Mammal Study 32: 105–114 (2007) © the Mammalogical Society of

Relationship between grooming and tick threat in sika deer Cervus nippon in habitats with different feeding conditions and tick densities

Miki Yamada1,* and Misako Urabe2,** 1 Graduate School of Human Culture, Nara Women’s University 2 Department of Ecosystem Studies, School of Environmental Science, The University of Shiga Prefecture, Shiga 522-8533, Japan

Abstract. Using two populations of sika deer Cervus nippon, the relationship between tick density within habitats and the duration of both self-grooming and allogrooming was investigated. At Mageshima Island, where the deer spend much of their time feeding and tick density is high, the time spent self-oral grooming frequency fluctuated with tick density within the deer’s habitat. However, in Nara Park, where the deer are fed artificially and tick density is low, neither oral nor self-scratch grooming correlate with tick density. Although tick density is much higher at Mageshima than at Nara Park, the overall duration of self-grooming did not differ between these sites, suggesting that this was affected by the time budget required for other behaviors, especially foraging. Allogrooming duration did not correlate with tick density at either study sites, suggesting that the most important function of this behavior is a social one, rather than the removal of ectoparasites.

Key words: allogrooming, self-grooming, sika deer, ticks, time budget.

The removal of ectoparasites, such as ticks, is an impor- required for its survival, such as foraging. Thus, the tant function of grooming behavior in ungulates (Hart frequency of grooming might depend upon cost/benefit 1990). In an experimental study with impala, Aepyceros ratio. As animals cannot simultaneously undertake both melampus, animals fitted with harnesses that restrained foraging and grooming, the cost/benefit ratio will vary self-oral grooming had 20 times more ticks than control with food availability, as animals require more foraging impala (Mooring et al. 1996a). The cost of tick infesta- time during the periods of famine (Mooring 1995). tion has been well demonstrated in domestic animals. It Mooring (1995) showed that grooming by female impala causes weight loss in calves (Little 1963) and mediates did not exhibit a negative correlation with feeding time infectious disease such as piroplasmosis (Yoshida 1980). whereas, for males, feeding time was negatively corre- Therefore, animals should groom more frequently when lated with self-oral grooming. This suggests that males ectoparasites are more prevalent within a habitat. This are limited by foraging compared with females and that has been experimentally demonstrated in impalas (Hart male grooming frequency is to some extent influenced et al. 1992) and the North American cervids, Alces by their foraging time budget. Furthermore, Mooring alces, Cervus elaphus, Odocoileus hemionus and O. and Hart (1995b) showed that territorial males of impala virginianus (Samuel 1991; Welch et al. 1991). groomed less than females carried a heavier tick load, Grooming behavior also involves costs. One of these suggesting that they sacrificed grooming time for ter- is the physical loss of material from the grooming organ ritorial behavior. However, an alternative explanation (e.g., saliva or teeth) (Ritter and Epstein 1974; McKenzie has been suggested: that testosterone may depress the 1990), while others include behavioral costs, such as immune system of males (Mooring and Hart 1995b). decreased vigilance against predators (Mooring and Hart Grooming itself is considered to be a programmed 1995a). Furthermore, grooming may reduce the time an behavior, because it does not become zero even in ecto- animal is able to devote to other behaviors that are parasite-free conditions (Mooring 1995; Mooring et al.

*Present address: Senbon-Izumi-sagaru, Juyonken-machi 394-1-708, Kamigyo-ku, Kyoto 602-8164, Japan **To whom correspondence should be addressed. E-mail: [email protected] 106 Mammal Study 32 (2007)

2004). Mooring et al. (2004) showed the evolutionary trend of ungulate grooming behavior in parasite-free conditions in relation to body plan and habitat type, They concluded that both body size and habitat influence grooming rates and the former explains more of the vari- ation in grooming. However, habitat also influences grooming rate significantly, and species found in more tick-dense, more closed habitat tended to groom more. They also showed that allogrooming delivered to other adult individuals is concentrated in lineages inhabiting closed forest habitats, especially in cervids. Grooming for another individuals (allogrooming) has also the function of ectoparasite removal (Mooring and Hart 1995b; Mooring et al. 1996a, b), but it has usually been interpreted in the social context. Allogrooming as a social behavior has been studied in many groups of mammals, especially in primates and ungulates. It is interpreted as an altruistic behavior for kin (e.g.

