Mammal Study 28: 1–10 (2003) © the Mammalogical Society of Japan

Foraging activity and night-roost usage in the Japanese greater horseshoe , Rhinolophus ferrumequinum nippon

Kimitake Funakoshi1,* and Fumikazu Maeda2

1 Biological Laboratory, Faculty of Intercultural Studies, The International University of Kagoshima, Kagoshima 891-0191, Japan 2 Kyusyu Natural Environment Research, Kikuyo-cho, 869-1102, Japan

Abstract. Nocturnal activity, foraging behavior and night-roost usage in Rhinolophus ferrum- equinum nippon were examined using bat-detectors, radio transmitters and fluorescent tags. Emergence from the cave roost began around dusk from March to November. The mean emergence time averaged 14 min after sunset, and the principal foraging period was within a few hours after sunset. The main foraging style was flycatching (perch-hunting). The height of the branches used by as feeding or roosting sites averaged 4.1 m above the ground. The length of bat’s stay per feeding site averaged 6.6 min. In June, August and September, once the bats emerged, they seldom returned to the cave until just before sunrise, but in July, lactating females returned to the cave before midnight. In November, most of the bats returned to the cave about two hrs after sunset. Home range of adult females averaged 1.5 ha, and they foraged mainly in open forests, woodland paths or forest edges. The distances traveled to their main foraging sites averaged 0.87 km. Maximum-recorded distance traveled was 2.3 km during one night, and they hunted up to a distance of 2.0 km from the cave. Bats used specific night roosts, and their roosting bouts in the night roosts extended for about six hrs in April and varied from one to five hrs in June and October.

Key words: foraging behavior, home range, night-roost usage, nocturnal activity, Rhinolophus ferrumequinum nippon.

In previous papers (Funakoshi and Uchida 1978a; features are readily investigated by using bat-detectors, Funakoshi and Takeda 1998; Funakoshi and Fukue radio-tracking and banding. The present study was 2001), we investigated the hibernation, winter activity, carried out to examine the nocturnal behavior and the diet, reproduction and growth in the Japanese greater use of night roosts. As a result of this research, we report Rhinolophus ferrumequinum nippon. the first information on the nocturnal activity and night However, their nocturnal activity, foraging habitat and roosting behavior in R. f. nippon and compare this with night-roost usage have not yet been examined in detail, data from the same species in England and other species except for the reports of Kuramoto (1972, 1977). Rela- of bats. tively few papers concerning the foraging behavior and nocturnal activity in rhinolophid bats have been pub- Study area and methods lished (Neuweiler et al. 1987; Jones and Rayner 1989; Jones and Morton 1992; Jones et al. 1995). Furthermore Principal investigations were made in and around little is known about the seasonal changes in nocturnal the Katano-do Cave in Kagoshima Prefecture (31°32’N, activity of R. ferrumequinum (but see Jones and Morton 131°10’E) from the spring of 1998 to the summer of 1992; Park et al. 1999, 2000). This species is an useful 2001 (Fig. 1). The vegetation of this region consists of subject for such study, because its night roosting sites secondary laurel forests and coppice forests including are easily accessible to observers. Furthermore, the above Quercus glauca, Castanopsis sieboldii, Lithocarpus

*To whom correspondence should be addressed. E-mail: [email protected] 2 Study 28 (2003)

