Part 20

Externals.--Size small to medium (total length 188 mm to 216 mm); tail longer than head and body (112 mm to 134 mm) and bicolored, pale brown to brownish-black above, white to yellowish-white below; hind feet long (28 mm to 31 mm), grayish-white above; back ochraceous to dark brown; sides paler than back with dark hair interspersed; lateral line usually present but sometimes indistinct or entirely absent (when present usually clear Ochraceous-Buff); ventral coloration white, sometimes with suffusion of ochraceous; guard hairs average 115 microns (96u to 140u) in diameter; underhair with pigment pattern in form of hollow, narrow rectangles; cuticular scales of underhair large and fewer than those of the underfur of Z. trinotatus, but underhair of Z. hudsonius otherwise resembles that of Z. trinotatus.

Baculum.--Size small (total length 4.5 mm to 4.9 mm); base medium in width (0.64 mm to 0.72 mm); tip narrow (0.24 mm to 0.26 mm) and dished out in dorsal aspect, blunted; shaft rounded, curving gently upward at tip.

Skull.--Small to medium and relatively narrow in relation to length; rostrum pointed and short; mastoid region relatively narrow; incisive foramina short; base of zygomatic process of squamosal narrow; coronoid process of mandible short, relatively weak. Upper premolar usually small (averaging .30 mm in length and .35 mm in breadth) sometimes functional (most often so in old adults), occlusal surface divided by single shallow re-entrant fold, which in worn teeth forms centrally located lake; tooth-row short as compared to that of other species; individual cheek-teeth usually smaller than those of other species; lower cheek-teeth shorter and narrower than those of other species; angle of mandible strongly inflected.

GEOGRAPHIC VARIATION

The species Z. hudsonius is divisible into 11 subspecies based on differences in color, relative proportions of the tail, hind feet, body, and size and shape of parts of the skull (zygomata, braincase, incisive foramina, auditory bullae, pterygoid fossae, rostrum, and interorbital breadth).

Color of the pelage varies, as a general rule, from dark-backed, dull-sided individuals in the northern parts of the geographic range of the species to light-backed, bright-sided individuals in the southern parts of the range.

Individuals from the southernmost geographic races (Z. h. americanus and Z. h. pallidus) are the smallest for the species and those from the northernmost subspecies (Z. h. alascensis) are the largest. One subspecies, Z. h. campestris, from the central part of the range of the species, however, seems to be out of the cline. This form inhabits the eastern foothills of the Rocky Mountains and is a robust animal approaching Z. princeps in size.

Seemingly there is no clinal variation in the several qualitative features of the cranium, for instance in the shape of the auditory bullae, shape of the incisive foramina, and shape of the postpalatal notch. On the other hand, the dimensions of the entire skull show that the larger crania are of the northernmost subspecies and the smaller of the southernmost subspecies.

NATURAL HISTORY

Habitat.--Zapus hudsonius occurs in low undergrowth usually of grasses or forbs or both, in open coniferous forests, deciduous hardwood groves, or in stands of tall shrubs and low trees, but most frequently in open, moist areas.

Quimby (1951:75) notes that jumping mice were more common in the moist lowlands than in the drier uplands. More were in the open type lowlands than in the forested type, and these mice favored habitats normally bordered by small streams affording moist to semi-aquatic living conditions. The reports of Goodwin (1924:255), Christian (1936:416), G. S. Miller (1899:329), Cory (1912:249), Lyon (1936:277), Stoner (1918:123), and others, although concerning widely different parts of North America, indicate that Z. hudsonius selects habitats in vegetation of like form, even though different assemblages of plant species may be involved.

An average of 11.91 mice per acre was recorded by Quimby (1951:91) from a study plot at Itasca Park, Clearwater County, Minnesota. He gives the monthly population densities per acre for Z. hudsonius at Centerville, Anoka County, Minnesota, as follows: June 2.78, July 3.57, August 3.10, and September 1.81. Blair's (1940:248) data on bi-monthly population density per acre for Z. hudsonius on the Edwin S. George Reserve, Livingston County, Michigan, are remarkably similar, when adjusted on a monthly basis, to those obtained by Quimby (loc. cit.). Blair's (loc. cit.) monthly population densities per acre are as follows: June 3.90, July 3.85, August 3.10, and September 2.00. Townsend (1935:90) estimated population densities per acre for Z. hudsonius in central New York state, at 11 to 72 individuals. As Quimby (1951:92) points out, Townsend's figures are probably too high, as commonly is the case when the moving quadrat technique is used because animals from neighboring areas enter the trapped area to take over the niches made available by their predecessors' removal.

