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UNITED STATES DEPARTMENT OF AGRICULTURE

BULLETIN No. 1091

Also Technical Bulletin No. 1 of the Agricultural Experiment Station University of Arizona

LIFE HISTORY OF THE KANGAROO RAT

Dipodomys spectabilis spectabilis Merriam

BY

CHARLES T. VORHIES, Entomologist
Agricultural Experiment Station, University of Arizona; and

WALTER P. TAYLOR, Assistant Biologist
Bureau of Biological Survey, U. S. Department of Agriculture

CONTENTS

WASHINGTON
GOVERNMENT PRINTING OFFICE
1922

LIFE HISTORY OF THE KANGAROO RAT,

Dipodomys spectabilis spectabilis Merriam.

By Charles T. Vorhies, Entomologist, Agricultural Experiment Station, University of Arizona; and Walter P. Taylor, Assistant Biologist, Bureau of Biological Survey, U. S. Department of Agriculture.

CONTENTS.

Note.—This bulletin, a joint contribution of the Bureau of Biological Survey and the Arizona Agricultural Experiment Station, contains a summary of the results of investigations of the relation of a subspecies of kangaroo rat to the carrying capacity of the open ranges, being one phase of a general study of the life histories of rodent groups as they affect agriculture, forestry, and grazing.

IMPORTANCE OF RODENT GROUPS.

As the serious character of the depredations by harmful rodents is recognized, State, Federal, and private expenditures for their control increase year by year. These depredations include not only the attacks by introduced rats and mice on food materials stored in granaries, warehouses, commercial establishments, docks, and private houses, but also, particularly in the Western States, the ravages of several groups of native ground squirrels and other noxious rodents in grain and certain other field crops. Nor is this all, for it has been found that such rodents as prairie dogs, pocket gophers, marmots, ground squirrels, and rabbits take appreciable and serious toll of the forage on the open grazing range; in fact, that they reduce the carrying capacity of the range to such an extent that expenditures for control measures are amply justified. Current estimates place the loss of goods due to rats and mice in warehouses and stores throughout the United States at no less than $200,000,000 annually, and damage to the carrying capacity of the open range and to cultivated crops generally by native rodents in the Western States at $300,000,000 additional; added together, we have an impressive total from depredations of rodents.

The distribution and life habits of rodents and the general consideration of their relation to agriculture, forestry, and grazing, with special reference to the carrying capacity of stock ranges, is a subject that has received attention for many years from the Biological Survey of the United States Department of Agriculture. As a result of the investigations conducted much has been learned concerning the economic status of most of the more important groups, and the knowledge already gained forms the basis of the extensive rodent-control work already in progress, and in which many States are cooperating with the bureau. If the work is to be prosecuted intelligently and the fullest measure of success achieved, it is essential that the consideration largely of groups as a whole be supplemented by more exhaustive treatment of the life histories of individual species and of their place in the biological complex. The present report is based upon investigations, chiefly in Arizona, of the life history, habits, and economic status of the banner-tailed kangaroo rat, Dipodomys spectabilis spectabilis Merriam (Pl. I).

INVESTIGATIONAL METHODS.

Some 18 years ago (in 1903) a tract of land 49.2 square miles in area on the Coronado National Forest near the Santa Rita Mountains, Pima County, southern Arizona, was closed to grazing by arrangement between the Forest Service and the Agricultural Experiment Station of the University of Arizona. Since that time another small tract of nearly a section has been inclosed (Griffiths, 1910, 7[1]). This total area of approximately 50 square miles is known as the United States Range Reserve, and is being devoted to a study of grazing conditions in this section and to working out the best methods of administering the range (Pl. II, Fig. 1).