Fig. 2. Map showing the location of Katano-do Cave and other day  Fig. 1. Location of the study areas. a, Katano-do Cave; b, Satsuma- roosts, and the distribution of feeding perches. Feeding perches were cho General Playground. found by following bats fitted with chemiluminescent tags, and with the use of bat detectors. , Katano-do Cave; , day roosts (deserted houses); , feeding perches; , feeding perches used by young. edulis and Cryptomeria japonica, with paddy fields (Fig. Horizontal hatched areas indicate farms or paddy fields. The areas 2). A small brook flows near the entrance of the cave. of horizontal broken lines indicate open forests. Thick broken lines The width of the entrance of the cave is about 7 m, and show the woodland paths. the length of its passage is about 70 m. Rhinolophus f. nippon, R. cornutus, Miniopterus fuliginosus, Myotis Kemihotaru; Lumica Corporation, Koga, Japan) were nattereri and Myotis macrodactylus were all found at the attached to hair on their backs with instantaneous adhe- cave from spring to fall (Funakoshi and Takeda 1998). sive before release. Light from the tags lasted for ap- The inner part of the cave is very large, and was used as proximately 6–10 hours. Due to this time period, 10–20 the maternity roost by R. f. nippon in summer. In order bats were captured by an insect net at the roost one hour to investigate the total colony size, nocturnal and sea- before sunset. Sex and age were recorded, and bats sonal activities, we recorded the number of bats emerg- were marked individually with flanged wing-bands, ing each minute around sunset and the number of bats before being fitted with the tags at the entrance of the returning before sunrise at the entrance of Katano-do cave. Fluorescent light did not seem to disturb the bats Cave by using bat detectors (S-25 or Mini-3; Ultra Sound in their feeding activity. Bats fitted with light tags were Advice, London, UK). In order to determine the flying followed during eight nights in June–October 1998, four routes, the bats flying out of the cave were counted at nights in June–August 1999, and one night in May 2000. several points outside the cave by using bat detectors. Home range and foraging site fidelity were investi- These points were decided in order of distance from the gated by radio-tracking. Three bats (adult females) were cave based on the observations of the flights at various fitted with transmitters (1.2 g; Holohil Systems Ltd., points in advance. Carp, Canada) which averaged 4.8% of the body weight Flight paths, foraging sites and foraging or night- of each bat. They were tagged at the entrance of the cave roosting behavior were observed by following bats fitted on 2 June 2001. Radio-fixes for perching bats were with chemiluminescent tags or radio tags, and with the obtained by triangulation with three portable receivers use of bat detectors. The cylindrical fluorescent tags (FT-290mk II; Vertex Standard Co. Ltd., Tokyo, Japan) (37 mm, 0.5 g, on average 2% of bats’ body weight; at intervals of two hours for six nights in June–July 2001. Funakoshi and Maeda, Foraging activity and night roosting in Rhinolophus ferrumequinum nippon 3

Fig. 4. Emergence times of the bats related to sunset at Katano-do Cave in 1998–2000. , and , a time when the first bat emerged from the cave in 1998, 1999 and 2000 respectively; --- , --- and --- , start and end of emergence; , and , mean emer- Fig. 3. Map showing the location of Satsuma-cho General Play- gence time. ground and the outhouses, and distribution of day roosts. G, Satsuma- cho General Playground; A, B and C, outhouses with two rooms respectively; , day roosts (artificial caves); , day roosts where the bats banded at the outhouses were recaptured. Diagonal shaded areas by R. f. nippon. indicate woodlands. Results In order to investigate the movements in one night on the basis of banding-recapture method, other captured Emergence time and flight direction bats were also marked with wing-bands and then At Katano-do Cave, activity before sunset and light- released at the entrance of the cave in 1998–1999. After sampling behavior were observed in R. f. nippon. Emer- that, other day-roosts were searched. gence began around dusk and continued for about 30 min It was difficult to observe the night-roosting behavior after sunset from March to November (Fig. 4). The after mid-night, because the power of fluorescence mean emergence time averaged 14 ± 1.7 (SE) min (range became weak. Therefore, it was desirable to find spe- 2–22 min, n = 24) after sunset. cific night roosts in order to examine their nonfeeding In June, R. f. nippon flew in every direction from the phases during the night. There were three outhouses cave, but most of the bats flew rather low over the used as night roosts by R. f. nippon at the second study ground (1–2 m) and headed eastward. Of 20 bats carry- area which was Satsuma-cho General Playground in ing fluorescent lights, 14 (70%) flew to the east, and of Kagoshima Prefecture (31°55’N, 130°33’E) (Fig. 1). 120 bats in the cave, 65 (54%) were counted at the east The distance between the outhouses A and B was about point by a bat detector (Fig. 5). In July, of 19 bats carry- 100 m, and that between B and C was about 200 m ing fluorescent lights, 11 (58%) flew to the southeast, (Fig. 3). Each outhouse had two separate rooms having and of 220 bats that were counted at the entrance of one entrance each. Nocturnal observations using bat the cave, 157 (71%) were counted at point A, while 49 detectors and/or banding were carried out for nine nights (22%) were counted at point B. In August, of 20 bats in May–November 1998, one night in September 1999, carrying fluorescent lights, 4 (20%) flew to the southeast one night in April 2000, and four nights in May–August to point C, and of 117 bats in the cave, 40 (34%) were 2002. In order to investigate use of night roosts, the counted at the same point. In September, of 100 bats in presence of prey remains beneath roosts was examined, the cave, 24 (24%) were counted at point D. Almost all and number of feces for a night was counted in each the bats flew directly towards the southeast or east, and room. In addition, their day roosts were searched in the then gradually dispersed. The flight paths of R. f. nippon study area. In the night roosts and day roosts, the were situated in relatively open space. ambient temperatures were measured at sites occupied 4 Mammal Study 28 (2003)