The population of Z. hudsonius may vary considerably from year to year as well as seasonally. Blair (1940:249) found notably fewer jumping mice on the George Reserve in 1938 than in 1939. Quimby (1951:94) found the numbers of Zapus to be highly variable and thought that there was a rapid turnover. Young animals were not caught until July when 25 per cent were either juveniles, young, or subadults; from this time on these age classes increased to a high of sixty-one per cent in September. Quimby (loc. cit.) found that separating the individuals into their proper age classes was more difficult in September, since the young from early litters are adultlike in appearance. His data indicate as he remarked, "That the over-wintering adults are, for the most part, gradually replaced by the young of the year as the summer progresses."

The sexes in Z. hudsonius vary only slightly from a one to one ratio. Quimby (1951:63) found a sex ratio of 110 females to 100 males and Blair (1940:245) records a sex ratio of 113 males to 100 females. Townsend (1935:42) records a sex ratio in central New York of 155 males to 100 females. Such a wide variation from a one to one ratio suggest that the moving quadrat technique, which Townsend (1935:90) employed in obtaining his data, may be, in some way unknown to me, more selective for the males.

Behavior.--The saltatorial powers of Z. hudsonius are well developed and often have been described in the literature. Stoner (1918:123) remarks that, "When disturbed hudsonius moves away by a series of leaps ... the distance traversed in one of these leaps is from six to eight feet."; Cory (1912:249) observed these mice to make surprisingly long leaps, and, according to him, a distance of 10 feet is by no means unusual; Handley and Patton (1947:49) credit these animals with jumping eight to ten feet at a single bound; Hamilton (1935:190) remarked that he noted an average of not more than four to six feet per jump; Townsend (1935:91) observed one individual make jumps of about two feet; and Harper (1932:29) records a jumping mouse leaping for distances of two to three feet. Quimby (1951:72) notes that he had never seen one jump farther than three feet. He found that the greatest jumps occurred initially and normally covered a distance of two to three feet; subsequent leaps were shorter but more rapid. A jumping mouse in full retreat progressed by jumps of about one foot.

Statements concerning the gait of Z. hudsonius are not in agreement but the consensus of opinion is that these animals when unfrightened progress by a series of hops of one to six inches, or, occasionally, with a slow creeping motion while the animal is on all fours. When frightened, however, their progress is by long bounds; the mice make a series of two or three such leaps to the nearest protective cover, and then sit motionless until pursued.

Concerning the use of the tail as a balancing organ, G. S. Miller (1899:330) describes the behavior of a jumping mouse from which the tail had been severed by the sickle of a mowing machine. "When I approached, it made violent efforts to escape, but the moment it was launched in the air, its body, deprived of its balancing power, turned end over end so that it was as likely as not to strike the ground facing the direction from which it had come."

Riparian animals such as Z. hudsonius need enter the water to escape from enemies or perhaps in search of food. Zapus hudsonius can and does swim. Hamilton (1935:190) found it to be a strong swimmer capable of remaining in the water for from four to five minutes. According to Hamilton (loc. cit.), when the mouse is swimming the head is held high, the tail is arched near its middle, and only the hind limbs are employed in propulsion. According to Sheldon (1938:327), Philip Allan, in northern Minnesota, saw many Z. hudsonius swimming three or four inches under the surface of the water. The mice swam upstream and only the hind legs were employed in the swimming movements. N. A. Preble (1944:200), at Archer's Pond, 3 miles southeast of Center, Ossipee County, New Hampshire, observed a jumping mouse swimming rapidly under water toward another portion of the shore 30 or 40 feet away. The mouse, swimming less than a foot beneath the surface, was vigorously using both forefeet and hind feet, but the long tail trailing limply behind, contributed in no way to the animal's movements. Quimby (1951:72) released five of the mice, one at a time, in the open water of a lake. He followed alongside in a boat and observed that, "In all instances the animals proved to be excellent swimmers both on and underneath the surface. The methods of progression were similar to land movements; i. e., the limbs were employed differently at various times depending upon the speed. When first placed in water they moved rapidly by lunges produced by sweeping strokes of the hind limbs employed simultaneously. This movement was accomplished similarly to the long jumps made on land ... Following the first excited lunges, they settled down to a steadier and slower gait using all four limbs one at a time. The anterior part of the body was held high in the water ... When approached too closely, they attempted to escape by diving. The maximum distance noted was about four feet ... One was able to swim vigorously for approximately three minutes after which it tired greatly and was in danger of drowning."

As concerns digging ability, Goodwin (1935:148) reports that Z. hudsonius makes its own burrows; these are short and close to the surface in the summer but longer, deeper, and below the frost-line in winter. Two captives used their forefeet and nails in digging a tunnel in the foot of soil that Goodwin (loc. cit.) had placed in their cage. Quimby (1951:72) remarks that captives excavate soil by means of the front feet and throw the soil out behind; as the burrow deepened the hind feet were also utilized to throw the loose soil out of the burrow.

