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Daily Activity Patterns

Data from tracking and activity recorders revealed ground squirrels were strictly diurnal. Activity data from both sexes were combined because similar percentages of male and female squirrels were active during each hourly period (P > 0.10). Ground squirrels typically became active between 0600 and 0900 hours (approx 1-2 hr after sunrise), remained active throughout most of the day, and entered burrows and became inactive between 1900 and 2100 hours (approx 1-2 hr before sunset). Data from 100 activity recordings of 19 ground squirrels revealed that the latest any squirrel entered its burrow was 2125 and the earliest any squirrel left its nighttime burrow was 0705 hours. Monthly variation in daily activity patterns was slight and associated with changing length of day.

Daily Movements

Daily movements of individual ground squirrels began when they emerged from their nighttime burrows and ended when they retreated into burrows for the night. The mean daily total distance traveled by males was greater than the mean daily total distance traveled by females (P < 0.05) (Table 1). Biweekly differences were significant during one 2-week period in 1981 and 3 2-week periods in 1982 (P < 0.05). Females traveled least during gestation (late May-early Jun) and during the 2-week period just before immerging for the winter (Aug). Distances traveled daily by males decreased during the 2-week period before immergence.

Table 1.  Mean daily distance traveled (m) by adult Franklin's ground squirrels during biweekly sample periods on a study area in North Dakota, 1981-82.
Yr     Period	M			F
	na		SD	na		SD
1981
10-23 May	13	219	138	7	140	78
24 May-6 Jun	17	139b	77	16	82b	34
7-20 Jun	15	223	104	16	167	49
21 Jun-4 Jul	16	213	77	15	205	76
5-18 Jul	16	210	135	12	214	88
19 Jul-1 Aug	11	152	37	18	148	76
2-13 Aug	8	134	57	15	152	53
biweekly	96	188		99	157
1982
25 Apr-8 May	9	282	147
9-22 May	23	263b	123	13	110b	56
23 May-5 Jun	34	234b	126	23	96b	38
6-19 Jun	26	205b	142	24	100b	52
20 Jun-3 Jul	18	197	139	17	169	64
4-17 Jul	26	259	155	27	197	86
18-31 Jul	33	241	132	45	184	92
1-13 Aug	19	137	72	24	149	55
biweekly	188	227		173	150
annual	284	213		272	153
a No. daily records. b Significantly different means between M and F (P < 0.05).

Home Ranges

Biweekly home-range sizes did not differ between years (P = 0.2418). Biweekly home-range sizes of 21 males averaged 7.2 ha (range = 0.5-38.4 ha) and were greater than those of 20 females that averaged 2.7 ha (range = 0.3-14.1 ha) (P = 0.001) (Table 2). The largest mean biweekly home-range sizes of males in 1981 were in early May during the breeding season, and in 1982 in late July. Biweekly home ranges for males were smallest before the males entered hibernation in 1981 and 1982. During both years, mean biweekly home ranges of females remained relatively small throughout gestation and lactation, then gradually increased during the post-lactation period, and decreased shortly before immergence. Mean activity radii of males during biweekly intervals ( = 1,106 m) were greater than those of females ( = 67 m) (P = 0.0001). Shifts in geometric centers of activity over biweekly intervals were also greater for males ( = 97 ± 103 [SE] m, n = 91) than for females ( = 59 ± 5.4 m, n = 86) (P = 0.0015).

Annual home-range size did not differ between years (P = 0.977). Annual home ranges of 21 males averaged 24.6 ± 11.5 ha and were greater than those of 20 females that averaged 8.7 ha (range = 2.5-22.7 ha) (P = 0.0001). Annual home ranges overlapped within and between sexes. The annual mean activity radius of males ( = 147.9 ± 69.4 m) was greater than that of females ( = 87.7 ± 35.4 m) (P = 0.0014).