Fig. 5. Flight directions of R. f. nippon departing from Katano-do Cave as observed at the entrance of the cave on 15, 27 June 1998 (a) and at sites A, B, C, and D on 22 July, 21 July, 17 August, and 3 September 1998 respectively (b). Arrows show the flight directions. Numerals indicate the number of the bats carrying fluorescent lights in Fig. 6. Changes in percentage proportions of numbers of active and each direction, and numerals in parentheses indicate the number of inactive bats at night in the summer of 1998–1999. Levels of activity flying bats counted by using a bat-detector. , Katano-do Cave. were observed by following bats fitted with chemiluminescent tags, and they were randomly found. Sample sizes are indicated above each bar. Foraging behavior and nocturnal activity Rhinolophus f. nippon emits pure tone frequencies. The constant frequency averaged 72 kHz, and the maxi- mum distance that we could receive their echolocation sounds by using bat detectors was about 17 m. They hunted by flycatching and aerial hawking, but flycatch- ing was more common. In flycatcher style, they emit sounds and detect insects from their perch. They then pursue and catch their prey and return to the same perch to eat it. The height of the perch used by the bats aver- aged 4.1 ± 0.32 (SE) m above the ground (range 2–10 m, n = 35). The length of their stay at the same perch averaged 6.6 ± 1.7 (SE) min (range 1–42 min, n = 25). The distance between the bats hanging on perches along the woodland paths averaged 103 ± 9 (SE) m (range 45– 210, n = 31) in July and August 1998 and July 1999. The active bats frequently echolocated, pursued and ate their prey at perches. The inactive bats, on the other hand, intermittently emitted feeble sounds while hanging on twigs and did not foraged. Each of the bats fitted with the light tags was counted at night. Out of the bats observed more than 90% were active within one hr after sunset (Fig. 6). But, three hrs after sunset, more than 70% of the bats became inactive (Fig. 6). Therefore, the principal foraging period was within a few hrs after emergence.  Fig. 7. Seasonal and nocturnal changes of the number of bats flying Changes in the return number at Katano-do Cave are into Katano-do Cave in 1998. Solid histograms show the return num- shown in Fig. 7. In June, all of the bats began to return to ber at 10 minutes intervals from post-midnight until dawn. , sunrise the cave one hr before sunrise. However, in the lactating time. season in July, flying activity became intermittent during the night, and most of the bats returned after mid-night. midnight, but most of the bats began to return at about From late July to mid-September, they seldom returned 5:00 am. In November 1998, most of the bats returned to the cave until one or two hrs before sunrise. In Octo- only about two hrs after sunset. ber, some of the bats returned to the cave two hrs after Of 15 lactating females fitted with fluorescent tags in Funakoshi and Maeda, Foraging activity and night roosting in Rhinolophus ferrumequinum nippon 5

