Habitat Use and Selection of the Siberian Weasel Mustela Sibirica Coreana During the Non-Mating Season

J. Mamm. Soc. Japan 19 (1) : 21-32 June 1994

Habitat Use and Selection of the Siberian Weasel Mustela sibirica coreana during the Non-mating Season

Hiroshi SASAKI* and Yuiti ONO** Laboratory of Ecology, Department of Biology, Faculty of Science, Kyushu University, 33, Fukuoka 812, Japan *Present address : Chikushi Jogakuen Junior College, Ishizaka, Daznifu, Fukuoka 818-01, Japan * *Emeritus professor (Accepted 10 December 1993)

Abstract. Habitat use and selection of the Siberian weasel Mustela sibirica during the non-mating season were studied, on the small island of Aoshima, Japan, from 1982 to 1989. Weasels occurred throughout the island. The village area was compared with the northern grassland which was less affected by human activity. Weasels in the village area were divided into two groups ; the "border weasels", which had their ranges around the periphery of the village, and the "central weasels", which occupied the center of the village. The population density was lower in the grassland area than in the village. The body weight of border weasels was the largest on Aoshima. Border weasels rested in barns and houses and were active in grasslands and on roads, while weasels in the grassland rested and were active mainly in grasslands. Weasels frequently used piles of hay in barns as resting sites in the village. Resting sites, offering good thermal conditions, are thought to be important factors of the range of the Siberian weasel, especially for females, during winter. The village periphery was assumed to be the most favorable weasel habitat because the village area provided good resting sites and a wide range of food.

Key words : Siberian weasel ; Habitat use ; Home range ; Resting site

The spacing pattern and social organization of mustelids have been discussed by several authors (Lockie 1966, King 1975, Powell 1979, Erlinge 1977a, Birks and Linn 1982). Least weasels Mustela nivalis and American mink M. vison are thought to have intrasexual territories (King 1975, Gerell 1970, Birks and Linn 1982). Erlinge and Sandell (1986) reported that male stoats M. erminea have intrasexual territories during the non-breeding season, and are non-territorial during the mating season. These studies focused on the distribution of food and of receptive females in relation to habitat selection. Erlinge (1983) showed that M. erminea populations are limited by food shortages, whereas Weber (1989) revealed the importance of resting sites for polecats M. putorius during winter. The high mortality of various species of the mustelidae during winter may be as a result of a combined shortage of food and of good resting sites. 22 Sasaki and Ono The Siberian weasel M. sibirica is widely distributed in east Asia, with M. s. coreana the subspecies endemic to the Korean Peninsula and to Tsushima, Japan. It invaded, and spread throughout the western part of Japan, during the 1900s (Milturiya 1969), and has also invaded urban areas. This 'urban' habit is not confined to Japan. Sheng and Lu (1982) showed that in China, female Siberian weasels concentrated in village areas during the breeding season. In our study area, Aoshima, off northwest Kyushu, we compared the village and semi-natural areas which offer different types of resting sites for Siberian weasels. We discuss the factors affecting habitat selection and range size during winter, that is during the non-mating season.

Study Area and Methods

The study was carried out on Aoshima (33" 25' N, 129" 40' E), an island some 700 m off in the northwest Kyushu (Fig. la). This island is 95 ha in area and its highest point is merely 58111 above sea level. At the center of the island, in area No. 4 (see Fig I.), there is a fishing village of about 500 people. Various habitats are represented on Aoshima : housing area, cultivated fields, paddy fields, grasslands (including abandoned cultivated and paddy fields) and secondary forests.

