A Population Study of the Prairie Vole (Microtus ochrogaster) in Northeastern Part 3

TABLE 3. DISTRIBUTION AMONG AGE GROUPS OF 21 VOLES USED IN THE STUDY OF VARIATION DUE TO AGE

====================================================================== Age in months 1-1/2 2 2-1/2 3 3-1/2 4 4-1/2 6 12 ---------------------------------------------------------------------- No. of individuals 1 4 5 1 3 2 3 1 1 ----------------------------------------------------------------------

In the series of voles studied, ten individuals were in the process of molting from subadult to adult pelage. Jameson (1947:131) reported the molt to occur between eight and 12 weeks of age and selected 38 grams as the lower limit of weight of adults. I also found all voles molting to be between eight and 12 weeks old but found none so large as 38 grams without full adult pelage. This may have been, in part, due to the dry weather delaying or inhibiting growth. Because of the small size of the sample and the influence of the unusual weather conditions, no conclusions concerning normal molting were drawn from the data described below. They are presented only as a description of a small sample drawn from a single population at one time. Table 4 summarizes these data.

TABLE 4. MEAN SIZES AND AGES OF VOLES MOLTING FROM SUBADULT TO ADULT PELAGE

===================================================================== Body length Condylo-basilar Weight minus tail length Age --------------------------------------------------------------------- Six males 32.67 gms. 106.16 mm. 23.78 mm. 9.67 wks.

(30-36) (96-116) (23.2-24.4) (8-12) Four females 29.0 gms. 100.25 mm. 23.45 mm. 10.5 wks.

(28-30) (98-102) (23.5-23.8) (8-12) Ten voles 31.2 gms. 103.8 mm. 23.73 mm. 10.0 wks.

(28-36) (96-116) (23.2-24.4) (8-12) ---------------------------------------------------------------------

The mean age of the ten voles molting was ten weeks (8-12). Six males averaged 9.67 weeks, almost a week younger than four females, who averaged 10.5 weeks. The difference in age at time of molting between the s.e.xes was not significant. Differences between the s.e.xes in other characteristics to be described also lacked significance. Mean weights at the time of molting were: males, 32.67 gms. (30-36); females, 29.0 gms. (28-30); and all individuals, 31.2 gms. (28-36). Because a piece of the tail of each vole had been removed in marking, the total length of the voles could not be determined. Body length, excluding tail, was used. Howell (1924:986) found this measurement subject to less individual variation than total length and thought body length was probably a better indicator of age. Mean body length at the time of molting was 103.8 mm. (96-116). Males averaged longer than females and were also more variable. The mean body length of males was 106.16 mm.

(96-116) and that of females was 100.25 mm. (98-102).

Of the subadults showing no signs of molting, none was above the mean age of molting. Twenty-five per cent of them were longer and heavier than the mean length and weight of those that were molting. Of the 20 adults in the series, one was below the mean weight of molting and one was shorter than the mean length of molting.

When Howell (_op. cit._:1014) studied skulls of _Microtus monta.n.u.s_ he found that the condylobasilar length was the most satisfactory means for arranging his series of specimens according to their age. When the skulls of my series were arranged according to their age (as determined from trapping records) the graph of the condylobasilar lengths showed a clear, though not perfect, relationship to age (Fig. 13). No separation of s.e.xes was made because the sample did not permit it. In Fig. 13 graphs of weight, as determined in the field, and of length (excluding tail) also were included because they are the most easily measured characters of live voles. The graphs indicate individual variation in these characters which limits their usefulness in determining age.

[Ill.u.s.tration: FIG. 13. Graphs of the condylobasilar lengths, body lengths and weights of a series of voles of known age. Within each age group, the youngest vole is on the left in the graphs.]

When other cranial measurements, and ratios of pairs of measurements, were plotted in the same order, individual variation obscured some of the variation due to age and the curves resembled those of weight and length of body rather than that of condylobasilar length. When the cranial measurements were averaged for the age groups the curves showed a relationship to age but the relationship of mean measurements is of little use in determining the age of individual specimens. The data described above indicated that a study of the relationship of the condylobasilar length and age in a large sample might provide useful information.

