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Habitat Variables of the Japanese Squirrel Identified by Regression Tree Model

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Habitat Variables of the Japanese Squirrel Identified by Regression Tree Model Mammal Study 31: 1–8 (2006) © the Mammalogical Society of Japan Habitat variables of the Japanese squirrel identified by regression tree model Noriko Tamura1,*, Norio Takahashi2 and Nobuhiko Satou3 1 Tama Forest Science Garden, FFPRI, Todori 1833, Hachioji, Tokyo 193-0843, Japan 2 National Research Institute of Far Seas Fisheries, 5-7-1 Shimizu Orido, Shizuoka 424-8633, Japan 3 Ecosystem Conservation Society-Japan, Otowa Bldg. 2-30-20 Nishi-ikebukuro, Toshima-ku, Tokyo 171-0021, Japan Abstract. Habitat variables of the Japanese squirrel, Sciurus lis Temminck, were analyzed by the regression tree method based on radio-tracking data. The results suggest that the squirrels tended to use the sites with more walnut trees from July to December, and more evergreen trees in middle layer from January to June for feeding. For nest sites, they tend to use sites with more evergreen trees in the upper layer, preferably having DBH (diameter at breast height) of at least 30 cm. These results are consistent with observations in previous field studies. Key words: habitat model, habitat requirement, Japanese squirrel, tree regression. Habitat destruction is one of the most serious factors (USFWS 1980). Habitat suitability index (HSI) models causing population decrease of various kinds of wildlife. (USFWS 1981), developed to quantify the habitat quality For wise landscape management to conserve species, in the HEP, have been widely used by several agencies habitat requirements should be identified through quan- for environmental impact assessments in the United titative measurement of environmental variables. States (Morrison et al. 1998). Japanese squirrels, Sciurus lis TEMMINCK, are en- Recently, discussion has started in Japan about the demic species occurring only on the islands of Honshu, importance of habitat evaluation and application of the Shikoku, and Kyushu in Japan (Wilson and Reeder HEP (Tanaka 2001; Ecosystem Conservation Society- 1993). Recently, however, the populations in Kyushu Japan 2004). Habitat units for the species in the HEP are and western Honshu have become extinct (Kawamichi determined as the product of “habitat quality” (calculated 1997). One potential cause of local extinction of the spe- by the HSI model) and “habitat quantity” (area) (USFWS cies is rapid habitat changes in forest environments, such 1981). As the first step to make an HSI model of the as replacement of natural forest with artificial plantations Japanese squirrel, we attempted to select habitat vari- and fragmentation of natural forests by roads (Yatake ables for their life requisites. and Tamura 2001). The regression tree method is a modern statistical The tree squirrels depend on forest resources for their technique, which allows flexible data analysis without food and nest sites, and thus changes in forest environ- making any assumption about statistical distribution. ments may greatly affect their occurrence. Specifically, This feature makes this method appropriate for analyzing the Japanese squirrels selectively use natural or second- data for wildlife-habitat relationship inherently involv- ary forests but seldom use artificial plantations (Tamura ing non-linearity and high-order interactions (De’ath and 1998). Therefore, conservation of this species is a key to Fabricius 2000). Applications of regression tree meth- protecting the forest environment. ods specific to ecological data and analyses of wildlife- Habitat Evaluation Procedure (HEP) is one approach habitat relationships, are found in De’ath and Fabricius to evaluate habitat value, that was developed in the (2000), Andersen et al. (2000), and Rejwan et al. (1999). 1970’s by the United States Fish and Wildlife Service De’ath and Fabricius’s paper also provides a good over- *To whom correspondence should be addressed. E-mail: [email protected] 2 Mammal Study 31 (2006) all review of tree models. 50 m × 50 m, including the quadrat in its center. The purpose of this study is to identify the habitat Field observation showed that the squirrels use the requirements of the Japanese squirrel using a regression large evergreen trees as nest sites (Yatake and Tamura tree model based on quantitative field data. 2001), so the number of large evergreen trees (DBH > 30 cm) and the mean DBH of evergreen trees in the upper Study area layer were added to the variables. A summary of the 10 vegetation variables considered in our habitat analyses is Field studies were conducted at the experimental for- shown in Table 1. est of the Forestry and Forest Products Research Insti- One of the 50 quadrats contained planted Japanese tute’s Tama Forest Science Garden (TFSG, elevation white pines (Pinus parviflora), a favorite food source of 170–265 m, 57 