For some years an intensive study of the forage and other vegetative conditions of this area has been made, the permanent vegetation quadrat, as proposed by Dr. F. E. Clements (1905, 161-175), being largely utilized. During the autumn of 1917 representatives of the Carnegie Institution and the Arizona Agricultural Experiment Station visited the Reserve and were impressed with the evidence of rodent damage to the grass cover. The most conspicuous appearance of damage was noted about the habitations of the banner-tailed kangaroo rat (Dipodomys spectabilis spectabilis Merriam), although it was observed also that jack rabbits of two species (Lepus californicus eremicus Allen and L. alleni alleni Mearns), which were very abundant in some portions of the reserve, were apparently affecting adversely the forage conditions in particular localities. Accordingly, the Biological Survey, the Agricultural Experiment Station of the University of Arizona, the Carnegie Institution of Washington, and the U. S. Forest Service have undertaken a study of the relation of the more important rodents to the forage crop of the Range Reserve in Arizona.

The present paper is a first step in this larger investigation.[2] In this work the authors have made no attempt to deal with the taxonomic side of the kangaroo rat problem. It is not unlikely that intensive studies will show that the form now known as Dipodomys spectabilis spectabilis is made up of a number of local variants, some of them perhaps worthy of recognition as additional subspecies. But it is felt that the conclusions here reached will be little, if at all, affected by such developments.

Color descriptions are based on Ridgway's Color Standards and Color Nomenclature published in 1912.

IDENTIFICATION.

There are only three groups of mammals in the Southwest having external cheek pouches. These are (a) the pocket gophers (Geomyidæ), which have strong fore feet, relatively weak hind feet, and short tail, as compared with weak fore feet, relatively strong hind feet, and long tail in the other two; (b) the pocket mice (Perognathus), which are considerably smaller than the kangaroo rats and lack the conspicuous white hip stripe possessed by all the latter; and (c) the kangaroo rats (Dipodomys).

Dipodomys spectabilis spectabilis Merriam requires comparison with three other forms of kangaroo rats in the same general region, namely, D. deserti Stephens, of approximately the same size, and D. merriami Mearns and D. ordii Woodhouse, the last two of decidedly smaller size. The range of deserti lies principally to the west of that of spectabilis, and the two do not, so far as known, overlap. On the other hand, merriami and ordii, and subspecies, occur over a large part of the range of spectabilis, living in very close proximity to its burrows; merriami is even suspected of pillaging the stores of spectabilis. The range of merriami, however, is much more extensive than that of spectabilis (Fig. 1), which argues against a definite ecological dependence or relationship. Separation of the four forms mentioned may be easily accomplished by the following key:

Key to Species of Dipodomys in Arizona.

a¹. Size much larger (hind foot and greatest length of skull more than 42 millimeters); tail tipped with white.

b¹. Upper parts dark brownish buffy; tail dark brownish or blackish with more sharply contrasted white tip; interparietal broader, distinctly separating mastoids (range in Arizona mainly southeastern part)Dipodomys spectabilis.

b². Upper parts light ochraceous-buffy; tail pale brownish with less sharply contrasted white tip; interparietal narrower, reduced to mere spicule between mastoids (range in Arizona mainly southwestern part)) Dipodomys deserti.

a². Size much smaller (hind foot and greatest length of skull less than 42 millimeters); tail not tipped with white.

b¹. Hind foot with four toes
Dipodomys merriami.

b². Hind foot with five toes
Dipodomys ordii.

On account of the small size, merriami and ordii do not require detailed color comparison with the other two. The general color of the upperparts of spectabilis is much darker than that of deserti; whereas spectabilis is ochraceous-buff or light ochraceous-buff grizzled with blackish, deserti is near pale ochraceous-buff and lacks the blackish.

The color of the upperparts alone amply suffices to distinguish spectabilis and deserti; but the different coloration of the tail is the most obvious diagnostic feature. The near black of the middle portion of the tail, the conspicuous white side stripes, and the pure white tip make the tail of spectabilis stand in rather vivid contrast to the pale-brown and whitish tail of deserti.