Muroyama 1991), affiliative relationship between indi- Fig. 1. Location of Mageshima Island and Nara Park. viduals (e.g. Tsukahara 1990; Muroyama 1994), or ten- sion reduction (Terry 1970; Feh and De Maziéres 1993; Das et al. 1997), although these explains are not exclu- tors. The island is covered with temperate and subtropi- sive each other. cal vegetations, which is heavily grazed by the deer. The In this study, the relationship between the seasonal dominant plant species of grassland on the southern part threat of tick infestation and grooming frequency was of the island are Imperata cylindrica and Pennisetum investigated in sika deer Cervus nippon at Mageshima alopecuroides, whereas forest on the eastern side is dom- Island and Nara Park. These sites have contrastive con- inated by Pinus thunbergii and Casuarina glauca (Sasaki ditions in tick density, and also in food availability for et al. 1960). deer, which may affect the time spent to grooming. Dif- The subspecies of deer on Mageshima, Cervus ferences in the patterns of correlation between grooming nippon mageshimae, is a little smaller in body size than frequency and tick density between sites are discussed in C. n. centralis of Nara Park. Body weight of C. n. relation to general behavioral time budgets. mageshimae is intermediate between those of the C. n. centralis (50–70 kg) and C. n. yakushimae (30–45 kg). Materials and methods It has become lighter in recent years because the nutri- tional condition of the deer has deteriorated following a Study sites population increase (Tatsuzawa 2000). The study was conducted in Mageshima Island, The study periods at Mageshima were 8–15 April , western Japan, and in Nara Park, 1998, 24 September–12 October 1998, 29 April–6 May Nara Prefecture, central Japan (Fig. 1). The mean annual 1999, 11–19 August in 1999 and 25 September–1 Octo- temperature is 21.1°C at Island (10 km east ber 1999. of Mageshima) and 16.0°C at Nara in 1998, and the Nara Park (Fig. 1) had a population of 1215 deer (737 annual precipitation is 2769 mm at Tanegashima Island females, 269 males and 209 fawns) in a 3 km2 area (more and 1693 mm at Nara in 1998 (data by Japan Meteoro- than 4 individuals/ha), according to the 20 July 1999 logical Agency). census of the Nara Deer Foundation. The park is not Mageshima Island is uninhabited, although people fenced, and deer generally can move freely although they often land to fish, sometimes with cars. About 500 wild are partly managed by the Nara Deer Foundation. Preg- sika deer inhabit a 9 km2 area (more than 5 individuals/ nant females are caught from April to July, and are ha: Tatsuzawa 2005). No large mammals other than the allowed to bear fawns in an enclosure within the park. deer occur on the island, so they have no natural preda- Males are caught from August to October to have their Yamada and Urabe, Relationship between grooming and tick threat in sika deer 107 antlers cut. After cutting, males are released from the distances of 100–500 m from the top of a hill using a enclosure during October. Injured or sick deer are also telescope. In Nara Park, we observed herds at distances confined to the enclosure. Most of the deer are well of 5–50 m. Focal animal observations of 10 min were domesticated and are often fed “deer cookies” sold in the employed to record the behavior of deer. Around 20–30 park, or vegetables served by tourists and residents. focal animals were selected at random during each day. Observations were carried out mainly at four sites: The total number of 10 min observation bouts at Daibutsu-ike, Ukimido, Hakuba and Koufukuji (Yamada Mageshima was 138 for ♂ adult-large, 143 for ♂ adult- and Urabe 1998). The vegetation at Daibutsu-ike is small, 103 for ♂ yearling, 417 for ♀ adult, 115 for ♀ Zoysia spp. grassland with scattered trees. Usually, deer yearling and 127 for fawn. In Nara Park, 202 bouts were moved around the site in a loose herd, grazing and rest- recorded for ♂ adult-large, 210 for ♂ adult-small, 232 ing. At Ukimido, Prunus spp., Pinus densiflora and for ♂ yearling, 909 for ♀ adult, 245 for ♀ yearling Castanopsis cuspidata trees occupy the half of the site and 248 for fawn. The behavioral categories both in and grassland the other. Deer were often fed with vege- Mageshima and Nara Park were as follows: self-oral tables or other foods in this site. Hakuba is surrounded grooming (SOG), performed with of the tongue and inci- by Cryptomeria japonica trees and there was little grass sor-canine teeth; self-scratch grooming (SSG), scratch- available to the animals. Deer gathered in a narrow area ing with the hoof of a hind leg; allogrooming delivered by a path to get deer cookies. These three sites are used (AGD), delivered grooming to another individual; allo- by mixed herds including both adult males and females grooming received (AGR), grooming received from during most observation days, In Koufukuji, there are another individual; feeding; lying down; others, all other Zoysia spp. grassland yards with Pinus trees, and deer, behavior, including moving, caution, display and mating. most of them are males, are fed by people most fre- Data for each class were pooled separately to calculate quently. the class average for each observation period (Mageshima) The study period at Nara Park