Edwin Perry Martin

“ 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. ”

“ The autumn rise in population may have been due, in part, to the addition of spring and early summer litters to the breeding population, but the rise occurred too late in the year to be explained by that alone. Another factor may have been the spurt in growth of grasses occurring in Kansas in early autumn, in September and October. There was a seeming correlation between high rainfall with rapid growth of grasses and reproductive activity, and, secondarily with high population densities of voles. These relationships are discussed in connection with reproduction. ”

“ The causative factor directly influences the population either by inhibiting reproduction or by increasing infant and prenatal mortality. The net reduction is due to an inadequate replacement of those voles lost by normal attrition.
Most voles, under natural conditions, live less than one year. Those individuals born in the autumn live longer, as a group, than those born at any other time. Since the heaviest mortality is in young voles, adults which become established in an area may live more than 18 months and, if they are females, may produce more than a dozen litters. ”

“ Those traps set in spots where there was little vegetation other than the dominant grass caught fewer voles than traps set in spots with a more varied cover. Poa pratensis formed an understory over most of the area studied, especially on House Field, and attained local dominance in shaded spots on both fields. The higher basal cover provided by the Poa understory seemed to support a vole population larger than those that occurred in areas lacking the bluegrass. Disturbed situations, such as roadsides, were characterized by the dominance of Bromus japonicus. ”

“ A home range may include several runway systems, and the ranges of individuals overlap extensively. Both home ranges and patterns of runway systems change constantly. Runways seem to be primarily feeding trails, and are extended or abandoned as the voles change their feeding habits. Groups of adult voles using a system of runways seem to have no special relationship. Juveniles tend to stay near their mothers, but as they mature, they shift their ranges and are replaced by other individuals. Males wander more than females, and shift their ranges more often. ”

“ The percentage of gravid females in the population could not be determined accurately by a live-trapping study and was not useful in this investigation. The percentage of juveniles trapped in the month following the sampling period tended to follow the curve of the percentage of adult females with a vaginal orifice. The ratio of trapped juveniles to adults trapped was a poor indicator of reproductive activity. ”

“ 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. ”

“ Most individuals were confined to the grassy areas around the field and made only occasional forays away from the edge. The dam of a small pond on the Reservation and low ground near the water were used by Microtus at all times. In the summer of 1949 no voles were taken anywhere on the Reservation but their runways were more abundant around the pond than in the other places examined. Of all the areas studied in the summer of 1949, only the pond area had been protected from grazing in previous years. ”

“ In House Field the herbaceous vegetation was much more lush than in Quarry Field and woody plants and weeds were more abundant. A graveled and heavily used road along one edge of House Field, leading to the Reservation Headquarters, was a barrier which voles rarely crossed. A little-used dirt road crossing the trapping plot in Quarry Field constituted a less effective barrier. The disturbed areas bordering the roads were likewise little used and tended to reinforce the effects of the roads as barriers. There were almost pure stands of Bromus japonicus along both roads. ”

“ A relationship between the condylobasilar length of the skull and the age of a vole was discovered and, with further study, may yield a method of aging voles more accurately than has been possible heretofore. Other characteristics, varying with age, were described. The most reliable indicator of age seemed to be the prominence of the temporal ridges.
Runway systems and burrows are used by groups of voles rather than by individuals. Most of the activity of voles is confined to these [411] runways and an exposed individual is seldom seen. ”

“ That this depressing effect involved more than direct competition for plant food was suggested by the fact that in House Field, with a heavy crop of vegetation and a seemingly high carrying capacity for both herbivorous rodents, the biomass of voles, and of all rodents combined, were lower than in Quarry Field which had less vegetation and fewer cotton rats. ”

“ As mentioned earlier, when House Field and Quarry Field were trapped out at the conclusion of the live-trapping program no unmarked voles were taken. This showed that the 30 foot interval between traps was short enough to cover the area as far as Microtus was concerned. The fact that unmarked adults were caught almost entirely in marginal traps is additional evidence. On the other hand, the Fitch traps were 50 feet apart and voles seemed to have lived within the grid for several months before being captured. ”

“ 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 grassland, 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. ”

“ Population trends in either direction may be reversed suddenly by changes in the factors discussed above. In the fall of 1951, a downward trend in the density of the voles was evident. At this time, populations of cotton rats were increasing rapidly and competition between cotton rats and voles was intensified. In February, 1952, the population of cotton rats was decimated suddenly by a short period of unusually cold weather. The voles were suddenly freed from the stress of competition and the population immediately began to rise. ”

“ 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. ”