The dens of the two larger species of Dipodomys—spectabilis and deserti—can be distinguished at a glance from those of the two smaller—merriami and ordii—by the fact that the mounds of the former are usually of considerable size and the burrow mouths are of greater diameter. On the Range Reserve merriami erects no mounds, but excavates its burrows in the open or at the base of Prosopis, Lycium, or other brush. The mounds of spectabilis are higher than those of deserti, the entrances are larger, and they are located in harder soil (Pl. III, Fig. 1). The dens of deserti are usually more extensive in surface area than those of spectabilis, and have a greater number of openings (Pl. III, Fig. 2).

DESCRIPTION.

Size large; ears moderate, ear from crown (taken in dry skin) 9 or 10 millimeters; eyes prominent; whiskers long and sensitive; fore feet short and weak; hind feet long and powerful, provided with four well-developed toes; tail very long, usually 30 to 40 per cent longer than the body. Cranium triangular, the occiput forming the base and the point of the nose the apex of the triangle, much flattened, auditory and particularly mastoid bullae conspicuously inflated.

COLOR.

General color above, brownish buffy, varying in some specimens to lighter buffy tints, grizzled with black; oblique hip stripes white; tail with dark-brown or blackish stripes above and below, running into blackish about halfway between base and tip, and with two lateral side stripes of white to a point about halfway back; tail tipped with pure white for about 40 millimeters (Pl. I). Underparts white, hairs white to bases, with some plumbeous and buffy hairs about base of tail; fore legs and fore feet white all around; hind legs like back, brown above, hairs with gray bases, becoming blackish (fuscous-black or chætura-black) about ankles, hairs on under side white to bases; hind feet white above, dark-brown or blackish (near fuscous) below.

Color variations in a series of 12 specimens from the type locality and points widely scattered through the range of spectabilis consist in minor modifications of the degree of coloration, length of white tip of tail, and length of white lateral tail stripes. In general the color pattern and characters are remarkably uniform. Young specimens, while exhibiting the color pattern and general color of adults, are conspicuously less brown, and more grayish.

There appears to be little variation in color with season. In the series at hand, most specimens taken during the fall, winter, and spring are very slightly browner than those of summer, suggesting that the fresh pelage following the fall molt is a little brighter than is the pelage after being worn all winter and into the following summer. But at most the difference is slight.

OIL GLAND.

Upon separating the hairs of the middle region of the back about a third of the distance between the ears and the rump, one uncovers a prominent gland, elliptical in outline, with long axis longitudinal and about 9 millimeters in length. The gland presents a roughened and granular appearance, and fewer hairs grow upon it than elsewhere on the back. The hairs in the vicinity are frequently matted, as if with a secretion. In worn stage of pelage the gland may be visible from above without separating the hairs. Bailey has suggested that this functions as an oil gland for dressing the fur, and our observations bear out this view. Kangaroo rats kept in captivity without earth or sand soon come to have a bedraggled appearance, as if the pelage were moist. When supplied with fine, dusty sand, they soon recover their normal sleek appearance. Apparently the former condition is due to an excess of oil, the latter to the absorption of the excess in a dust bath. The oil is doubtless an important adjunct to the preservation of the skin and hair amid the dusty surroundings in which the animal lives.

MEASUREMENTS AND WEIGHTS.

External measurements include: Total length, from tip of nose to tip of tail without hairs, measured before skinning; tail vertebræ, length of tail from point in angle when tail is bent at right angles to body to tip of tail without hairs; and hind foot, from heel to tip of longest claw.

The following are measurements of a series from the U. S. Range Reserve:

Average measurements of 30 adult specimens of both sexes: Total length, 326.2 millimeters (349-310); tail vertebræ, 188.4 (208-180); hind foot, 49.5 (51-47); the average weight of 29 adult specimens of both sexes was 114.5 grams (131.9-98.0).

Averages for 17 adult females: Total length, 326.4 millimeters (349-310); tail vertebræ, 188.8 (208-179); weight (16 individuals), 113.7 (131.9-98.0); excluding pregnant females, 13 individuals averaged 112.9 grams (131.9-98.0).