Fig. 1. The study area. (a) Areas selected for the distribution survey. (b) Two study areas used for the radio-telemetry survey. I : the northern grassland, I1 : the village area. The dotted area shows the residential area. Habitat Use of Siberian Weasel 23 The distribution and density of the Siberian weasel was surveyed across the whole area of Aoshima. Two study areas were chosen for radio- telemetry : the northern grassland, and the village area, where houses were concentrated (Fig. lb). Outside the village the habitat was divided into grassland 43.5%, cultivated fields 21.6%, secondary forest 21.4% and paddy fields 10.0% for the grassland. The grasslands consist predominantly of Miscanthus sinensis. The island's secondary forests are characterized by Pittosporum tobira, Litsea japon- ica and Rhaphiolepis uvzbellata, and are often covered by Pueraria lobata. The undergrowth of the secondary forest is sparse, but is dominated by Tra- chelospermum jasminoides. The cultivated and paddy fields are mostly sur- rounded by stone walls of ca. 1 m high. Most of the village area is residential, consisting of houses, barns, school buildings, shrines and a harbor.

Trapping Wooden box live-traps were set at a density of four per hectare in the evenings and closed in the mornings during the distribution survey. Traps were checked at midnight to reduce risks of injury to the weasels during the radio-telemetry survey. At each trap site, one trap was placed either on a path or along a stone wall. Dough, spread with mayonnaise, was used as a bait during the distribution survey. During the radio-telemetry survey, the same bait was used in the grassland, but chicken meat with bones was used in the village area, to improve trappability. Captured weasels were anesthetized by an intramuscular injection of Ketamin hydrochloride, Xylazine hydrochloride, and Atropin sulphate. Then they were sexed, weighed, and marked by toe clipping (during the distribution survey), or by freeze-branding and tattooing on the back (during the radio- telemetry survey). Weasels were released after recovery at their capture sites. Judging from the condition of the testis, the non-mating season starts in October and ends in February on Aoshima.

Estimation of Population Density Aoshima was divided into six areas (Fig. la), which were trapped in turn from June to September 1982 during the distribution survey. Trappings was also conducted at the beginning of each research period during the radio- telemetry survey. Trapping took place over four or five nights, and the weasel densities were estimated using the removal method (Hayne 1949). The exposed rates were calculated from the number of captured weasels divided by the estimated population number.

Radio - telemetry The areas least affected by human activity were the northern and southern areas, but the density of weasels in the southern area was not high enough for telemetric study, thus radio-telemetry was carried out only in the northern 24 Sasaki and Ono grassland (from November 1982 to December 1985), and in the village area (from January 1988 to February 1989). Weasels in the village area were further divided into two groups : those which were located frequently enough to be able to estimate their range sizes, and whose whole range were included in the residential area ("central weasels"), and those which had part of their range in the residential area ("border weasels"). Male weasels were fitted with radio-collars weighing about 20 g, females with collars of about 10 g. The transmitters' frequencies ranged from 50 to 54 MHz. Tracking took place over either for two or three weeks. Weasels were tracked in turn and thus tracking time was broken up over each day. Track- ing times per day varied depending on the number of weasels being tracked. The locations of weasels were determined by triangulation using portable receivers and adcock antennae, throughout the study, however, from 1982 to 1984 three- or five-element Yagi antennae were also used. Location points identified using fixed antennae (the three- or five-element Yagi antennae) were used only for estimating range sizes. Range sizes were calculated using the minimum convex polygon method. Weasels which provided insufficient location points were excluded from the analysis of home range size and habitat use. The weasels which were used for the calculation of range size are listed in Appendix A. Weasel movements, which were observed during tracking, were also recor- ded and used for estimating range sizes. Only the first point of observation on each occasion, and only records one hour or more apart, were used for the analysis of habitat use.

Food Habits Feces of Siberian weasels were collected from 1983 to 1985 from the grassland area. Feces were washed over a 1 mm mesh sieve and the remains were identified by stereoscopic microscope.

Results

Population Density The total number of weasels captured was 38 (29 males and 9 females), and the exposed rate was 92%, excluding area No.6, during the distribution survey. The density in the village, area No.4, was the highest on the island. Newborn males were excluded from the calculation, and we were unable to estimate the density in area No. 6 because trappability was very low there. The population density of the Siberian weasel in each area on Aoshima is shown in Table la. Weasel densities were lower in the grassland than in the village (Mann- Whitney U-test, U=O, p <0.05, see Table lb). The average exposed rates were 95.4% in the grassland, and 89.9% in the village area, respectively.

