasionally was made in the stylized form of animals such as the mountain sheep. The silhouettes of sheep and deer persist as pictographs or petroglyphs on walls of kivas and on rocks near prehistoric dwellings. Mammalian bones from archeological sites reveal that the fauna of Mesa Verde was much the same in A. D. 1200, when the Pueblo Indians were building their magnificent cliff dwellings, as it is today. One of the native mammals is the ubiquitous deer mouse, Peromyscus maniculatus. The geographic range of this species includes most of the United States, and large parts of Mexico and Canada.
Another species of the same genus, the pinyon mouse, P. truei, also lives on the Mesa Verde. The pinyon mouse lives mostly in southwestern North America, occurring from central Oregon and southern Wyoming to northern Oaxaca. This species generally is associated with pinyon pine trees, or with juniper trees, and where the pinyon-juniper woodland is associated with rocky ground (Hoffmeister, 1951:vii).
P. maniculatus rufinus of Mesa Verde was considered to be a mountain subspecies by Osgood (1909:73). The center of dispersion for P. truei was in the southwestern United States, and particularly in the Colorado Plateau area (Hoffmeister, 1951:vii). The subspecies P. truei truei occurs mainly in the Upper Sonoran life-zone, and according to Hoffmeister (1951:30) rarely enters the Lower Sonoran or Transition life-zones. P. maniculatus and P. truei are the most abundant of the small mammals in Mesa Verde National Park, which comprises about one-third of the Mesa Verde land mass.
Under the auspices of the Wetherill Mesa Archeological Project, the flora of the park recently was studied by Erdman (1962), and by Welsh and Erdman (1964). These studies have revealed stands of several distinct types of vegetation in the park and where each type occurs. This information greatly facilitated my study of the mammals inhabiting each type of association. The flora and fauna within the park are protected, in keeping with the policies of the National Park Service, and mammals, therefore, could be studied in a relatively undisturbed setting.
Thus, the abundance of these two species of Peromyscus, the botanical studies that preceded and accompanied my study, the relatively undisturbed nature of the park, and the availability of a large area in which extended studies could be carried on, all contributed to the desirability of Mesa Verde as a study area.
My primary purpose in undertaking a study of the two species of Peromyscus was to analyze a number of ecological factors influencing each species—their habitat preferences, how the mice lived within their habitats, what they ate, where they nested, what preyed on them, and how one species influenced the distribution of the other. In general, my interest was in how the lives of the two species impinge upon each other in Mesa Verde.
The Mesa Verde consists of about 200 square miles of plateau country in southwestern Colorado, just northeast of Four Corners, where Colorado, New Mexico, Arizona and Utah meet. In 1906, more than 51,000 acres of the Mesa Verde were set aside, as Mesa Verde National Park, in order to protect the cliff dwellings for which the area is famous.
The Mesa Verde land mass is composed of cross-bedded sandstone strata laid down by Upper Cretaceous seas. These strata are known locally as the Mesaverde group, and are composed, from top to bottom, of Cliff House sandstone, the Menefee formation, the Point Lookout sandstone, the well known Mancos shale, and the Dakota sandstone, the lowest member of the Cretaceous strata. The Menefee formation is 340 to 800 feet thick, and contains carbonaceous shale and beds of coal.
There are surface deposits of Pleistocene and Recent age, with gravel and boulders of alluvial origin; colluvium composed of heterogeneous rock detritus such as talus and landslide material; and alluvium composed of soil, sand, and gravel. A layer of loess overlays the bedrock of the flat mesa tops in the Four Corners area. The earliest preserved loess is probably pre-Wisconsin, possibly Sangamon in age (Arrhenius and Bonatti, 1965:99).
The North Rim of Mesa Verde rises majestically, 1,500 feet above the surrounding Montezuma Valley. Elevations in the park range from 8,500 feet at Park Point to about 6,500 feet at the southern ends of the mesas. The Mesa Verde land mass is the remnant of a plateau that erosion has dissected into a series of long, narrow mesas, joined at their northern ends, but otherwise separated by deep canyons. The bottoms of these canyons are from 600 to 900 feet below the tops of the mesas.