was from 20 April to 27 or month (Nara Park). Next, seasonal variation of mean November 1998. Data from all four sites in the park duration was tested for each grooming category by were combined for the following analysis. ANOVA, and then the mean duration of grooming and tick density were tested for Spearman’s rank correlation Size-sex classification of deer in all those cases for which the seasonal variation was The deer were classified into the following six classes significant. Because the deer were not identified at both by sex, body size and shape of antlers following Miura study sites, our data might include pseudoreplication by (1984): ♂ adult-large, males estimated to be 4 years or focusing on one deer several times (Hairston 1989) and older with a large body and usually 4-point antlers we have a risk of type I error in ANOVA. However, the (corresponding to age class 3 of Miura (1984)); ♂ adult- rank of mean duration itself is not influenced by the small, males estimated to be 2–4 years old with medium- result of ANOVA, and thus, the correlation analysis is sized body and 2- or 3-point antlers (age class 2 of still valid. Miura); ♂ yearling, males estimated to be 12–24 months old with a small body and 1-point spikes (age class 1 of Tick sampling Miura); ♀ adult, females estimated to be 2 years or older We measured the ixodid tick density in those areas with medium or large body; ♀ yearling, females esti- frequented by deer using the drag method (Mooring mates to be 12–24 months old with a small body; fawn, 1995). Ixodid ticks have a three-host life cycle. The under 12 months. For most females give birth to fawns ticks require a new host for each life stage (larva, nymph in June, the data from January to May in each year indi- and adult) and detach from the host following blood- cate the individuals born in the previous year, and the feeding in order to molt into the next stage. We collected data from June to December indicate the individuals born host receptive ticks of all developmental stages by pull- in that year. ing a 0.9 × 1.5m white flannel cloth along 5 m transects. At Mageshima, we established two 55 m transects from Behavioral observation grassland into a forest along deer trails. We conducted Behavioral observations were generally conducted six 5 m drags along each transect and replicated twice on between 0600 and 1800 at both sites. At Mageshima, we different days. In Nara Park, eight points were estab- mainly observed the herds on the southern grassland at lished at each of the Daibutsu-ike, Ukimido and Hakuba 108 Mammal Study 32 (2007) sites, with two replicates. In Nara Park, tick density was Relationship between seasonal tick threat and grooming surveyed monthly from April 1998 to October 1999, frequency with sampling undertaken between 1300 and 1600 on The recorded tick density on the ground was always fine days. At both site, air temperature and humidity on higher at Mageshima than at Nara Park (Fig. 3). At the ground, which have a significant effect on the activ- Mageshima, density was highest during April 1998, ity and survival of ticks (Yoshida 1980), were recorded whereas at Nara Park density was highest during Sep- by a digital thermometer. tember in both 1998 and 1999 (Fig. 3). A one-way Ticks collected were Haemaphysalis mageshimaensis ANOVA revealed significant seasonal variation in tick at Mageshima, and H. longicornis in Nara Park. At the density at both study sites (Mageshima, F = 2.517, df = 4, latter site, the drag method occasionally also collected P = 0.045; Nara Park, from April to December 1998, F = Lipoptena fortisetosa, a dipterous bloodsucker. The life 3.345, df = 8, P = 0.023; Nara Park, from January to cycle of this species has not been described. October 1999, F = 3.314, df = 9, P = 0.0014). The results of grooming frequency were shown in Results Figs. 4–7, and the statistical results were summarized in Table 1. At Mageshima, there was significant seasonal General time allocation of behavior at Mageshima variation in SOG and SSG by females and fawns, with Island and Nara Park the exception of SSG by ♀ yearling (ANOVA: SOG by Figure 2 shows the mean time spent feeding and lying ♀ adult, F = 4.456, df = 4, P = 0.0016; SOG by ♀ year- down at Mageshima and Nara Park. Grooming time was ling, F = 3.347, df = 4, P = 0.0126; SOG by fawn, F = relatively short in both sites and not shown in the figure. 9.408, df = 4, P < 0.0001; SSG by ♀ adult, F = 6.926, At Mageshima, feeding time accounted for two-thirds of df = 4, P < 0.0001; SSG by fawn, F = 7.395, df = 4, P < the total observation time during all observation periods, 0.0001) (Fig. 4). Tick density was significantly posi- and there was no clear pattern of variation. At Nara tively correlated with the frequency of SOG by ♀ adult Park, feeding time was about half that at Mageshima, and and fawn and SSG by ♀ adult (Spearman’s rank correla- the deer lay down more frequently in place of feeding. tion test: SOG by ♀ adult, P = 0.0455; SOG by fawn, Feeding duration at this site was shortest during April P = 0.0455; SSG by ♀ adult, P = 0.0455). and increased slightly in November, but was almost con- For males at Mageshima, there was a significant sea- stant from May to October. sonal variation in the duration of SOG by ♂ adult-large

