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.

Signals.

If a scratching or tapping sound be made at the mouth of a burrow, even in the daytime, one is likely to hear a muffled tapping in response, and this may at times be heard while one is engaged in excavating a mound. It has a chirring or fluttering quality, described by Fisher as resembling the noise of a quail flying. Bailey (1905, 148) is of the opinion that it is used as a signal of alarm, call note, or challenge, a view which the present authors believe to be correct. During the winter of 1920-21, however, both Bailey and Vorhies discovered that this sound, or a very similar one, is made by the rapid action of the forefeet in digging. On one occasion in the laboratory the sound was given by one of a pair and was responded to at once by the other, the two being in separate but contiguous cages. This observation, however, could not be repeated. (Vorhies MS.)

One evening, while working in the vicinity of the Burro Mountains, N. Mex., Goldman heard a kangaroo rat near camp making this thumping noise. Taking a lantern, he approached the den, very cautiously, until within 10 feet. The kangaroo rat was just outside the entrance of one of its burrows, and though moving about more or less restlessly at first showed little fear, and kept up the thumping or drumming at intervals. When making the noise the animal was standing with the forefeet on the ground and the tail lying extended. The noise seemed to be made with the hind feet only, and the vibration of the feet could be seen. The tapping was kept up for a second or two at a time, the sounds coming close together and being repeated rhythmically after a very short interval, suggesting the distant galloping of a horse. After continuing in this way for a short time, the animal turned quickly about, with its head in the opposite direction, and began tapping. It appeared to pay little attention to the light, but finally gave a sudden bound and entered one of its holes about 4 feet from the one in front of which it had been standing.

Vorhies has repeatedly noted when watching for the appearance of a kangaroo rat at night that this sound invariably precedes the rodent's first emergence into the open, and often its appearance after an alarm, though when the storage season has begun and the kangaroo rat is carrying loads of grass heads or other material into its den, it regularly comes out without preliminary signaling. Vorhies has also observed it making the sound while on top of the mound, and certainly not digging, but was unable to see how it was made.

Voice.

No data concerning any call notes or sounds other than those described above are at hand, with the following exception: Price (in Allen, 1895, 213), who studied the habits of the animal in the moonlight, at Willcox, Ariz., says that a low chuckle was uttered at intervals; and Vorhies has had one captive female that would repeatedly utter a similar chuckle in a peevish manner when disturbed by day, and one captive male which, when teased into a state of anger and excitement, would squeal much like a cornered house rat. Vorhies has spent many moonlight hours observing kangaroo rats, but without ever hearing a vocal sound uttered by free individuals.

DAILY AND SEASONAL ACTIVITY.

The kangaroo rat is strictly nocturnal. An observer watching patiently by a den in the evening for the animal's first appearance is not rewarded until darkness has fallen completely, and unless the moon is shining the animal can hardly be seen. Were it not for the white tail-brush of spectabilis and its white belly when upright on the hind legs and tail, one could not as a rule see the animal at all when it makes its first evening appearance. With the first streak of dawn activity usually ceases completely and much more abruptly than it began with the coming of darkness, but on a recent occasion Vorhies observed that a kangaroo rat which did not appear until near morning remained above ground until quite light, but not fully daylight. On removal of the plug from the mouth of a kangaroo rat burrow, one may sometimes see a fresh mass of earth and refuse shoved into the opening from within. As often as not, however, even this unwelcome attention does not elicit any response by day, the great majority of the burrow openings of this species, as observed by the authors, remaining permanently open.The ordinary activities of the kangaroo rat in southern Arizona can scarcely be said to show any true seasonal variation. The animals are active all the year in this region, there being neither hibernation nor estivation, both perhaps being rendered unnecessary by the storage habit, to be discussed in full later (pp. 15-16), and by the mildness of the winter climate. On any particular night that the weather is rainy, or the ground too wet and cold, activity is confined to the interior of the burrow system, and for this reason one has no opportunity to see a perfect imprint of the foot in freshly wet soil or in snow. On two or three of the comparatively rare occasions on which there was a light fall of snow on the Range Reserve a search was made for tracks in the snow. At these times, however, as on rainy nights, the only signs of activity were the pushing or throwing out of fresh earth and food refuse from within the burrow. This is so common a sight as to be complete evidence that the animals are active within their dens during stormy weather but do not venture outside. Trapping has again and again proved to be useless on rainy nights, unless the rain is scant and a part of the night favorable, in which case occasional individuals are taken. These statements apply to the Range Reserve particularly; the facts may be quite different where the animals experience more winter, as at Albuquerque, N. Mex., although in November, 1921, Vorhies noted no indications of lessened activity in that region.