The entire Mesa Verde land mass tilts southward; Park Headquarters, in the middle of Chapin Mesa (Fig. 1), is at about the same elevation as is the entrance of the park, 20 miles by road to the north.
Mesa Verde is characterized by pinyon-juniper woodlands that extend throughout much of the West and Southwest. Although the pinyon-juniper woodland dominates the mesa tops, stands of Douglas fir occur in some sheltered canyons and on north-facing slopes. Thickets of Gambel oak and Utah serviceberry cover many hillsides and form a zone of brush at higher elevations in the park. Aspens grow in small groups at the base of the Point Lookout sandstone and at a few other sheltered places where the supply of moisture suffices. Individual ponderosa pine are scattered through the park, and stands of this species occur on some slopes and in the bottoms of some sheltered canyons.
Tall sagebrush grows in deep soils of canyon bottoms, and in some burned areas, and was found to be a good indicator of prehistoric occupation sites.
The climate of Mesa Verde is semi-arid, and most months are dry and pleasant. Annual precipitation has averaged about 18.5 inches for the last 40 years. July and August are the months having the most rainfall. Snow falls intermittently in winter, and may persist all winter on north-facing slopes and in valleys. In most years, snow is melting and the kinds of animals that hibernate are emerging by the first of April.
Because of the great differences in elevation between the northern and southern ends of the mesas, differences in climate are appreciable at these locations. Winter always is the more severe on the northern end of the park, owing to persistent winds, lower temperatures, and more snow. The northern end of the park is closer to the nearby La Platta Mountains where ephemeral storms of summer originate. They reach the higher elevations of the park first, but such storms dissipate rapidly and are highly localized. The northern end of the park therefore receives much more precipitation in summer and winter than does the southern end.
The difference in precipitation and the extremes in weather between the northern and southern ends of the mesas affect the distribution of plants and animals. Species of mammals, plants, and reptiles are most numerous on the middle parts of the mesas, as also are cliff-dwellings, surface sites, and farming terraces of the prehistoric Indians.
Anderson (1961) reported on the mammals of Mesa Verde National Park, and Douglas (1966) reported on the amphibians and reptiles. In each of these reports, earlier collections are listed and earlier reports are summarized.
I lived in Mesa Verde National Park for 28 months in the period July 1961 to September 1964, while working as Biologist for the Wetherill Mesa Archeological Project, and the study here reported on is one of the faunal studies that I undertook.
This study could not have been completed without the assistance and encouragement of numerous persons. I am grateful to Dr. Olwen Williams, of the University of Colorado, for suggesting this study and helping me plan the early phases of it.
Mr. Chester A. Thomas, formerly Superintendent, and Mrs. Jean Pinkley, formerly Chief of Interpretation at Mesa Verde National Park, permitted me to use the park's facilities for research, issued collecting permits, and in 1965 appointed me as a research collaborator in order that I might complete my studies.
Dr. H. Douglas Osborne, California State College, Long Beach, formerly Supervisory Archeologist of the Wetherill Mesa Project, took an active interest in my research and provided supplies, transportation and laboratory and field assistance under the auspices of the Wetherill Project. His assistance and encouragement are gratefully acknowledged.
Mrs. Marilyn A. Colyer of Mancos, Colorado, ably assisted in analyzing vegetation in the trapping grid; Mr. Robert R. Patterson, the University of Kansas, assisted me in the field in October of 1963 and in August of 1965. Mr. James A. Erdman, United States Geological Survey, Denver, formerly Botanist for the Wetherill Mesa Project, and Dr. Stanley L. Welsh, Brigham Young University, identified plants for me in the field, and checked my identifications of herbarium specimens. I owe my knowledge of the flora in the park to my association with these two capable botanists.