Fig. 2. Mean time spent for feeding and lying down in Mageshima (upper) and in Nara Park (lower). Shaded area, feeding; solid area, lying down; open area, others. Yamada and Urabe, Relationship between grooming and tick threat in sika deer 109

Fig. 3. Seasonal change in tick density, temperature and humidity at ground surface. Bar shows SD. Solid line: temperature, broken Line: humidity. The meteorological data at Mageshima in April 1998 are lacking.

Table 1. Summary of Figs. 4–7

SOG SSG AGD AGR SV COR SV COR SV COR SV COR Mageshima ♀ adult * * *** * n.s. n.s. ♀ yearling * n.s. n.s. n.s. * n.s. fawn *** * *** n.s. n.s. n.s. ♂ adult-large ** * ** n.s. * n.s. *** n.s. ♂ adult-small *** * n.s. *** n.s. * n.s. ♂ yearling n.s. n.s. n.s. n.s. Nara Park ♀ adult n.s. * n.s. * n.s. n.s. ♀ yearling n.s. n.s. n.s. n.s. fawn n.s. n.s. n.s. *** n.s. ♂ adult-large *** n.s. *** n.s. ** n.s. n.s. ♂ adult-small n.s. n.s. n.s. n.s. ♂ yearling * n.s. n.s. n.s. n.s. SV, Seasonal variation; COR, Correlation with the tick density. * P < 0.05; ** P < 0.01; *** P < 0.001; n.s., not significant. and ♂ adult-small, SSG by ♂ adult-large (ANOVA: SOG 0.7240). by ♂ adult-large, F = 3.627, df = 4, P = 0.0077; SOG by For males at Nara Park, a significant seasonal varia- ♂ adult-small, F = 6.090, df = 4, P = 0.0002; SSG by ♂ tion was revealed for SOG by ♂ adult-large and ♂ year- adult-large, F = 4.190, df = 4, P = 0.0032) (Fig. 4). Tick ling, and for the duration of SSG by ♂ adult-large (SOG density was significantly positively correlated with the by ♂ adult-large, F = 4.094, df = 7, P = 0.0006; SOG by duration of SOG by ♂ adult-large and ♂ adult-small ♂ yearling, F = 2.338, df = 7, P = 0.0268; SSG by ♂ (Spearman’s rank correlation test: SOG by ♂ adult- adult-large, F = 4.629, df = 7, P = 0.0002) (Fig. 5). Tick large, P = 0.0455; SOG by ♂ adult-small, P = 0.0455). density did not have a significant positive correlation For females at Nara Park, significant seasonal varia- with any of these values (Spearman’s rank correlation tion was detected only for SSG by ♀ adult (F = 2.609, df test: SOG by ♂ adult-large, P = 0.1313; SOG by ♂ year- = 7, P = 0.0117) (Fig. 5), and tick density did not corre- ling, P = 0.1453; SSG by ♂ adult-large, P = 0.5661). late with this (Spearman’s rank correlation test: P = ANOVA revealed a significant seasonal variation for 110 Mammal Study 32 (2007)