Body Weight Males from the village area were significantly heavier than those from the Habitat Use of Sibe~ianWeasel 2 5

Table 1. Densities of Siberian weasels on Aoshima as estimated by the removal methods (Hayne 1948). (a) Distribution survey in 1982. Males which were less than 470 g were treated as "newborn", as the smallest body weight of reproductive males during the mating season was 470 g ; newborn males were excluded from the calculations. (b) Densities during the non-mating seasons from 1982 to 1989. (G : the grassland, V : the village area). (a) Research No. of weasels captured Estimated Area size Density Y Area period male female number (ha) (No./ha) 1 23-30 Jun. 3 0 2.9 11.43 0.25 -0.913 2 24-30 Jun. 4 3 9.3 12.19 0.76 -0.745 3 17-23 Jul. 3 1 3.6 11.16 0.32 -0.871 4 15-20 Sep. 14 4 19.9 24.26 0.82 -0.968 5 13-21 Aug. 5 1 5.4 12.70 0.43 -0.918 6 23-29 Aua. 2 0 - 22.46 - -

No. of weasels captured Estimated Area size Density Area Research period male female number (ha) (No./ha) G 27 Nov. -10 Dec. 1982 4 2 6.0 23.17 0.26 -1.000 G 4-10 Oct. 1983 4 2 6.1 23.87 0.25 -0.780 G 24 Nov. -6 Dec. 1983 4 2 6.1 26.09 0.23 -0.735 G 12 Oct. -25 Oct. 1984 7 1 9.3 26.09 0.36 -0.877 G 16 Nov. -2 Dec. 1985 3 5 8.5 24.93 0.34 -0.964 V 30 Jan. -11 Mar. 1988 11 8 26.4 20.00 1.32 -0.742 V 28 Dec. 1988-13 Feb. 1989 15 7 20.4 20.75 0.98 -0.606 grasslands (U-test, p<0.05, Table Za), however, there was no significant difference between the weights of the females from the same two areas. Both male and female border weasels were heavier (though not significantly so) than central weasels, though border weasels were significantly heavier than those in the grassland areas (p< 0.05 for male ; p <0.05 for female, Tables Za, b).

Table 2. Body weights of Siberian weasels during the non-mating season on Aoshima. Data were collected from the grassland from 1982 to 1985, and from the village area from 1988 to 1989. Differences were tested using Mann-Whittney U-test. (average body weightkSD, g) (4 Grassland Village area Male 670+89(n=22) 750*150(~=28) z=-2.218 p<0.05 Female 329+53(n=10) 346f52(n=15) U=59 N.S. z= -4.472 z= -5.351 p<0.001 p<0.001

Central Border Male 789f 69(n=7) 831*146(n= 12) U=29 N.S. Female 327f 70(n=3) 382f 31 (n=7 ) U=6 N.S. U=O U=O p<0.05 p<0.05 26 Sasaki and Ono The body weight of males was about twice that of females (p < 0.001 in the village area, p < 0.001 in the grassland, p < 0.05 for central weasels and p < 0.05 for border weasels, see Tables Za, b).

Home Range Size The range size of males was larger in the grassland than in the village area (p < 0.05, Table 3a). And in the village area, the range size of border males was larger than that of central ones (p <0.05, Table 3b). There was no significant difference in the range size of females between areas, nor between central and border females (Tables 3a, b). Central weasels occupied significantly smaller ranges than did weasels in the grassland (p<0.05 for males; p<0.05 for females). The differences in range size between the sexes were not significant in either area, nor in central or border weasels. The range size of central males was not correlated to their body weight, unlike that of border males which was correlated (p<0.05, Fig. 2). In the grassland, there was no correlation between male range size and body weight. But if the very lightest weasel was excluded from the calculation, range size was strongly correlated to body weight (p < 0.05, Fig. 2). There was no correlation between female range size and body weight.

Table 3. Range sizes of Siberian weasels during the non-mating season on Aoshima. Data were collected from the grassland from 1982 to 1985, and from the village area from 1988 to 1989. Differences were tested using Mann-Whittney U-test. (average range size+SD, ha) (a) Grassland Village area Male 4.37+3.87(n=8) 1.42f 1.62(n=19) U=29 p<0.05 Female 1.67f 0.91(n=4) 1.31+1.12(n=10) U=15 N. S. U=7 U=85 N. S. N. S.