Anyone who has examined mammalian skulls knows of many other characters which vary with age but which are difficult to measure and describe with precision. Figs 14 and 15 are drawings of skulls of voles of known age.

The most obvious change, related to aging, evident in the dorsal view of the skulls (Fig. 14) is the increasing prominence and closer approximation of the temporal ridges in older specimens. The lambdoidal ridge is also more prominent in older voles, and their skulls have a generally rougher and more angular appearance. The individual variation evident in these ridges is probably due to variations in the development of the muscles operating the jaws (Howell, 1924:1003). There is an increased flattening of the roof of the skull of older voles.

[Ill.u.s.tration: 1-1/2 months 2-1/2 months 3 months 3-1/2 months

4 months 4-1/2 months 6 months 12 months

All 3.

FIG. 14. Dorsal views of skulls of voles of known age.]

[Ill.u.s.tration: 1-1/2 months 2-1/2 months 3 months 3-1/2 months

4 months 4-1/2 months 6 months 12 months

All 3.

FIG. 15. Palatal views of skulls of voles of known age.]

From a palatal view (Fig. 15) the skulls of voles also showed age variation which was apparent but not easily correlated with precise age.

The median ridge on the basioccipital bone increases in prominence in older voles. The shape of the posterior margin of the palatine bones changes from a V-shape to a U-shape. On the skull of the oldest (12 months) vole the pterygoid processes are firmly fused to the bullae, a condition not found in any of the other specimens. The anterior spine of the palatine approaches the posterior projection of the premaxillae more closely as age increases and, in the oldest vole is firmly attached and forms a complete part.i.tion separating the incisive foramina.

Tooth wear during the life of a vole causes a considerable variation in the enamel patterns, especially of the third upper molar. Howell (1924:1012) considered such variation to be independent of age, but Hinton (1926:103) related the changes to age and interpreted them as a recapitulation of the evolution of microtine molars. In my series, an indentation on the medial margin of the posterior loop of the third upper molar seemed to be related to age. This indentation was absent in the youngest vole (one and one-half months), absent or indefinite in those voles less than 3-1/2 months of age, and progressively more marked in the older voles.

Food Habits

The prairie vole, like other members of the genus _Microtus_, feeds mostly on growing gra.s.s in spring and summer. Piles of cuttings in the runways are characteristic sign of the presence of voles. The voles cut successive sections from the bases of gra.s.ses until the young and tender growing tips are within reach. The quant.i.ty of gra.s.s destroyed is greater than that actually eaten, a fact which will have to be considered in any attempt to evaluate the effects of voles upon a range.

In all piles of cut plants that were examined, _Bromus inermis_ was the most common gra.s.s, and _Poa pratensis_ was the gra.s.s second in abundance. These were, by far, the most common gra.s.ses present on the areas studied; in most places, _B. inermis_ was dominant. Other gra.s.ses present on the areas were occasionally found in the piles of cuttings.

Jameson (1947:133-136) found no utilization of _B. inermis_ by voles but that gra.s.s was present in a relative abundance of only one per cent in the areas studied by him. The voles that he studied ate alfalfa in large amounts and alfalfa was, perhaps, the most common plant on the particular areas where his voles were caught. Seemingly, the diet of voles is determined mostly by the species composition of the habitat.

Other summer foods included pokeberries, blackberries and a few forbs and insects. Forbs most commonly found in the piles of cuttings were the leaves of the giant ragweed (younger plants only) and dandelion. Insect remains were found in the stomachs of voles killed in summer and occurred most frequently in those killed in August and September. At no time did insects seem to be a major part of the diet but they were present in most vole stomachs examined in late summer. Laboratory experiments with summer foods gave inconclusive results but suggested that the voles chose gra.s.ses on the basis of their growth stage rather than according to their species. Young and tender gra.s.ses were chosen, regardless of species, when various combinations of _Triodia flava_, _Bromus inermis_ and _Poa pratensis_ were offered to the voles. At times the voles showed a marked preference for dandelion greens, perhaps because of their high moisture content; the voles' water needs were satisfied mostly by eating such succulent vegetation.