Fig. 8. The home ranges of three adult females of R. f. nippon in June–July of 2001. Locations of feeding or roosting sites are indicated Fig. 9. Seasonal changes in night-roost usage in the outhouses at individually by open circles (R1), triangles with apex down (R2), and Satsuma-cho General Playground. --- , --- and --- ; squares (R3). The areas of horizontal broken lines indicate open start and end of the usage in 1998, 1999 and 2000 respectively. forests. Points a, b and c represent the main foraging sites for six days respectively. , Katano-do Cave. distances averaged 0.87 km. Minimum area convex June 1999, 9 (60%) had returned one hr after sunset, and polygons show the home ranges of the bats tracked 14 (93%) had returned to the cave two hrs after sunset. during the summer (Fig. 8). In some areas, the home Of 20 adult females fitted with tags in May 2000, 8 ranges overlapped each other. The home range sizes (40%) had retuned four hrs after sunset, and 13 (65%) of R1, R2 and R3 were 2.7 ha, 0.6 ha and 1.2 ha, respec- had returned to the cave five hrs after sunset. tively, and the ranges averaged 1.5 ha (n = 3).

Foraging areas and home ranges Night-roost usage The bats foraged mainly in open forests, woodland The bats began to enter the outhouses at Satsuma-cho paths or forest edges (Fig. 2). They were not found in General Playground to roost two hrs after sunset in April dense forests, open areas or near brooks. The adult to early June and October (Fig. 9). From mid-June to females carrying fluorescent-tags foraged up to a dis- September, however, they began to roost from three to tance of 2.0 km from Katano-do Cave. Early in August, five hrs after sunset (Fig. 9). Especially, from mid-June the newly weaned young foraged within 0.5 km of the to July the period of usage was only about three hrs. roost, and sometimes returned to the roost together with Seasonal and nocturnal changes of the frequency of their mothers at midnight. In the study area, four day- night-roost usage are shown in Fig. 10. In May, the roosts were found around the Katano-do Cave (Fig. 2). number of bats roosting during the night had two peaks Maximum distance traveled was recorded as 2.3 km with the first one three hrs after sunset and the second during one night on the basis of banding-recapture one eight hrs after sunset (Fig. 10). Similar pattern was method at the day-roosts. Three bats (R1, R2 and R3) also shown in May 2002. In June, the number of roost- were tracked for six nights (Fig. 8). The greatest distances ing bats reached a peak about three hrs before sunrise. traveled by each radio-tracked individual were 1.98 km From June to September, such peaks were concentrated for R1, 0.48 km for R2 and 1.03 km for R3. The princi- after midnight. In July, the number of bats roosting pal foraging areas, however, were widely separated. The during the night was reduced (Fig. 10). In October, the distances traveled to such main foraging sites were 1.2 bats started using the night roosts two hrs after sunset, km for R1, 0.4 km for R2 and 1.0 km for R3, and the and many of them began to leave the roosts four hrs after 6 Mammal Study 28 (2003)