ha) located in Hachioji, western Tokyo, the squirrels. Because this pine is not native to the low- Japan. The experimental forest comprises six types of elevation mountains of the Kanto district, the analysis vegetation; (1) natural forest dominated by Abies firma was conducted using data from 49 quadrats excluding and Quercus glauca, (2) secondary forest dominated by this Japanese white pine plantation. evergreen trees, such as Q. glauca, mixed with decidu- ous Q. serrata, (3) conifer plantations of Cryptomeria Radio-tracking japonica and Chamaecyparis obtusa, (4) deciduous for- Trapping was conducted 2–4 months intervals from est dominated by Q. serrata and Prunus jamasakura, (5) September 1991 to April 2001. To capture squirrels, 15 an arboretum, and (6) shrub/grassland (Tamura 1998). to 20 live-traps were set on tree branches (1–3 m in Tree species, the number of trees planted, and the year of height) and placed at 50–100 m intervals throughout the planting in each lot were recorded in the Afforestation entire trap area. Walnuts with peanut butter were used as Plan of TFSG. bait. Each trapped squirrel was weighed and the sex, maturity and reproductive status were recorded by the Methods following methods. The length of the scrotum was measured to discriminate between mature (≥ 25 mm) and Vegetation survey immature (< 25 mm) males. For females, the size and Fifty quadrats (10 m × 10 m) for the vegetation sur- shape of teats were used as criteria to estimate whether vey, were set up at random points throughout the study they had experienced reproduction. We defined an indi- area (Fig. 1). Each lot had at least one quadrat; two or vidual that was captured more than two times in different more quadrats were set in large lots containing different months as a resident individual. types of vegetation. We identified the species and mea- A collar with a radio-transmitter (50 MHz band, ZTS- sured the DBH (diameter at breast height) and the height 7D, Nakane Studio, Kamakura, Japan) was placed on 10 for all of the woody plants existing in a quadrat from males and 9 females. All radio-collared squirrels were April to June in 1996. We divided vegetation into three mature residents. Location was obtained by triangula- layers: Upper layer (DBH ≥ 23 cm, or height ≥ 10 m), tion methods using a receiver (FT-690 mk2, Yaesu) and Middle layer (3 cm ≤ DBH < 23 cm, or 3 m ≤ height < a dipole receiving antenna (Nakane Studio). The dis- 10 m), and Lower layer (DBH < 3 cm, or height < 3 m). tance between a squirrel and a receiver ranged from 20 The number of woody plants and evergreen plants in to 50 m. The location of each individual was traced from each layer were recorded. The number of species in the the time it left its nest in the morning to the time it middle and upper layers was counted because the species returned to the nest in the evening. richness may be important for squirrels as an indicator of We traced each squirrel for a total of 5 days within a food availability. In addition, the number of walnut trees period of 50 days. Data for females whose parturition was counted, because walnuts are the main food of squir- had occurred within the previous 2 weeks were omitted rels in this study site (Tamura 2004). Japanese squirrels from analysis, because they seldom left their nests. often transport walnuts to hoard and eat later and the The Japanese squirrels move from the nest sites to the mean transport distance was found to be ca. 20 m from feeding sites soon after sunrise and stay there for an hour the source tree (Tamura and Shibasaki 1996). Therefore, or more in the early morning (Nishigaki and Kawamichi the counting of walnut trees was expanded to the area 20 1996). Thus, we defined a site where the squirrel stayed m beyond each side of a quadrat; the total area was thus for eating for at least 1 hr in the early morning as a feed- Tamura et al., Habitat of Japanese squirrel 3 Fig. 1. Habitat utilization of the Japanese squirrels estimated by radio-tracking. Squares are the quadrates of vegetation study established in each lot. White squares indicate the quadrates in lots where squirrels did not use, while black squares do ones where squirrels used as feeding sites in Period 1 (a), Period 2 (b), and nest sites (c), respectively. ing site. After the early morning, squirrels occasionally sional feeding sites in the analyses, because it does not foraged in a short time during resting, grooming, nest- always follow that squirrels selected the sites for feeding. making, and other activities. We omitted such occa- In cases where after feeding at one site, a squirrel moved 4 Mammal Study 31 (2006) Table 1. The mean value of 10 vegetation variables for 49 quadrats (10 m × 10 m) at TFSG, Hachioji, Western Tokyo. Upper layer (DBH ≥ 23 cm, or height ≥ 10 m), middle layer (3 cm ≤ DBH < 23 cm, or 3 m ≤ height < 10 m), lower layer (DBH < 3 cm, or height < 3 m). Variables Mean SE Min Max NU: No.
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