Averages for 13 adult males: Total length, 326 millimeters (345-311); tail vertebræ, 187.8 (202-168); weight, 116.8 grams (129-100).

There appears to be no significant difference in the measurements and weights of males and females, with the possible exception of the comparison of adult males and adult nonpregnant females.

OCCURRENCE.

Dipodomys spectabilis spectabilis is found in southeastern Arizona, in northwestern, central, and southern New Mexico, in extreme western Texas, in northern Sonora, and in northern and central Chihuahua (Fig. 1). A subspecies, D. s. cratodon Merriam, has been described from Chicalote, Aguas Calientes, Mexico, the geographic range of which lies in central Mexico in portions of the States of Zacatecas, San Luis Potosi, and Aguas Calientes.

HABITAT.

In the Tucson region spectabilis is typically a resident of the Lower Sonoran Zone. This is perhaps the principal zone inhabited over its entire range, but the animal is often found in the Upper Sonoran also, and in the Gallina Mountains of New Mexico Hollister found it invading the yellow pine Transition where the soil was dry and sandy and the pine woods of open character. The same observer found it common in grassy and weed-grown parks among the large junipers, pinyons, and scattering yellow pines of the Bear Spring Mountains, N. Mex. Bailey calls attention to the fact that the animal apparently does not inhabit the lower half of the Lower Sonoran Zone, as it extends neither into the Rio Grande Valley of Texas nor the Gila Valley of Arizona. In extreme western Texas it is common at the upper edge of the arid Lower Sonoran Zone, and in this region does not enter the Upper Sonoran to any extent.

In July, 1914, Goldman found this kangaroo rat common on the plain at 4,600 feet altitude, near Bonita, Graham County, Ariz., and noted a few as high as 5,000 feet altitude on the warm southwestern slopes of the Graham Mountains, near Fort Grant. Apparently spectabilis reaches its upper altitude limit in the Burro Mountains, N. Mex., where Bailey has found it sparingly on warm slopes up to 5,700 feet, and at the western base of the Sandia Mountains, east of Albuquerque, N. Mex., where dens occur at approximately 6,000 feet.

About Tucson it is undoubtedly more common in the somewhat higher portions of the Lower Sonoran Zone, above the Covillea association, than elsewhere (Pl. IV, Figs. 1 and 2). A few scattered dens are to be seen in the Covillea belt, but as one rises to altitudes of 3,500 to 4,000 feet, and the Covillea is replaced by the cat's-claws (Acacia sp. and Mimosa sp.) and scattered mesquite (Prosopis), with the Opuntia becoming less abundant, kangaroo rat mounds come more and more in evidence. Here is to be found the principal grass growth supporting the grazing industry, and the presence of a more luxuriant grass flora is probably an important factor in the greater abundance of kangaroo rats, both spectabilis and merriami. In this generally preferred environment the desert hackberry (Celtis pallida) is one of the most conspicuous shrubs; clumps of this species are commonly accompanied by kangaroo rat mounds.

In order to ascertain whether the banner-tailed kangaroo rat has any marked preference for building its mounds under Celtis or some other particular plant, all the observable mounds were counted in a strip about 20 rods wide and approximately 4 miles long, an area of approximately 160 acres, particular note being taken of the kind of shrub under which each mound was located. Of 300 mounds in this area, 96 were under Prosopis, 95 under Acacia, 65 under Celtis, 11 under Lycium, 31 in the open, 1 about a "cholla" cactus (Opuntia spinosior), and 1 about a prickly pear (Opuntia sp.). There is apparently no strongly marked preference for any single species of plant. While both desert hackberry and the cat's-claws afford a better protection than mesquite—since cattle more often seek shade under the latter, and in so doing frequently trample the mounds severely—it appears that the general protection of a tree or shrub of some sort is sought by kangaroo rats, rather than the specific protection of the thickest or thorniest species.