PUGNACITY AND SOCIABILITY.

So far as their reactions toward man are concerned, kangaroo rats are gentle and make confiding and interesting pets; this is especially the case with merriami. This characteristic is the more surprising in view of the fact that they will fight each other so readily and so viciously, and yet probably it is explained in part by their method of fighting. They do not appear to use their teeth toward each other, but fight by leaping in the air and striking with the powerful hind feet, reminding one most forcibly of a pair of game cocks, facing each other and guarding in the same manner. Sometimes they carry on a sparring match with their fore feet. Biting, if done at all, is only a secondary means of combat. When taken in hand, even for the first time, they will use their teeth only in the event that they are wounded. The jaws are not powerful, and though the animals may lay hold of a bare finger, with the apparent intention of biting, usually they do not succeed in drawing blood. As Bailey says (1905, 148), they are gentle and timid, and, like rabbits, depend upon flight and their burrows for protection.

The well-traveled trails elsewhere described (p. 10) indicate a degree of sociability difficult to explain in connection with their pugnacity toward each other. While three or four individuals may sometimes be trapped at a single mound, more than two are seldom so caught, and most often only one in one night. Trapping on successive nights at one mound often yields the larger number, yet in some cases the number is explained by the fact that two or three nearly mature young are taken, and the capture of several individuals at a single mound can not be taken to indicate that all are from the one den. Our investigations tend strongly to the conclusion that only one adult occupies a mound, except during the period when the young are in the parental (or maternal) den. In the gassing and excavating of 25 or more mounds we have never found more than one animal in a den, except in one instance, and then the two present were obviously young animals.

SENSE DEVELOPMENTS.

Without making special investigations through a study of behavior or other special methods, one can speak in only general terms regarding what appear to be the special sense developments of kangaroo rats. The eyes are large, as is very often the case in nocturnal animals, and when brought out into the bright light of day the rats perhaps do not see well. Yet, if an animal leaves a den which is in process of excavation, and follows one runway, even in bright sunlight, it makes excellent speed to the next opening, often a distance of several yards. Whether this is accomplished chiefly by the aid of sight or in large measure by a maze-following ability, such as experiments have shown some rodents to have, can not be stated without precise experimentation. Marked ability to follow a maze would not be at all surprising in view of the labyrinthine character of the underground passages which make up the normal habitation.

When watching beside a mound by moonlight one is impressed with the fact that the rats possess either a very keen sense of hearing or of sight, probably both. The very slightest movement or noise on the part of the observer results, with a timid individual, in an instantaneous leap for safety, a disappearance into the burrow so sudden as to be almost startling. All attempts to obtain flashlight photographs at the mounds were failures, the animal either having gotten completely out of the field before the light flashed following the pull of the trigger, or leaving merely an indistinguishable blur on the plate as it went, and this in spite of carefully hiding the trigger chain behind a screen. A slight noise accompanying the trigger action gave the alarm in one case, and in another the length of time of the flash was sufficient for the get-away. The marvelous quickness of the animal clearly indicates a remarkably short reaction time. Occasionally a bold individual is found, as in the case of one which came out repeatedly, even after being flashed twice in the same night.