“ Although a few cotton rats were taken in Quarry Field and still fewer in Reithro Field, the population of those hilltop areas did not approach, at any time, the levels reached on House Field, which produced a more luxuriant cover. Only when the levels of population were exceptionally high did the cotton rats spread into less productive habitats. At all times, there were areas on the Reservation used by Microtus which could not support a population of Sigmodon. ”

“ The correlation coefficient between the percentage of captures involving both sexes and the level of reproductive activity was 0.58. Among those pairs of individuals caught together more than once, 61 per cent were composed of both sexes. Among those pairs taken together three or more times 76 per cent were male and female and among those pairs taken together four or more times 80 per cent were male and female. When adult voles stayed together any length of time their relationship usually appeared to be connected with sex. ”

“ The mean number of embryos per female was higher than the mean number of young per litter delivered in captivity and the mean number of young per litter delivered in traps was lower than in those delivered in captivity. The differences were not statistically significant. In some instances females that delivered young voles in traps may have delivered others prior to entering the trap or the mother or her trapmates may have eaten some of the newborn voles before they were discovered. ”

“ Presumably these animals, born in autumn and early winter, were more vigorous than their older competitors and were therefore better able to survive the shrinking habitat of winter. Their continued survival after large numbers of younger voles had been added to the population probably was permitted by the expanding habitat of spring and summer. The percentage of the population surviving the winter of 1951-1952 was approximately double the percentage surviving the winter of 1950-1951. ”

“ From 1948 to 1950 and again in 1952 and 1953 I trapped in various habitat types in a mixed prairie near Hays, Kansas. Before the great drought of the thirties, Microtus ochrogaster was the most common species of small mammal in that area. Since 1948, at least, it has been taken only rarely and from a few habitats. No voles have been taken from grazed sites. In a relict area, voles were trapped in a lowland association dominated by big bluestem. ”

“ Other grasses 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 grass 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. ”

“ Females seemingly acquired the habit of entering traps more readily than did males. No correlation between any seasonally variable factor and the number of captures per individual was apparent. To a large degree, the formation of trap habits by voles was an individual peculiarity.
In order to study the extent of utilization of various habitats by Microtus, a number of areas were sampled with Museum Special snap-traps. These traps were set in linear series approximately 25 feet apart. The number of traps used varied with the size of the area sampled and ranged from 20 to 75. ”

“ The male was already an adult when first captured; consequently its life span must have exceeded 650 days. No evidence of any decrease in vigor or fertility was observed to accompany old age.
Of the 45 marked voles snap-trapped in August of 1952, 21 had been captured first as juveniles. The ages of these voles could be estimated within a few days, and the series presented a unique opportunity for studying individual and age variation. Only individuals weighing less than 18 grams when first captured were used, and their ages were estimated according to the growth rate described above. ”

“ House Field was from 1.5 to 3 times as productive as Quarry Field during the growing seasons of 1951 and 1952. Although House Field, being more productive, usually supported a larger population of voles than Quarry Field the reverse was true at the time of the clipping (Fig. 5) .
Table 1. Relationship Between Yield and Various Population Data
Although no explanation was discovered which accounted fully for the seeming aberration, two sets of observations were made that may bear on the problem. ”

“ Several instances of cotton rats eating voles, caught in the same live-trap, were noted. There is reason to believe that young voles, unable to leave the nest, are subject to predation by cotton rats. This would accentuate any competitive advantage gained otherwise by the cotton rats. ”

“ The mean percentages for the thirty month period for which data were available were: adults, 77.72 ± 4.48 per cent; subadults, 14.06 ± 3.14 per cent; and juveniles, 8.22 ± 2.62 per cent. Seasonal and yearly changes in the population structure occurred, with notable variation in the ratio of [374] breeding females to the entire population, as discussed in this report under the heading of reproduction.
Since some of the juveniles did not move enough to be readily trapped, the real percentage of juveniles in the population was probably far greater than that shown by trapping data. ”

“ Such high temperatures rarely occurred early in the day, however, so that removal of the animals from traps between eight and ten a. m. almost eliminated mortality. Those individuals captured in the night were not yet harmed, but it was already hot enough to reduce the activity of the voles and prevent further captures until late afternoon. When it was necessary to run trap lines earlier, the traps were closed in the morning and reset in late afternoon.
Reactions of small mammals to live-traps and the effects of prebaiting were described by Chitty and Kempson (1949) . ”

“ Few adults (of any species) marked before the heavy rains of July, 1951, were trapped in September, 1951, when trapping was resumed after a lapse of one month. Several subadults and some juvenal cotton rats did survive, however, [405] and provided a breeding population from which the area was repopulated. Cotton rats are born fully furred and able to move well, and are often weaned at ten days (Meyer and Meyer, 1944:123-124) . Voles, on the other hand, are born naked and helpless and are often not weaned for three weeks. ”