Central Border Male 0.23+0.12(n=7) 2.11f 1.69(n=12) U=4 p<0.05 Female 0.33+0.13(n=3) 1.73f 1.10(n=7 ) U=2 N. S. U=5 U = 39 N. S. N. S.

Habitat Use Weasel activity was recognized, from the stability of received radio pulses when located, and classified either as active or resting. The percentages of location points in each habitat are shown in Table 4. Outside the village area weasels rested and were active mainly in grasslands. There were only three barns, and no houses, in the grassland area, but female weasels used the barns frequently. Border weasels rested mainly in Habitat Use of Sibelian Weasel

BODY WEIGHT (g) Fig. 2. The body weights and range sizes of male weasels during the non-mating season on Aoshima. Circles : grassland weasels, triangles : border weasels, squares : central weasels. The regression lines are : Y = 0.037 X - 22.147, r = 0.807, n = 7, 0 < 0.05, for grassland males when the lightest weasel ( ) was excluded from calculation ; Y = 0.007X - 3.476, r = 0.583, n = 12, p < 0.05, for border males ; Y = 0.0004 X -0.118, Y = 0.262, n= 7, N. S., for central males. Data were collected from the grassland from 1982 to 1985, and from the village area from 1988 to 1989.

Table 4. The share (%) of the location points in the habitat component during the non- mating season on Aoshima. Data were collected from the grassland from 1982 to 1985, and from the village area from 1988 to 1989. Piles of goods include only those outside of barns and houses. G : the grassland. Grass- Forest Cultivated Bar11 Pile of Ship Stone Road No. of No. of Grave Shore Underground land field house goods wall hole inds. plots Resting state G male G female Border male Border female Central male Central female Active state G male G famalte Border male Border fe~nale Central male Central fe~llale 28 Sasaki and Ono barns and houses, but were active both in grasslands and along roads in the residential area. Central weasels rested and were active mainly in barns and houses. The most important resting sites for weasels in the village area were barns. Of the weasels which rested in either houses or barns, most rested in barns (95.6% of border males, 75.0% of border females, 75.5% of central males and 58.5% of central females). Weasels used piles of hay as resting sites in many barns. Direct observation, during tracking, could be made more frequently in the village area than in the grassland, as weasels were conspicuous when they used roads for movement.

Table 5. Distribution of location points of border weasels in relation to habitat types during the non-mating season on Aoshima. Data were collected in the village area from 1988 to 1989. The expected number of location points was calculated from the average ratio of the sizes of housing, and the other areas, to the home range size. "Housing area" includes houses, barns, ports, and roads in residential areas. Housing area Other area x P Broder male Active obs. exp . Resting obs. exp . Broder female Active obs. exp . Resting obs. exp .

Table 6. Frequency occurrence of foods of the Siberian weasel in the grassland of Aoshima from 1983 to 1985. The quantity of each food item was scored : 1 ; less than 25%, 2 ; 25-50%, 3 ; 50-75%, 4 ; 75-100%. Each figure in parentheses shows the average score of each food item when it occurs in feces. Winter : Jan. - Mar., Spring : Apr. - Jun., Summer : Jul. - Sep., Autumn : Oct. - Dec. Winter Spring Summer Autumn Total Insects 40.0(1.0) 39.1(1.0) 55.6(1.9) 87.5(2.3) 68.7(2-1) Larvae 0 8.7(1.0) 0 311.0 3.50.0) Centipedes lO.O(l.0) 8.7(2.0) 0 3.1(2.0) 4.4(1.8) Fish 60.0(1.8) 21.7(2.1) 0 4.7(1.7) lZ.Z(l.9) Frogs 0 30.4(2.1) 0 12.5(1.5) 13.0(1.8) Reptiles 0 13.0(2.0) 38.9(2.1) 4.7(1.0) 9.6(1.7) Birds lO.O(l.0) 4.4(3.0) 0 0 1.7(2.0) Rodents eo.o(l.7) 13.0(3.0) 16.7(1.3) 23.4(1.7) 23.5(1.8) Unidentified vertebrates 0 8.7(1.0) 0 0 1.7(1.0) Pieces of meat 20.0(2.0) 0 33.3(1.0) 32.8(1.1) 25.2(1.2) Fruits 0 13.0(2.3) 22.2(2.0) 1.60 7.0(2.0) Man-made 20.0(1.0) 13.0(2.7) 5.6(1.0) 1.6(1.0) 0-l(1-7) No. of feces 10 23 18 64 115 Habitat Use of Siberian Weasel 29 The average percentages of housing areas in the home ranges of border weasels were 27.9% for males, and 30.7% for females. Both active and resting border males favored the housing area (x2-test,p < 0.01 for active males, p < 0. 001 for resting males). Active border females were impartial in their use of home ranges, but resting border females were concentrated in the housing area (p<0.001). The bias of home range use during active and resting behavior is shown in Table 5.