Fig. 4. Seasonal change in the mean duration of self-grooming in Mageshima. Gray back-ground shows the rutting season. Bar shows SD. The number above bars in the left-side graphs shows data size, and it is applied to the right-side graphs. Means are not connected with broken lines when the cohorts are changed between two months. Probability shows the result of one-way ANOVA (see Table 1). duration of AGR by ♀ yearling at Mageshima (F = tion between any of these measures and tick density 2.903, df = 4, P = 0.0250) (Fig. 6). Once again, tick den- (Spearman’s rank correlation test: AGD by ♀ adult, P = sity was not significantly positively correlated with this 0.4014; AGR by fawn, P = 0.1627). factor (Spearman’s rank correlation test: P = 0.2301). For males at Nara Park, significant seasonal variation For males at Mageshima, there was significant sea- was found in the duration of AGD by ♂ adult-large (F = sonal variation in the duration of AGD by ♂ adult-large 2.978, df = 7, P = 0.0076) (Fig. 7), and this did not corre- and ♂ adult-small, and in the duration of AGR by ♂ late with tick density (Spearman’s rank correlation test: adult-large and ♂ adult-small (AGD by ♂ adult-large, P = 0.2719). F = 2.898, df = 4, P = 0.0244; AGD by ♂ adult-small, F = 5.135, df = 4, P = 0.0007; AGR by ♂ adult-large, Seasonal pattern of allogrooming for each size-sex class F = 6.053, df = 4, P = 0.0002; AGR by ♂ adult-small, At Mageshima, in both years, the mean duration of F = 2.929, df = 4, P = 0.0231) (Fig. 6). However, tick AGD by ♀ adults tended to increase during the rutting density was not significantly positively correlated with season (Fig. 6), while the mean duration of AGR by ♀ any of these measures (Spearman’s rank correlation test: adults fluctuated regardless of season. The duration of AGD by ♂ adult-large, P = 0.4237; AGD by ♂ adult- AGR by ♀ yearlings decreased to some extent during small, P = 0.2501; AGR ♂ adult-large, P = 0.6171; AGR the rutting season. Both AGD and AGR by fawns did by ♂ adult-small, P = 0.2301). not fluctuate significantly, but AGR tended to increase in For females at Nara Park, ANOVA revealed sig- the rutting season. nificant seasonal variation in the duration of AGD by ♀ At Mageshima, none of the ♂ adult-large delivered adult and AGR by fawn (AGD by ♀ adult, F = 2.539, df nor received allogrooming during the rutting season = 7, P = 0.0140; AGR by fawn, F = 8.361, df = 7, (Fig. 6). The mean duration of AGD and AGR by ♂ P < 0.0001) (Fig. 7). There was no significant correla- adult-small at this site also tended to decrease during the Yamada and Urabe, Relationship between grooming and tick threat in sika deer 111

Fig. 5. Seasonal change in the mean duration of self-grooming in Nara Park. See Fig. 4 for the explanation notes.

Fig. 6. Seasonal change in the mean duration of allogrooming in Mageshima. See Fig. 4 for the explanation notes. 112 Mammal Study 32 (2007)