Zapus hudsonius climbs; Sheldon (1934:293) observed captive animals to climb over small evergreen trees in their cages. They moved with surprising sureness and agility, chasing each other among the branches or sitting for several minutes at a time on one of the limbs. Hamilton (1935:190) found that the mice ran over limbs and brush which were placed in their outdoor enclosure.

Ordinarily Z. hudsonius is nocturnal, appearing in the early dusk and remaining active until pre-dawn. Occasional individuals are abroad in daylight hours. Sheldon (1934:293) found in Nova Scotia that Z. hudsonius is most active from early dusk through the night, but that it may be abroad in daylight as well. Her statements are based on trapping results, field observations, and observations made on captive individuals. Quimby (1951:73) found that Z. hudsonius in Michigan is mostly nocturnal; however, he saw mice on a few occasions in the daytime. Diurnal activity seems to be increased in cloudy or damp weather; Quimby (loc. cit.) almost invariably trapped more of these mice on cloudy, damp days than on other days.

This jumping mouse usually is silent but does utter various sounds. Sheldon (1934:295) records squeaking and clucking noises. Quimby (1951:73) records the clucking noise described by Sheldon (loc. cit.) and mentions also the squeaking and suckling sounds produced by the small young. This mouse is most vociferous when young or when about to go into hibernation. Sheldon (1938:327) writes that Z. hudsonius makes a drumming noise by vibrating the tail against dry leaves.

Many data are available concerning the hibernation of Z. hudsonius. In general it seems necessary for the mice to put on a certain amount of fat preparatory to hibernation. This fat is deposited in a thin layer over the inside of the skin, over the back, and in the body cavities. The thickest deposits are in and about the inguinal region.

Quimby (1951:83) noted that gain in weight was accelerated in a brief period prior to entrance into hibernation. This relationship of rapid gain in weight to hibernation allows a person to estimate the date of hibernation. Cold weather seems to hasten hibernation, but less so than the correct physiologic condition which is foreshadowed by a rapid gain in weight. For example, Quimby's (1951:84) data reveal that mice that were moved to a heated room gained weight and hibernated in a fashion similar to those in unheated surroundings. Hamilton (1935:193) states that, "It seems necessary for the mouse to lay on a certain amount of fat before it is capable of hibernation." Hamilton (loc. cit.) reported that 18 specimens of Z. hudsonius taken [presumably in an active state] near Ithaca, New York, on November 13, were without a trace of fat.

Data that are available concerning the hibernation sites of Z. hudsonius show that almost invariably these mice seek shelter in burrows beneath the surface of the ground and there construct nests of grass, leaves, or some other vegetation. Nicholson (1937:103) found a hibernating Z. hudsonius on the George Reserve, Livingston County, Michigan, on October 20. The mouse was in a nest, composed of 10 to 12 damp elm leaves, in a sand bank two feet three inches vertically and three feet nine inches horizontally from the surface. On April 11, 1948, Schwartz (1951:228) found five nests (three with occupants) of Z. hudsonius at Jefferson City, Cole County, Missouri. All nests were one foot beneath the surface of a pile of coal-ash, which was about three and one-half feet high and five feet in diameter. The nests were spherical, approximately four inches in diameter and consisted of dried oak leaves and bits of dried grass. Grizzell (1949:74) found two hibernating jumping mice at the Patuxent Research Refuge, Laurel, Maryland, in January, 1948. The mice were in separate woodchuck dens; one mouse was 40 inches below the surface and the other was 26 inches below the surface. The mice were curled up in the center of masses of dead leaves, and thus, were well insulated against the cold. On April 29, 1944, at Ithaca, New York, Eadie (1949:307) uncovered a hibernating jumping mouse. The nest, about the size of a baseball, was compactly made of fine grasses and was 10 inches below the surface of the ground in a mound of earth that was approximately six by four feet at the base and three feet high.

From the foregoing reports on hibernation sites it is evident that well drained areas are utilized. Sheldon (1934:300) remarks that the burrows used for hibernating are dug in a bank or some place from which the rain water and melted snow probably drains off.

Eadie (1949:307), Grizzell (1949:75), Sheldon (1934:299), Schwartz (1951:228), and Sheldon (1938:331) all agree that the hibernating mouse rolls up into a ball-like shape (resting on its head and pelvis) with the head between the hind legs, the nose against the lower belly, the forefeet curled on the chest, and the tail curled around the head and body.

A marked loss of weight occurs immediately after hibernation begins, and then reduction in weight is slow and regular. (See Hamilton, 1935:194 and Quimby, 1951:84.)