Food Habits The food habits of weasels were revealed by the frequency occurrences of food items in the feces (see Table 6). Their main foods preyed were insects and rodents in the grassland area. It was difficult to find feces for analysis from the village area, but weasels were observed eating abandoned fish around the harbor and from ships, and sugar cakes, bread, mayonnaise, and many kinds of cooked food from houses.

Discussion

Home Range Size Home range size is assumed to be related to body weight, and thus food resources may be the primary determinant of range size (McNab 1963, Hares- tad and Bunnell 1979, Gittleman and Harvey 1982). Especially during the non- mating season range size might strongly reflect the availability of food resources. Erlinge (197713) reported the social rank of male stoats to be positively correlated to body weight and age. In our study area range size was certainly correlated to body weight for border males, and also for grassland male if the lightest individual was excluded. Siberian weasels are assumed to have intrasexual territories during the non-mating season, though overlap of home ranges sometimes occurs as a result of the dispersion of young weasels of the year on Aoshima (Sasaki unpubl. data). The lightest Siberian weasels may be young and thus not yet sedentary. Central males may hold ranges of a minimum viable size and try to stay in the village area where the females are concentrated. The fact that the central weasels had the smallest range sizes suggested that food was abundant in and around barns and houses. Good resting sites, for example, piles of hay in barns, were also concentrated in the village area. But, no significant difference in female range size between areas was observed, and the range size of females was independent of their body weight. This suggests that neither food nor resting sites may regulate female range size. 30 Snsaki and Ono Habitat Selection Border weasels were the largest on Aoshima and held ranges in the area fringing the village. They mainly used barns and houses as resting sites. Sheng and Lu (1982) found that the density of female Siberian weasel nests was high in village areas with abundant water, and 85% of nests during the breeding season were found in piles of hay. In our study area, weasels also frequently used piles of hay in barns as resting sites, and these were distributed only in the village. Piles of hay would be beneficial to weasels not only as shelter for breeding, but also to minimize heat loss in winter, and small mustelids are known to be prone to heat loss. The metabolism of cold stressed longtailed weasels M. frenata is, for example, 50-100% greater than that of normal shaped mammals of the same body weight, because they have a greater relative surface area, shorter fur, and are unable to adopt a spherical resting posture (Brown and Lasiewski 1972). The basal metabolic rate of mustelids with body weights below 1 kg has been shown to be higher than that of heavier species (Iversen 1972). The Siberian weasel is thought to need good resting sites in order to reduce heat loss, and this is particularly true for females which are only half the weight of males (Table 2). Predation is also an important factor in the selection of resting sites. The main predators of weasels are assumed to be cats on Aoshima, and they are abundant in the residential area ; nevertheless weasels frequently used barns and houses as resting sites. Food for weasels is assumed to be abundant both in and around barns and houses, yet border weasels were found to be active mainly in grasslands and along road sites in the residential area. Insects, which were abundant in grasslands, were one of the weasels most important foods in the grassland in winter (Table 6). A home range including both housing area and grassland is thought to offer a wider range, and more abundant food. Disturbance by humans may also affect weasel activity. Sandell (1989) assumed that whereas food availability determines the distribution of females, the spacing of males is determined by the distribution of females, at least during the mating season, in solitary carnivores. But, Weber (1989) reported that the distribution of polecats in Switzerland may be affected by the availability of suitable winter resting sites, and that they change these sites seasonally. Birlts and Linn (1982) also reported the importance of resting sites for the habitat selection of American mink. In our study, the most favored habitat proved to be the periphery of the village, because the density of weasels was higher (than in the grassland), and they were heaviest. Home ranges around the periphery of the village offered good resting sites and high food availability in the village and additional foraging areas outside the village. During the non-mating season Siberian weasels are assumed to select their ranges not merely based on food abundance, but also on the supply of resting sites. Habitat Use of Siberian Weasel