The following records indicate particular habitat preferences of spectabilis as noted at different points in its range:

A. Brazier Howell notes that spectabilis occurs in harder soil than does deserti. This observation is confirmed by others, and seems to afford a conspicuous habitat difference between the two, for deserti is typically an animal of the shifting aeolian sands.

Usually, as on the Range Reserve, the rodents are widely distributed over a considerable area. Occasionally, as in the vicinity of Rio Alamosa, N. Mex., as reported by Goldman, they occur only in small colonies.

HABITS.

One traveling over territory thickly occupied by the banner-tailed kangaroo rat is certain to note the numerous and conspicuous mounds so characteristic of the species, particularly if the region is of the savannah type, grassy rather than brushy. These low, rounded mounds occupy an area of several feet in diameter, and rise to varying heights above the general surface of the surrounding soil, the height depending rather more upon the character of the soil and the location of the mound as to exposure or protection than upon the area occupied by the burrow system which lies within and is the reason for the mound.

A den in sandy soil in the open may be of maximum size in area occupied and yet scarcely present the appearance of a mound in any sense, due probably both to the fact that the sandy soil will not heap up to such a height over a honeycomb of tunnels as will a firmer or rocky soil, and also to its greater exposure to the leveling action of rains and the trampling of animals. These mounds are in themselves large enough to attract some attention, but their conspicuousness is enhanced by the fact that they are more or less completely denuded of vegetation and are the centers of cleared areas often as much as 30 feet in diameter (Pl. V, Fig. 1); and further that from 3 to 12 large dark openings loom up in every mound. The larger openings are of such size as to suggest the presence of a much larger animal than actually inhabits the mound. Add to the above the fact that the traveler by day never sees the mound builder, and we have the chief reasons why curiosity is so often aroused by these habitations.

On the Range Reserve the mounds are usually rendered conspicuous by the absence of small vegetation, but Nelson writes that in the vicinity of Gallego, Chihuahua, they can be readily distinguished at a distance because of a growth of weeds and small bushes over their summits, which overtop the grass. In the vicinity of Albuquerque, N. Mex., Bailey reports (and this was recently confirmed by Vorhies) that the mounds about the holes of spectabilis are often hardly noticeable. Hollister writes that in the yellow-pine forests of the Gallina Mountains the burrows are usually under the trunk of some fallen pine, both sides of it in some cases being taken up with holes, there being some eight or ten entrances along each side, the burrows extending into the ground beneath the log. In the vicinity of Blanco, N. Mex., Birdseye says that occasionally spectabilis makes typical dens but more often lives in old prairie-dog holes (Cynomys), or in holes which look more like those of D. ordii.

Runways and Tracks.

Still other features add to the interest in the dwelling places of spectabilis. Radiating in various directions from some of the openings of the mounds well-used runways are to be seen, some of them fading out in the surrounding vegetation, but others extending 30, 40, or even 50 or more yards to neighboring burrows or mounds (Pl. V, Fig. 2; Pl. VI, Fig. 1). These runways and the entrances to the mounds are well worn, showing that the inhabitants are at home and are at some time of day very active. The worn paths become most conspicuous in the autumnal harvest season, when they stand out in strong contrast to surrounding grass. One usually finds not far distant from the main habitation one or more smaller burrows, each with from one to three typical openings, connected by the trail or runway system with the central den, and these we have called "subsidiary burrows" (Pl. VI, Fig. 2). These will be again referred to in discussing the detailed plan of the entire shelter system.

Examination of the runways and of the denuded area about a mound discloses an abundance of almost indecipherable tracks. The dust or sand is ordinarily much too dry and shifting to record clear footprints, and there are no opportunities to see footprints of this species recorded in good impressionable soil. Very characteristic traces of kangaroo rats may be readily observed in the dust about the mounds, however, and these are long, narrow, sometimes curving, furrows made by the long tails as the animals whisk about their work or play.