Winter foods consisted of stored hay and fruits and of underground plant parts. _Bromus inermis_ made up nearly all of the hay and was stored in lengths of up to ten inches in underground chambers specially constructed for storage. Underground parts of plants were reached by tunnelling and were an especially important part of the voles' diet in January and February. The fruit of _Solanum carolinense_ was eaten throughout the winter and one underground chamber, opened in February, 1952, was packed full of these seemingly unsavory fruits. Fisher (1945:436), in Missouri, found this fruit to be an important part of the winter diet of voles. An occasional pod of the honey locust tree was found partly eaten in a runway. Fitch (1953, _in litt._) often observed girdling of honey locust and crab apple (_Pyrus ioensis_) root crowns on the Reservation but I saw no evidence of bark eating, perhaps because my study plots were mostly gra.s.sland. On two occasions when two voles were in the same trap one of them was eaten. In both traps, all of the bait had been eaten and the captured voles probably were approaching starvation. Because the trapping procedure offered abundant opportunity for cannibalism, the low frequency of its occurrence suggested that it was not an important factor in satisfying food requirements under normal conditions.

Runways and Nests

Perhaps the most characteristic sign of the presence of _Microtus ochrogaster_ were their surface runways and underground tunnels. Only rarely was a vole observed to expose itself to full view. When a trapped vole was released it immediately dove out of sight into a runway. Once in a runway, the vole showed no further evidence of alarm and was usually in no hurry to get away. The runways seemed to provide a sense of security and the voles were familiar with their range only through runway travel. The urge to seek a runway immediately when exposed has obvious survival value.

Surface runways were usually under a mat of debris. In areas where debris was scanty or lacking, runways were usually absent. Jameson (1947:136) reported that in alfalfa and clover fields the voles did not make runways as they did in gra.s.sland, even in fields where trapping records showed voles to be abundant. Typical surface runways are approximately 50 mm. wide, only slightly cut into the ground and bare of vegetation while in use. Usually they could be distinguished from the runways of the pine vole, which were cut more deeply into the ground, and those of the cotton rat which were wider and not so well cleared of vegetation. Some runways ended in surface chambers and some of these were lined with gra.s.s. Their size varied from a diameter of 90 mm. to 250 mm. and they seemed to be used primarily for resting places.

A runway system usually consisted of a long, crooked runway and several branches. Two typical systems are ill.u.s.trated in Fig. 16. The runway systems often were not clearly limited; they merged with other systems more or less completely. One map showed a runway system extending across 140 square meters and including 12 underground burrows. All of these runways seemed to be part of a single runway system but the system probably was used by more than one vole or family group of voles.

Sixteen of the 22 maps that were made extended across areas between 50 and 90 square meters. One map, mentioned above, was larger and the remaining five smaller. The smallest extended across only 20 square meters. Of course, the area encompa.s.sed by a set of runways changed almost daily, as the voles extended some runways, added some and abandoned others in the course of their daily travels.

[Ill.u.s.tration: FIG. 16. Maps of runway systems of the prairie vole. The runways follow an irregular course and are frequently changed. The solid lines represent surface runways and the dotted lines underground pa.s.sages.]

Each runway system contained underground nests. These were in chambers from 70 mm. to 200 mm. below the surface and were up to 200 mm. in diameter. Most systems that were mapped had from two to six of these burrows. Most of these were lined with dried gra.s.s and seemed to be used for delivering and nursing litters. Each burrow was connected to a surface runway by a tunnel. Often the tunnel was short and the hole opened almost directly into the burrow from the surface runway. Others had tunnels several meters long. Jameson (1947:137) reported every burrow to have two connections with the surface. In the present study, however, I found three arrangements in approximately equal frequency of occurrence: (1) one hole to one tunnel leading to a burrow; (2) two holes to two short tunnels which joined a long tunnel leading to a burrow; and (3) two separate tunnels from the surface to a burrow. The size, depth and number of underground burrows in the systems that I studied varied and so did those reported in the literature. Jameson (_loc. cit._) found burrows in eastern Kansas as deep as 18 inches, far deeper than any found in my study. Fisher (1945:435) reported none deeper than five inches in central Missouri. The soil data in my study, as well as in the two reports cited immediately above, were not adequate to permit conclusions, but the type and condition of the soil probably determine the extent of burrowing by the voles of any given locality.