Certain peculiar physical characteristics suggest a relationship to sense reactions. On these, however, the authors are not prepared to do more than offer suggestions for future work. The extremely large mastoids found in kangaroo rats suggest a connection in some way with special developments of the sense of hearing or of balance. It may be noted that an intermediate condition between the kangaroo rats and the majority of rodents in respect to this character is to be found in the pocket mice (Perognathus), which belong to the same family. Herein lies a field for some interesting experimentation and discovery.

The small, pointed nose might suggest a not overkeen sense of smell, and there appears no reason to believe that this sense is particularly well developed. However, the turbinals are very complex. The vibrissÆ are long and sensitive, and may indicate a special development of the sense of touch as an adaptation to nocturnal habits and to life in an underground labyrinth. The long, well-haired tail doubtless serves as an important tactile organ as well as a balance.

MOVEMENTS AND ATTITUDES.

Movements and attitudes are characteristic. As a kangaroo rat emerges from the burrow a reason for the relatively large size of the opening is seen in the fact that, kangaroolike, the animal maintains a partially upright position. Its ordinary mode of progression is hopping along on the large hind legs, or, when in the open and going at speed, leaping. When moving slowly about over the mound, as if searching for food, it uses the fore legs in a kind of creeping movement. It appears to be creeping when pocketing grain strewn about, but close observation shows that the fore feet are then used for sweeping material into the pockets, reminding one somewhat of a vacuum cleaner. When it assumes a partially upright position the fore limbs are usually drawn up so closely that they can be seen only by looking upward from a somewhat lower level than that occupied by the animal. The slower movements of searching or playing about the mound are occasionally interrupted by a sudden leap directly upward to a height of 1-1/2 to 2 feet, often with no apparent reason other than play. This is, however, a fighting or guarding movement, though indulged in for play. The play instinct seems to be well developed, and in evidence on any moonlight night when actual harvesting operations are not going on.

STORING HABITS.

Probably no instinct is of greater importance to the kangaroo rat than that of storing food supplies. When a crop of desirable seeds is maturing the animal's activities appear to be concentrated on this work. During September, 1919, when a good crop of grass seed was ripening following the summer rains, a kangaroo rat under observation made repeated round trips to the harvest field of grass heads. Each outward trip occupied from 1 to 1-1/2 minutes, while the unloading trip into the burrow took only 15 to 20 seconds.

One individual in a laboratory cage, which had not yet been given a nest box, busied itself in broad daylight in carrying its grain supply into the darkest corner of the cage. When a nest box is supplied the individual will retreat into its dark shelter, and will only come forth after darkness has fallen unless forcibly ejected, but will store the food supplied.

In another case an animal escaped while being handled, and sought refuge behind a built-in laboratory table, where it could not be recovered without tearing out the table. For four days and nights it had the run of the laboratory. On the first night of its freedom it found and entered a burlap bag of grass seed that had been taken from a mound. A trail of seed and chaff next morning showed that it had been busily engaged in making its new quarters comfortable with bedding and food. After four nights of freedom it was captured alive in a trap, and later it was found that it had moved from the corner behind the table to the space beneath a near-by drawer, where it had stored about 2 quarts of the grass seed and a handful of the oatmeal used for trap bait.

BREEDING HABITS.

Observations on breeding habits have consisted mainly in taking records from the females trapped at all seasons of the year throughout the course of the investigation, and from examinations made during poisoning operations, and yet from this source the number of pregnant females taken or of young discovered is disappointingly small. The records indicate a breeding period of considerable length, extending from January to August, inclusive. It is possible that the length of the period may be increased by a second litter from the earliest breeding females in summer, but the large percentage of nonpregnant or nonbreeding animals which occurs throughout the season would indicate a wide variation in the time of breeding of different individuals.