I am grateful to the following persons for identification of invertebrates: D. Eldon Beck, fleas and ticks; Paul Winston, mites; V. Eugene Nelson, mites; William Wrenn, mites; Wayne W. Moss, mites; William B. Nutting, mites (Desmodex); Marilyn A. Colyer, insects; John E. Ubelaker, endoparasites; Veryl F. Keen, botflies. George A. King, Architect, of Durango, Colorado, prepared the original map for Figure 1.
Mr. Harold Shepherd of Mancos, Colorado, Senior Game Biologist, Colorado Department of Fish, Game and Parks, obtained permission for me to use the department's trapping grid near Far View Ruins, and provided me with preserved specimens of mice.
Mr. Fred E. Mang Jr., Photographer, National Park Service, processed large numbers of photomicrographs of plant epidermis. Dr. Kenneth B. Armitage, The University of Kansas, offered valuable suggestions for the study of water consumption in the two species of Peromyscus, and permitted me to use facilities of the Zoological Research Laboratories at The University of Kansas. Dr. Richard F. Johnston, The University of Kansas, permitted me to house mice in his controlled-temperature room at the Zoological Research Laboratories. I am grateful to all of the above mentioned persons for their aid.
I acknowledge with gratitude the guidance, encouragement, and critical assistance of Professor E. Raymond Hall throughout the course of the study and preparation of the manuscript. I also extend my sincere thanks to Professors Henry S. Fitch, Robert W. Baxter, and William A. Clemens for their helpful suggestions and assistance.
To my wife, Virginia, I am grateful for encouragement and assistance with many time-consuming tasks connected with field work and preparation of the manuscript.
Travel funds provided by the Kansas Academy of Science permitted me to work in the park in August, 1965. The Wetherill Mesa Project was an interdisciplinary program of the National Park Service to which the National Geographic Society contributed generously. I am indebted to the Society for a major share of the support that resulted in this report. This is contribution No. 44 of the Wetherill Mesa Project.
Trapping was begun in September of 1961 in order to analyze the composition of rodent populations within the park. I used the method of trapping employed by Calhoun (1948) in making the Census of North American Small Mammals (N. A. C. S. M.). It consisted of two lines of traps, each 1,000 feet long having 20 trapping stations that were 50 feet apart. The lines were either parallel at a distance of 400 feet from each other, or were joined to form a line 2,000 feet long. Three snap traps were placed within a five-foot radius of each station, and were set for three consecutive nights. More than a dozen areas were selected for extensive trapping (Fig. 1). Some of these were retrapped in consecutive years in order to measure changes in populations.
One circular trapline of 159.5 feet radius was established in November 1961, and was tended for 30 consecutive days to observe the effect of removing the more dominant species (Calhoun, 1959).
Other mouse traps and rat traps were set in suitable places on talus slopes, rocky cliffs, and in cliff dwellings. Most of these traps were operated for three consecutive nights.
In order to test hypotheses concerning habitat preferences of each of the species of Peromyscus, several previously untrapped areas that appeared to be ideal habitat for one species, but not for the other, were selected for sampling. In the summers of 1963 and 1964 snap traps were set along an arbitrary line through each of these areas. Traps were placed in pairs; each pair was 20 feet from the adjacent pairs.
A mixture of equal parts of peanut butter, bacon grease, raisins, roman meal and rolled oats was used as bait. Rolled oats or coarsely ground scratch feed was used in areas where insects removed the mixture from the traps.
Rodents trapped by me were variously prepared as study skins with skulls, as flat skins with skulls, as skeletons, as skulls only, or as alcoholics. Representative specimens were deposited in The University of Kansas Museum of Natural History. In the course of my study, traps were set in the following areas:
In July 1959 a fire destroyed more than 2,000 acres of pinyon-juniper forest (Pinus edulis and Juniperus osteosperma) in the eastern part of the park. The burned area extends from Morfield Canyon to Waters Canyon, encompassing several canyons, Whites Mesa, and a ridge between Morfield Canyon and Waters Canyon that is known locally as Morfield Ridge (Fig. 1). Beginning on September 4, 1961, three pairs of traplines were run on this ridge at elevations of 7,300 to 7,600 feet.