The number of voles using a runway system at one time was difficult to ascertain. In one system, however, four adult individuals were trapped in a ten day period. In August, 1952, at the conclusion of the live-trapping program, a runway system was mapped which had included two trapping stations. In the preceding ten days, four adult voles (three males and one female) had been taken in both traps. During that time, therefore, the runway system was shared by at least four voles. The voles used an area that was considerably larger than that encompa.s.sed by any one runway system, a fact obvious when the sizes of home ranges as computed from trapping data were compared with the sizes of the runway systems mapped. A runway system seemed not to be a complete unit, but was only a part of the network of runways used by a single individual.

Activity

Although no special investigation of activity was made, some conclusions concerning it were apparent in the data gathered. There have been a few laboratory studies of the activity pattern of _Microtus_ by various methods. Calhoun (1945:256) reported _M. ochrogaster_ to be mainly nocturnal with activity reaching a peak between dark and midnight and again just before dawn. Davis (1933:235), working with _M. agrestis_, and Hatfield (1935:263), working with _M. californicus_, both found voles to be more nocturnal than diurnal. In a field study of _M.

pennsylvanicus_, Hatt (1930:534) found the species to be chiefly nocturnal, although some activity was reported throughout the day.

Hamilton (1937c:256-259), however, reported the same species to be more active in the daytime. Agreement on the activity patterns of these species of _Microtus_ has not yet been attained.

From occasional changes in the time of tending a trap line, and from running lines of traps at night a few times in the summer of 1951, I gained the impression that these voles were primarily diurnal.

Relatively few of them were caught in the hours of darkness. In summer, however, their activity was mostly limited to the periods between dawn and approximately eight o'clock and between sunset and dark. In colder weather, there was increased activity on sunny days.

PREDATION

Although voles were a common item of prey for many species of predators on the Reservation, no marked effect on the density of the population of this vole could be attributed to predation pressure. Only when densities reached a point that caused many voles to expose themselves abnormally could they be heavily preyed upon. Their normally secretive habits, keeping them more or less out of sight, suggest that they are an especially obvious ill.u.s.tration of the concept that predation is an expression of population vulnerability, rising to high levels only when a population is ecologically insecure, rather than a major factor regulating population levels (Errington, 1935; 1936; 1943; Errington _et al_, 1940).

Scats from predatory mammals and reptiles and pellets from raptorial birds were examined. Most of these materials were collected by Dr. Henry S. Fitch, who kindly granted permission to use them. The results of the study of the scats and pellets are summarized in Table 5. Remains of voles were identified in 28 per cent of the scats of the copperhead snake (_Ancistrodon contortix_) examined. Copperheads were moderately common on the Reservation (Fitch, 1952:24) and were probably important as predators on voles in some habitats. Uhler _et al_ (1939:611), in Virginia, reported voles to be the most important prey item for copperheads. A vole was taken from the stomach of a rattlesnake (_Crotalus horridus_) found dead on a county road adjoining the Reservation. Rattlesnakes were present in small numbers on the Reservation but were usually found along rocky ledges rather than in areas where voles were common (Fitch, _loc. cit._). The rattlesnakes probably were less important as predators on voles than on other small mammals more common in the usual habitat of these snakes. The blue racer (_Coluber constrictor_) was common in gra.s.sland situations on the Reservation (Fitch, 1952:24) and twice was observed in the role of a predator on voles; one small blue racer entered a live-trap in pursuit of a vole and another blue racer was observed holding a captured vole in its mouth. The blue racer seems well adapted to hunt voles and probably preys on them extensively. The pilot black snake (_Elaphe obsoleta_) has been reported as a predator on _M. ochrogaster_ in the neighboring state of Missouri (Korschgen, 1952:60) and was moderately common on the Reservation (Fitch, _loc. cit._). _M. pennsylvanicus_, with habits similar to those of _M. ochrogaster_, has been reported as a prey for all of the above snakes (Uhler, _et al_, 1939).

TABLE 5. FREQUENCY OF REMAINS OF VOLES IN SCATS AND PELLETS