Start of night-roost usage and roosting bout were varied with individuals (Fig. 11). Roosting bouts lasted for 1–5 hrs, and average duration of the bouts was 136 ± 21 (SE) min (n = 14) in early June (Fig. 11). In other seasons, roosting bouts lasted for three hrs in October 1998, and six hrs in April 2000. The total number of feces on the floors of the three outhouses reached 319 for a night in June, whereas those of feces decreased to 80 in September and 65 in October 1998. Similarly, 74 feces were found in September 1999. No prey remains beneath roosts were found in any season. The ambient tempera- tures at night-roosts by R. f. nippon ranged from 17 to 22°C in June, while those in day roosts ranged from 14 to 16°C. During daytime searches, new 12 day-roosts were found around the playground (Fig. 3). Of 26 bats banded at the outhouses at night, 10 of them were recaptured at the two day-roosts (artificial caves). Thus, the distance traveled to the night roosts varied from 0.6–0.8 km. Seasonal and nocturnal changes of the members of bats in each night-roost were shown in Fig. 12. In July, Fig. 10. Seasonal and nocturnal changes in the total number of bats hanging on the ceiling of the outhouses at Satsuma-cho General Play- only one bat roosted in each room at different times after ground in 1998. , sunrise time. midnight, and three bats captured were adult males (Fig. 12). In August, of nine bats captured, all of them were adult females and young (Fig. 12). Their nursery colony, however, was not found around the night roosts at the study area. In September, of nine bats captured, seven were females, and only one adult male was found (Fig. 12). In room A1, four bats were found at the same time. In October, of 10 bats captured, six were adults and four were adult males (Fig. 12). Especially, in room A1 and C1, a pair of male and female was found respectively. On the night of 18–19 September 1999, eight bats roosted in the outhouses (A1, A2, B2, C1 and C2), one of which was recaptured (F45 rested in room A1). On the night of 11–12 May 2002, six bats (two adult males, two adult females, one subadult male and one subadult female) roosted individually in rooms B1 and C1 at different times. One of them (adult male G83) was recaptured in room C1, and has since been captured in the same room in July and September 1998. On the Fig. 11. Roosting patterns of the bats (a–n) in the outhouses at Satsuma-cho General Playground on 6–7 June 1998. A, B and C night of 15–16 June 2002, five bats roosted in rooms A1, indicate the outhouses with two rooms respectively. Solid bars show A2, B2, and C1 at different times. Three of them were duration of roosting. , sunrise time. adult males, one subadult female and the other one sub- adult male. The subadult male (F811) was recaptured in sunset (Fig. 10). On the other hand, in this season some room C1, and has since been captured there on 12 May of the bats began to return the Katano-do Cave five hrs 2002. One owl (Strix uralensis) was observed near the before sunrise (Fig. 7). In November, there were no bats outhouse (C1, C2) at 0:40 am on 16 June, 2002. On the in the outhouses at night. night of 21 August 2002, three adult females and one Funakoshi and Maeda, Foraging activity and night roosting in Rhinolophus ferrumequinum nippon 7