Vegetation in the trapping area consisted of dense growths of grasses and herbaceous plants, which had covered the ground with seeds. In this and in the following accounts, the generic and specific names of plants are those used by Welsh and Erdman (1964). The following plants were identified from the trapping area on Morfield Ridge:
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Lithospermum ruderale Chenopodium pratericola Achillea millefolium Artemisia tridentata Aster bigelovii Chrysothamnus depressus Chrysothamnus nauseosus Helianthus annuus Helianthella sp. Lactuca sp. Lepidium montanum Quercus gambelii Agropyron smithii Bromus inermis Bromus japonicus Oryzopsis hymenoides Calochortus nuttallii Linum perenne Sphaeralcea coccinea Polygonum sawatchense Solidago petradoria Wyethia arizonica Nicotiana attenuata Fendlera rupicola Penstemon linarioides |
Only Peromyscus maniculatus, Perognathus apache and Reithrodontomys megalotis were taken in this area (Table 1). Many birds inhabit this area, including hawks, ravens, towhees, jays, juncos, woodpeckers, doves, sparrows and titmice. Rabbits, badgers and mule deer also live in the area. Only two reptiles, a horned lizard and a collared lizard, were seen.
Two parallel trap lines were established on October 4, 1961, in the area immediately south of Far View Ruins (Fig. 1). In altitude, latitude and geographical configuration the area is similar to that trapped in the Morfield burn, but the Chapin Mesa site had not been burned.
Canopy vegetation is pinyon-juniper forest. A dense understory was made up of Amelanchier utahensis (serviceberry), Cercocarpos montanus (mountain mahogany), Purshia tridentata (bitterbrush), and Quercus gambelii (Gambel oak). The ground cover consisted of small clumps of Poa fendleriana (muttongrass), and Koeleria cristata (Junegrass), intermingled with growths of one or more of the following:
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Artemisia nova Solidago petradoria Sitanion hystrix Astragalus scopulorum Lupinus caudatus Eriogonum alatum Penstemon linarioides Eriogonum racemosum Eriogonum umbellatum Polygonum sawatchense Amelanchier utahensis Purshia tridentata Comandra umbellata |
Seeds of Cercocarpos montanus covered the ground under the bushes in much of the trapping area, and large numbers of juniper berries were on the ground beneath the trees. Individuals of P. truei and P. maniculatus were caught in this area (Table 1).
Several deer, rabbits, one coyote, and numerous birds were seen in the area. No reptiles were noticed, but they were not searched for. A mountain lion was seen in this general area two weeks after trapping was completed.
Three pairs of traplines were run west of Far View Ruins in an area comparable in vegetation, altitude, general topography, and configuration to the area previously described. The elevations concerned are typical of the middle parts of mesas throughout the park. This area differs from the trapping area south of Far View Ruins and the one on Morfield Ridge in being wider and on the western side of the mesa.
The woody understory was sparse in most places, and where present was composed of Cercocarpos montanus, Purshia tridentata, Fendlera rupicola (fendlerbush), Amelanchier utahensis, Quercus gambelii, and Artemisia tridentata (sagebrush). The herbaceous ground cover was dominated by Solidago petradoria (rock goldenrod), and grasses—including Poa fendleriana, Oryzopsis hymenoides, and Sitanion hystrix. Other herbaceous species were as follows:
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Echinocercus coccineus Achillea millefolium Aster bigelovii Wyethia arizonica Lepidium montanum Lupinus caudatus Yucca baccata Linum perenne Eriogonum racemosum Eriogonum umbellatum Polygonum sawatchense Delphinium nelsonii Penstemon linarioides |
Fresh diggings of pocket gophers were observed along the trap lines. Badger tunnels were noted in numerous surface mounds that are remnants of prehistoric Indian dwellings, but no badgers were seen. Numerous deer and several rabbits were present. Juncos, two species of jays, and woodpeckers were seen daily. No reptiles were observed.