Trapping in February and March for the purpose of securing greater numbers of female specimens, begun with the idea that these months were most likely to be the breeding months, has invariably yielded an unsatisfactory number of nonbreeding specimens and males. Unfortunately, the numbers of females secured in some months were not sufficient to be significant if worked out in percentages of breeding and nonbreeding individuals, and this, coupled with the fact that the importance of recording carefully all nonbreeders was not at first recognized, makes it impossible to tabulate such information reliably. The total of females taken in April, for example, is only 3, of which 1 was breeding; while in June, during the course of poisoning operations, 45 females were examined, of which 21 were breeding.

Five breeding females were taken in January, all during the last three days of the month. One of these was a suckling female, the young of which were secured alive and were probably at least a week old when taken. This must have been exceptionally early for young, since of a number of adult kangaroo rats taken during the first week of January none have been found to be breeding. Two records from Vernon Bailey are as follows: May 19-June 8, 1903, young specimen in nest (Santa Rosa, N. Mex.); June 12, 1889, one female, two embryos (Oracle, Ariz.).

The considerable proportion (which we believe to be more than 50 per cent) of nonbreeding females taken during all those months in which breeding has been found to occur may also indicate an extended period of breeding, with a small percentage breeding at any one time. This period also furnishes ample time for the rearing of two litters a year by some females, but we have no evidence as to the occurrence of two litters. Young of the year, practically grown, are taken during and after the month of April.

The mammae are arranged in three pairs, pectoral, 1/1; inguinal, 2/2.

Kangaroo rats are among those rodents in which the vagina becomes plugged with a rather solid material, translucent, and of the consistency of a stiff gelatine, after copulation. This must occur very soon after coitus, since in those individuals taken in this condition no definite evidence of the beginning of development of embryos could be detected by examination.

The length of the gestation period of spectabilis is unknown. The young are born naked, a fact inferred by failure to find any fetus showing noticeable hair development, and from the conditions observed in such young as have been seen. A suckling female was taken by Vorhies, January 31, 1920, and her den immediately excavated in the hope of securing the young. Two juveniles were found in a special nest chamber (see p. 30). These were estimated to be perhaps two weeks old. A serious effort was made to raise the little animals by feeding milk with a pipette and keeping them warm with a hot water bottle, but they survived only 10 days, without the eyes having opened. The uneven temperature as well as the character of the food was probably responsible for their deaths. On February 3 they were measured and weighed, with the following results:

At this stage the young were partially clothed with a coat of fine velvety fur, more especially on the bodies, the tails being still nearly naked. The body color was dark plumbeous, just the color of the dark underfur of the adult, or a shade darker, while the characteristic white markings of the adult stood out sharply as pinkish-white areas against the dark background (see Pl. IX, Fig. 2, at p. 32). The proportions were much as in the adult, except that the tails were relatively much shorter and the feet relatively longer.

Only one other record of young is at hand, that by Bailey, who secured the young after capture of a suckling female at Santa Rosa, N. Mex. In this case the litter contained only one. This was squeaking when found, but was not large enough to crawl away. Its eyes and ears were closed, and its soft, naked skin was distinctly marked with the pattern of the adult, the colors being as given for the other two. This juvenile lived only a week. Young less than half grown were not trapped or noted in our poisoning operations outside the dens.

Kangaroo rats, if spectabilis be representative, reproduce at a slow rate as compared with many other small rodents. We have records of 67 females with embryos or scars showing the number produced, and of the two litters of young described above. Of the 69 females thus recorded, 15, or 21.7 per cent, had but one offspring each; 52, or 75.3 per cent, but two each; while only 2 individuals, or 2.9 per cent, had three. Three young is the maximum litter recorded. This, taken in connection with the protracted breeding season and lack of sure evidence of the production of two broods a year, gives a surprisingly low rate of reproduction, indicating relative freedom from inimical factors.

Our breeding records for merriami are fewer than for spectabilis, but are very similar in every way so far as they go, both as to the time of year and number of young.