equinum. In addition, the flight paths of R. f. nippon were located in rather open spaces. Rinolophus f. nippon is probably more susceptible to aerial predation than R. cornutus. On the other hand, M. fuliginosus which hunts in open spaces far above the woodland canopy emerged later than the other species. Miniopterus fuliginosus may avoid the risk of predation by diurnal or nocturnal preda- tors in the sky. The flight paths of the emerging bats were evidently specific. They flew rather low above the ground (1–2 m), as stated also by Ransome (1990). Most of the bats flew directly toward the southeast or east (Fig. 5). In these directions, there were densely distributed foraging sites (Fig. 2). In addition, they repeatedly used certain foraging areas. It seems that their food supply distribu- tion was patchy in space over the study area. Foraging in flycatcher style presented here is con- sistent with a previous field study of R. ferrumequinum (Jones and Rayner 1989), and is similar to that in rela- tively small Rhinolophus rouxi (Neuweiler et al. 1987). Rhinolophus ferrumequinum sometimes hunts by aerial hawking: bats detect, pursue and eat their prey on the wing (Kuramoto 1972; Funakoshi 1978; Jones and Fig. 12. Seasonal and nocturnal changes of the members of bats Morton 1992; this study), or by ground gleaning: landing present in the night roosts at Satsuma-cho General Playground in 1998. A, B and C indicate the outhouses with two rooms respectively. on the ground and catching flightless insects (Stebbings , adult females; , subadult females; , young females; , adult 1977). It is conceivable that R. ferrumequinum is flex- males; , young males; , unknown age and sex; —, no bat roosting. ible in its foraging style according to habitat. Thick arrows indicate sunrise times. Thin arrows indicate individual Jones and Rayner (1989) suggested that flycatching movement. evolved in species feeding on large insects that the bats find difficult to handle during flight. Relatively large subadult female grouped in room C1, two of the adults Megadermatid bats forage in flycatcher style (Vaughan were recaptured (F47 and F49). During the above study and Vaughan 1986; Norberg and Rayner 1987). Many of period, adult males were not found together in the same them perch on low branches and listen for ground-dwell- room. ing prey (Vaughan 1976; Fiedler 1979). The smaller (R. hipposideros), in contrast, is Discussion predominately an aerial hawker (Jones and Rayner 1989). Large insects generally occur at lower densities, Foraging strategies and the energetic cost of perching and waiting for large The emergence time in R. f. nippon approximately prey may be considerably lower than continual flying followed sunset (Fig. 4). On average, Rhinolophus f. while hunting. nippon emerged about ten min earlier than Miniopterus The principal foraging period was within a few hrs fuliginosus, Myotis nattereri and Myotis macrodactylus, after sunset, and was followed by resting bouts (Fig. 6). but about five min later than R. cornutus. Emergence at Similar activity has been shown in R. ferrumequinum in this time seems to be related with the appearance of England (Jones and Morton 1992) and in R. rouxi in Sri insect prey, and with the feeding sites of relatively light Lanka (Neuweiler et al. 1987). In addition, one or two open forests and with predation risk (Jones and Rydell short foraging bouts were shown between midnight and 1994; Funakoshi and Takeda 1998). This idea is con- sunrise (Jones and Morton 1992). Rhinolophus f. nippon sistent with the smaller R. cornutus, which forages in probably forage on their way from night roosts to day darker dense forests, emerging earlier than R. ferrum- roosts just before sunrise in our study areas. In any case, 8 Mammal Study 28 (2003) such a foraging pattern appears to be closely correlated defended against intruders (Neuweiler et al. 1987). with the emergence of insect prey, as shown in M. fuliginosus (Funakoshi and Uchida 1975) and P. Use of night roosts abramus (Funakoshi and Uchida 1978b). In addition, The timing and duration of nightly rest periods vary seasonal dietary shifts and food choice was consistent with seasons, according to ambient temperature, repro- with the changes of food conditions throughout the year ductive condition and prey availability (Kunz 1982). In (Funakoshi and Takeda 1998). this study, the bats began to use the night roosts around Greater horseshoe bats foraged mainly in open forests, foraging sites or day roosts two hrs after sunset in spring woodland paths or forest edges, and were not found in (Fig. 9). In some cases, the duration of roosting bouts dense forests, open area or over brooks (Fig. 2). At at night lasted for six hrs in April. Most of the bats Akiyoshi-dai Pateau, they foraged also over the river or returned to their day roosts before midnight in May. above the grassland (Kuramoto 1972). In England, many Such short feeding bouts may be closely related to short- R. ferrumequinum are found in patches of trees where age of prey for R. f. nippon including large Coleoptera they hunt by flycatching (Jones and Rayner 1989). They