A small mound of _Dipodomys s. spectabilis_ in early autumn, showing runways radiating from the den. Evidences of activity may be noted in and about the surface of the mound.]

[Ill.u.s.tration: PLATE VI. FIG. 1.--RUNWAY OF DIPODOMYS S. SPECTABILIS.

Well-traveled path leading from the main den, in the foreground, to a subsidiary burrow (see Fig. 2, below), about 30 feet distant, at apparent end of runway.]

[Ill.u.s.tration: PLATE VI. FIG. 2.--SUBSIDIARY BURROW OF DIPODOMYS S.

SPECTABILIS.

Located at the end of the 30-foot runway shown in Figure 1, above. This has three openings, two in the foreground and the third a little to the rear and indicated by an arrow.]

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 m.u.f.fled 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 gra.s.s 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 Willc.o.x, 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 ma.s.s 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 c.o.c.ks, 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 ga.s.sing 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 pa.s.sages 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 vibrissae 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 ATt.i.tUDES.

Movements and att.i.tudes 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 a.s.sumes 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 gra.s.s seed was ripening following the summer rains, a kangaroo rat under observation made repeated round trips to the harvest field of gra.s.s 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 gra.s.s 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 s.p.a.ce beneath a near-by drawer, where it had stored about 2 quarts of the gra.s.s 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 v.a.g.i.n.a 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:

---------------------------------------------------Measurements (in millimetres).

Weight-------------------------------(inTotalTailHindgrams).length.vertebrae.foot.

----------------------------------------------- No. 113.3903824 No. 212.6933824 ---------------------------------------------------

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.

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