Both Peromyscus maniculatus and P. truei were caught in this area (Table 1).
A circular trapline, 1,000 feet in circumference, was established on November 16, 1961, in a stand of big sagebrush, and was operated for 30 consecutive nights.
The vegetation of the trapping area was predominantly Artemisia tridentata (big sagebrush), interspersed with a few scattered seedlings of pinyon and juniper. This stand was burned in 1858 (tree-ring date by David Smith) and some charred juniper snags still stood. The deep sandy soil also supported a variety of grasses and a few other small plants. The following species were common in this area:
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Bromus inermis Oryzopsis hymenoides Poa fendleriana Sitanion hystrix Solidago petradoria Orthocarpus purpureo-albus |
The 15 to 20 acres of sagebrush were surrounded by pinyon-juniper forest. The trapping station closest to the forest was approximately 100 feet from the edge of the woodland. More P. truei than P. maniculatus were caught here (Table 1).
The trapping area lies north of Cliff Palace, eastward of the loop road, at elevations of 6,875 to 6,925 feet. Two pairs of traplines were run from January 9, 1962, to January 12, 1962, and from February 13 to 15, 1962.
Vegetation was pinyon-juniper woodland with an understory of mixed shrubs. One to four inches of old snow covered the ground during most of the trapping period, but the ground beneath trees and shrubs was generally clear, providing suitable location for traps.
Numerous juncos and jays were seen in this area; deer and rabbits also were present.
Individuals of P. truei and of P. maniculatus were taken (Table 1).
Navajo Hill is the highest point (8,140 feet) on Chapin Mesa. The top of the hill is rounded and the sides slope gently southward and westward until they level out into mesa-top terrain at elevations of 7,950 to 8,000 feet. The northern and eastern slopes of the hill drop abruptly into the respective canyon slopes of the East Fork of Navajo Canyon and the West Fork of Little Soda Canyon. The gradually tapering southwestern slope of the hill extends southward for one mile and is bisected by the main highway, which runs the length of the mesa top.
Heavy growths of grasses cover the ground; Amelanchier utahensis, Cercocarpos montanus, and Fendlera rupicola comprise the only tall vegetation. Trees are lacking on this part of the mesa, except on the canyon slopes, where Quercus gambelii forms an almost impenetrable barrier.
Four traplines were run from May 4-7, 1962, and from May 9-12, 1962. P. maniculatus was taken but P. truei was not present here in 1962, or in 1964 or 1965 when additional trapping was performed as a check on populations (Table 1).
Other species trapped include the montane vole, long-tailed vole, and Colorado chipmunk. Mule deer and coyotes were abundant in the area. Striped whipsnakes, rattlesnakes and gopher snakes are known to occur in this vicinity (Douglas, 1966).
In 1934 a widespread fire deforested large areas of pinyon-juniper woodland on the northern end of Wetherill Mesa. The current vegetation consists of shrubs with a dense ground cover of grasses. Many dead trees still remain on the ground, providing additional cover for wildlife.
The trapping area was a wide, grassy meadow, three and a half miles south of the northern end of the mesa. A pronounced drainage runs through this area and empties into Rock Canyon. Four traplines were run parallel to each other. The first lines were established on May 23, 1962, and the second pair on June 3, 1962.
Another pair of lines was run in a grassy area two miles south of the northern escarpment of Wetherill Mesa. This area was one and a half miles north of the above-mentioned area. These lines ran along the eastern side of a drainage leading into Long Canyon. The vegetation was essentially the same in both areas, and they will be considered together.