Fig. 7. Seasonal change in the mean duration of allogrooming by males in Nara Park. See Fig. 4 for the explanation notes. rutting season. AGD and AGR by ♂ yearlings did not Discussion change significantly. At Nara Park, the mean duration of AGD and AGR by At Mageshima, tick density was significantly posi- ♀ adults peaked during June (Fig. 7), when many tively correlated with the duration of SOG in four out of females gave birth to fawns. The duration of AGD and five age-sex classes, and with duration of SSG in one out AGR by ♀ yearlings was much lower for ♀ adults, and of three age-sex classes, for which it was possible to test did not fluctuate significantly. The duration of AGD by for this. Furthermore, the seasonal variation in the dura- fawns was low, and they did not groom others in the tion of SOG by ♂ yearling shows the same pattern as the rutting season. The duration of AGR by fawns had a other age-sex classes, although this class was not statis- prominent peak in June, when they were newborns. tically significant. In Nara Park, the duration of SOG During all seasons, the duration of allogrooming by by ♂ yearling and ♂ adult-large, and of SSG by ♀ adult males at Nara Park was lower than that at Mageshima. and ♂ adult-large showed significant seasonal variations, ♂ adult-large, ♂ adult-small and ♂ yearlings did not but none of these were shown to have any significant deliver or receive allogrooming during the rutting season association with tick density. and none of these classes displayed a clear peak in the From these results, it can be concluded that the pattern duration of AGD or AGR during the non-rutting season of self-grooming, at least at Mageshima, reflects tick (Fig. 7). density in the habitat. No correlation between self- grooming duration and tick density at Nara Park may be Yamada and Urabe, Relationship between grooming and tick threat in sika deer 113 due to the low density of ticks there. Seasonal fluctua- Mooring (1995) showed that allogrooming bouts by tions in tick density at Nara Park were significant, but female impala had a significant positive association with on a very much smaller scale than was the case at the density of nymphs, and he concluded that allogroom- Mageshima. ing also had a relationship to tick density, because However, when the results of both study sites are com- nymphs and larvae favor the neck and ear regions that pared the relationship between self-grooming duration are not accessible to oral self-grooming. By inference and tick density is not clear. During April 1998, tick from Mooring (1995), self-scratch grooming, which density at Nara Park was 1/30 that at Mageshima, while grooms almost the same body region as allogrooming, the average duration of self-grooming was two-thirds should show much the same pattern as allogrooming if that at the island site. During September 1998, when tick allogrooming is influenced by the density of immature density at Nara Park was one-third that at Mageshima, ticks. In the present study, however, seasonal change in the average duration of self grooming was greater than the duration of SSG did not coincide with that of AGD or that at Mageshima. This result does not coincided with AGR in all cases, but rather showed a similar pattern to the previous studies in impala, which revealed the corre- SOG. Thus, allogrooming does not seem to be influ- lation with tick density and grooming frequency by the enced by the density of ticks on head and neck regions of data from several regions with different tick density the deer in our study. (Mooring 1995; Mooring and Samuel 1998). Some social reasons of seasonal pattern of allogroom- This inconsistency may due to differences in the time ing are easily inferred. Allogrooming by ♀ adult to allocation of behaviors between the study sites. At fawns prominently increased at Nara Park during June, Mageshima, foraging accounted for two-thirds of the when females have newborn fawns. Adult males rarely total observed time, whereas for deer at Nara Park, for- performed allogrooming during the rutting season at both aging occupied approximately half the time it did for sites, as the result of Yamada and Urabe at Nara Park deer at Mageshima, and “lying down” concomitantly (1998). Thus, our results strongly suggest that the fre- increased. This result may be modified to some extent, quency of allogrooming of sika deer is interpreted in a because our study was carried out only in daytime and in social context, rather than in connection to tick density. open areas both at Mageshima and Nara Park. The There is a remarkable difference between Mageshima behavioral pattern of deer changes temporally in a day and Nara Park for the point that who behaves as an and spatially among habitat type (Tatsuzawa and Fujita allogroomer. At Mageshima, adult males in the non- 2001). However, the reduced foraging time of deer in rutting season, and yearlings and fawns of both sexes Nara Park is almost certain, and it is clearly due to artifi- allogroomed frequently, while only♀ adults were active cial feeding. The deer in Nara Park live much longer allogroomers at Nara Park. These suggest that most than wild deer (Ohtaishi 1975), which suggests they are allogrooming observed in Nara Park may carry out from not threatened by the most important cause of death for mothers to their offspring, but in Mageshima, almost wild deer - starvation. On the other hand, the nutritional members, even fawns, behave as an allogroomer. More- condition of the deer at Mageshima has deteriorated over, reciprocal allogrooming were observed in about in recent years and their body size has become smaller one-third of the series of contiguous allogrooming bouts (Tatsuzawa 2000). The greatly reduced foraging time in (Yamada and Urabe, unpublished data), while it was Nara Park means that the deer have more than adequate very rare at Nara Park (Yamada and Urabe 1998). It is time to groom. an interesting theme if the social relationship of deer At Mageshima, the duration of AGD by ♂ adult-large through allogrooming is influences by their habitat con- and ♂ adult-small, and of AGR by ♀ yearling, ♂ adult- dition, such as tick density. large and ♂ adult-small showed significant seasonal variations, but none of them had a significant association Acknowledgments: We are grateful Dr. S. Tatsuzawa, with tick density. In Nara Park, the duration of AGD by Hokkaido University, and Ms. H. Kuroda, Osaka City ♀ adult and ♂ adult-large, and of AGR by fawn also University, for their great help for our field study in showed significant seasonal variation, but none of them Mageshima. We also thank Dr. T. Thunoda, Chiba Uni- had a significant association with tick density, too. versity, for his kind advice for the sampling of ticks. Ms. These results lead a conclusion that the duration of allo- S. Nakamura and M. Toyonishi kindly help our study in grooming has no relationships to tick density. Nara Park. 114 Mammal Study 32 (2007)

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