Table 2.  Mean biweekly and annual home-range sizes (ha) of adult Franklin's ground squirrels on a study area in North Dakota 1981-82.
Yr     Period	M			F
	n		SD	n		SD
1981
27 Apr-9 May	6	9.7	2.7
10-23 May	8	8.4	2.9	8	2.9	1.6
24 May-6 Jun	8	4.7	2.8	9	1.3	0.7
7-20 Jun	6	8.4	4.9	8	3.1	1.8
21 Jun-4 Jul	8	6.0	2.8	7	3.9	0.9
5-18 Jul	7	6.0	3.2	7	3.6	1.7
19 Jul-1 Aug	6	4.2	3.4	9	4.0	4.0
2-13 Aug	6	3.6	2.2	6	2.2	1.3
biweekly	55	6.4		56	3.0
annual	10	19.3		11	9.4
1982
25 Apr-8 May	8	6.1	3.0
9-22 May	8	8.4	6.1	4	0.8	0.7
23 May-5 Jun	8	5.5	5.5	6	1.7	1.4
6-19 Jun	8	5.3	4.4	8	1.1	1.0
20 Jun-3 Jul	8	7.0	9.0	8	2.7	1.1
4-17 Jul	10	11.6	3.6	9	3.6	2.5
18-31 Jul	8	14.8	8.1	9	4.0	3.0
1-13 Aug	6	3.6	2.7	8	2.2	1.3
biweekly	64	8.4		52	2.5
annual	11	29.4		9	7.9

Habitat Use

The distribution of 4,430 telemetry locations of active ground squirrels during both years combined showed unequal use of habitats on the WPA (Fig. 1). The most intensively used portion of the WPA comprised the northern portion of TB and adjacent portions of UN19 and DC6. Two-thirds of the radio-marked ground squirrels were located in TB at least once. The least used portions of the WPA upland were DC5, the center of DC8, and the southeastern part of DC6.

When inactive at night, ground squirrels were underground in what appeared to be abandoned burrow systems of northern pocket gophers (Thomomys talpoides). Because burrows were also used for refuge during the day, above ground movements of the ground squirrels were influenced by the location of suitable burrows. The spatial distribution of burrows used at night by habitat block was similar to the distribution of locations of squirrels during the day (Table 3).

Electivity indexes calculated for the 8 habitats using only active locations showed that TB and UN were used greater than expected and that DC3, NP, DC8, and DC5 received less use than expected (Table 3). During each year, analysis of visual obstruction and litter depth measurements on the 7 habitats dominated by herbaceous cover (TB excluded) indicated some difference among habitats during each of the 3 sampling periods (ANOVA, P < 0.001), but when least square means were examined, no pattern to the differences among sampling periods or habitats was evident.

All habitats differed in age of the stand and frequency of cultivation after the initial seeding of the WPA in 1963. The 2 habitats that received greater use than expected were characterized by relatively long periods of no disturbance to vegetation or soil. Although the vegetation and soil on NP were undisturbed longer than in any other habitats, that area was often surrounded by water that deterred use by ground squirrels and accounted for its relatively low electivity index. The 3 habitats that received less use by ground squirrels than expected had relatively recent disturbances to the soil and vegetation. Habitat block DC3 and DC5 were farmed for ≥2 years between 1974 and 1979. Habitat block DC8 was farmed continuously from 1966 to 1975. Although DC6 received less use by ground squirrels than expected, avoidance was not as great as anticipated based on cultivation history. However, most use of that block occurred adjacent to heavily used TB and an adjacent sunflower stubble field often visited by squirrels in spring 1981.

The electric fence probably had little effect on the low relative use of DC8 because the portion of UN19 enclosed by the fence received heavy use similar to that portion of UN19 outside the fence (Fig. 1). A previous study (Lokemoen et al. 1982) and telemetry data and observations obtained during our study showed that ground squirrels moved freely through these fences.

Ground squirrels used only the fringes of the cropland adjoining the WPA (Fig. 1). The most intensively used cropland was located immediately northeast of TB. A narrow band of trees extending north of TB provided the ground squirrels with a source of protective cover at the field edge. Nineteen of the radio-equipped ground squirrels used that area at least once. Most use occurred in April 1981 when waste sunflower seeds from the previous year's crop were available, and during late summer 1982 when wheat planted in the field was ripening. Use of other fields adjoining the WPA and containing the same crop was limited to < 1 % of the locations. These fields were not close to shrub cover on the WPA. Seven of 49 ground squirrels used the uninhabited farmstead adjacent to the WPA.

Mowing and burning of portions of the WPA affected ground squirrel movements. Habitat block DC5 was mowed the summer preceding this study. Ground squirrels avoided habitat block DC5 throughout the study. In contrast, DC3 was mowed in late summer 1981 and burned the following spring. Ground squirrels used habitat block DC3 extensively before it was mowed and burned but used it infrequently until the vegetation regrew to a height of about 30 cm in mid-May.