Acknowledgments

The authors express gratitude to the members of the Department of Biology, Faculty of Science, Kyushu University, for their assistance. Special thanks are due to the people of Aoshima for their kind help during the field work. We thank Dr. Alan Newsome for his critical comments on the manuscript and also two anonymous referees for their constructive comments. The authors wish to express special thanks to Dr. T. Uchida and Dr. S. Shiraishi to introduce the study area. Dr. M. Brazil kindly improved the English of the manuscript. This study was financially supported in part by a Grant-in-Aid for Special Project Research on the Biological Aspects of Optimal Strategy and Social structure from the Japanese Ministry of Education, Science and Culture (No. 61133003).

References

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Appendix A Weasels for calculation of range sizes Area Animals Sex Tracking period No. of location points Grassland area M4 Male 27 Nov.-10 Dec. 1982 M14 Male 27 Nov.-10 Dec. 1982 M65 Male 27 Nov.-6 Dec. 1983 M62 Male 13-31 Oct. 1984 M66 Male 12-30 Oct. 1984 MT1 Male 12-31 Oct. 1984 MT3 Male 13-20 Oct. 1984 MT4 Male 14-25 Oct. 1984 F11 Female 27 Nov.-10 Dec. 1982 FT2 Female 12-23 Oct. 1984 FT4 Female 16 Nov.-3 Dec. 1985 FT5 Female 16 Nov.-3 Dec. 1985

Village area MT23 Male 30 Jan.-11 Mar. 1988 (Border weasels) MT36 Male 30 Jan.-10 Mar. 1988 MT39 Male 10 Feb.-11 Mar. 1988 MT26 Male 13 Jan.-5 Feb. 1989 MT30 Male 13 Jan.-6 Feb. 1989 MT41 Male 13 Jan.-13 Feb. 1989 MT43 Male 28 Dec. 1988-5 Feb. 1989 MT45 Male 28 Dec. 1988-2 Feb. 1989 MT47 Male 28 Dec. 1988-1 Feb. 1989 MT48 Male 6 Jan.-$ Feb. 1989 MT49 Male 13 Jan.-12 Feb. 1989 FT14 Female 10 Feb.-3 Mar. 1988 FT21 Female 3 Feb.-5 Mar. 1988 FT31 Female 12 Feb.-3 Mar. 1988 FT33 Female 28 Dec. 1988-12 Feb. 1989 FT34 Female 28 Dec. 1988-27 Jan. 1989 FT35 Female 28 Dec. 1988-10 Jan. 1989 FT36 Female 28 Dec. 1988-26 Jan. 1989

Village area MT24 Male 30 Jan.-11 Mar. 1988 (Central weasels) MT30 Male 7 Feb.-11 Mar. 1988 MT32 Male 1 Feb.-10 Mar. 1988 MT35 Male 30 Jan.-10 Mar. 1988 MT37 Male 1 Feb.-10 Mar. 1988 MT35 Male 9 Jan.-6 Feb. 1989 MT46 Male 28 Dec. 1988-5 Feb. 1989 FT26 Female 5 Feb.-3 Mar. 1988 FT30 Female 7 Feb.-5 Mar. 1988 FT37 Female 28 Dec. 1988-4 Feb. 1989