The vegetation was composed predominantly of grasses. Quercus gambelii and Amelanchier utahensis were the codominant shrubs. Artemisia tridentata and Chrysothamnus depressus (dwarf rabbitbrush), were common. Plants in the two areas included the following:
Individuals of P. maniculatus and of Reithrodontomys megalotis were caught (Table 1).
Table 1—Major Trapping Localities in Mesa Verde National Park, Colorado. Vegetational Key as Follows: 1) Pinyon-Juniper-Muttongrass 2) Pinyon-Juniper-Mixed Shrubs 3) Juniper-Pinyon-Bitterbrush 4) Juniper-Pinyon-Mountain Mahogany 5) Grassland with Mixed Shrubs 6) Big Sagebrush 7) Pinyon-Juniper-Big Sagebrush 8) Grassland.
| Locality | Date | No. trap nights | P. truei | P. man. | Type of vegetation |
| Morfield Ridge | Sept. 1961 | 1080 | 0 | 83 | 5 |
| Oct. 1963 | 360 | 0 | 13 | 5 | |
| S. of Far View | Oct. 1961 | 360 | 10 | 13 | 2 |
| W. of Far View | Oct. 1961 | 1080 | 22 | 17 | 2 |
| South Chapin Mesa | Nov.-Dec. 1961 | 3600 | 16 | 9 | 6 |
| East Loop Road | Jan. 1962 | 720 | 6 | 2 | 2 |
| Navajo Hill | May 1962 | 720 | 0 | 18 | 5 |
| Aug. 1964 | 20 | 0 | 2 | 5 | |
| Aug. 1965 | 50 | 0 | 8 | 5 | |
| N. Wetherill Mesa | May-June 1962 | 1080 | 0 | 57 | 5 |
| Bobcat Canyon Drainage | June 1962 | 360 | 0 | 0 | 6 |
| N. of Long House | June 1962 | 1080 | 3 | 4 | 1 |
| Mug House—Rock Springs | Aug. 1962 | 720 | 8 | 14 | 4 |
| Aug. 1963 | 720 | 9 | 7 | 4 | |
| S. Wetherill Mesa | Aug. 1962 | 720 | 0 | 5 | 3 |
| 1 mi. SE Park Entr. | June 1963 | 50 | 0 | 16 | 7 |
| 1/4 mi. SE Park Entr. | July 1963 | 100 | 0 | 7 | 8 |
| M-2 Weather Sta. | May 1964 | 25 | 2 | 0 | 1 |
| 8 mi. S North Rim Moccasin Mesa | Aug. 1964 | 100 | 0 | 3 | 8 |
| 10 mi. S North Rim Moccasin Mesa | Aug. 1964 | 25 | 2 | 0 | 2 |
Bobcat Canyon, a large secondary canyon on the eastern side of Wetherill Mesa, is a major drainage for much of the mesa at its widest part. The mesa top drains southeast into a pour-off at the head of Bobcat Canyon. A stand of big sagebrush, Artemisia tridentata, grows in the sandy soil of the drainage, and extends northwest for several hundred yards from the pour-off. The sagebrush invades the pinyon-juniper forest at the periphery of the area.
Two traplines were set in the drainage, with trapping stations at intervals of 25 feet. The lines traversed elevations of 7,000 to 7,100 feet, and were run from June 26 to 29, 1962.
Grasses are the most abundant plants in the ground cover. Artemisia dracunculus is common in the drainage, and A. nova grows around the periphery of the drainage. Other species occurring in this stand include:
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Aster bigelovii Tetradymia canescens Tragopogon pratensis Bromus tectorum Poa fendleriana Sitanion hystrix Stipa comata Lupinus argenteus Calochortus gunnisonii Sphaeralcea coccinea Phlox hoodii Eriogonum umbellatum Peraphyllum ramosissimum Purshia tridentata Penstemon linarioides |
No mice were caught in three nights of trapping (360 trap nights), and only one mammal, a Spermophilus variegatus, was seen.