and Diptera during this season (Funakoshi and Takeda forage in ancient woodland during the spring, and over 1998). pasture containing trees during late summer (Jones and In summer, once the bats emerged at sunset, they Morton 1992). The seasonal shift in habitat use may be seldom returned to the cave until one or two hrs before associated with the seasonal changes in food abundance sunrise, except for lactating females early and mid-July. and prey (Jones 1990; Jones et al. 1995). In contrast, R. Most of the bats became inactive on a perch three hrs rouxi forage only in dense forest and are not found in after sunset (Fig. 6). Roosting bouts during the night open woodland in October (Neuweiler et al. 1987). For roosts varied from one to five hours (Fig. 11). Nocturnal R. f. nippon, the flexible food choice (Funakoshi and flying insects were most abundant in June and July Takeda 1998) and the foraging site selection mentioned (Funakoshi and Takeda 1998), and in this season, the above may well fit in with the seasonal changes of food feces on the floor of the outhouses at Satsuma-cho availability in warm-temperate regions. General Playground were great in number. Thus, in In the study area, the adult females foraged up to a dis- summer most of the bats may forage or roost on a perch tance of two km from the cave, which was much shorter intermittently during the night, and not frequently use than that of the same species in England (5 km; Jones night roosts. and Morton 1992). Such short distances in our study In all of the outhouses, prey remains were not found area may be associated with the distances traveled from beneath roosts. The results indicate that the outhouses the day roosts to feeding areas, or with abundance of are exclusively used as roosting places at night, and do food resources around the day roosts. The newly weaned not serve as masticating sites like feeding perches. young foraged up to distances of 0.5 km, similar to those Warmer ambient temperatures of the night roosts may reported in the same species (Jones et al. 1995). Mother promote the digestion of food or absorption of nourish- and young frequently leave the day-roost together in the ment, as stated by Kunz (1982). In addition, if night flying stage (Matsumura 1988), but during weaning the roosts are within easy reach of the feeding sites, the cost development of foraging behavior in R. ferrumequinum of the movement between them may be reduced. Night involves no tuition from the mother, and young forage roosts may also act as protection from predators, such as independently of their mothers (Jones et al. 1995). owls (Kuramoto 1977). The home ranges of individuals overlapped each other In July, in the breeding season, lactating females had to some extent, but each principal foraging site that was short foraging bouts just after sunset, and most of them repeatedly used was separate (Fig. 8), and bats hanging had returned to the nursery cave by two hrs after sunset. from trees were widely spaced along the woodland paths. After that, they embraced infants until sunrise (Sano We observed no territorial behavior at foraging sites, 2000; Funakoshi and Fukue 2001). As a result, lactating however, and thus, it seems that R. f. nippon has no clear females did not use night roosts. Only adult males were feeding territory, as stated by Jones and Morton (1992). captured at the night roosts after midnight in July (Fig. They may avoid the struggle for defending food by 12), and some of them were recaptured in September spacing. Rhinolophus rouxi, perch on specific twigs, and October. It is possible that adult males change their hunt in flycatcher style and its foraging areas are not night roosts seasonally, and do not defend these roosts Funakoshi and Maeda, Foraging activity and night roosting in Rhinolophus ferrumequinum nippon 9 for extended periods in summer. Adult males were 62. always solitary in night roosts, and a few used the same Funakoshi, K. and Uchida, T. A. 1975. Studies on the physiological and ecological adaptation of temperate insectivorous bats. I. roost repeatedly. In August, the night roosts were Feeding activities in the Japanese long-fingered bats, Miniopterus occupied by the postlactating females and their young. schreibersi fuliginosus. Japanese Journal of Ecology 25: 217– In September, subadult females and adult males began 234 (in Japanese with English synopsis). to use the night roosts, together with the adult females Funakoshi, K. and Uchida, T. A. 1978a. Studies on the physiological and ecological adaptation of temperate insectivorous bats. II. and young (Fig. 12). Hibernation and winter activity in some cave-dwelling bats. In October, the bats used night roosts from about two Japanese Journal of Ecology 28: 237–261. hrs after sunset, and began to return to their day roosts Funakoshi, K. and Uchida, T. A. 1978b. Studies on the physiological and ecological adaptation of temperate insectivorous bats. III. five hrs before sunrise (Figs. 7 and 9). Possibly, they use Annual activity of the Japanese house-dwelling bat, Pipistrellus night roosts or day roosts for many hours at night when abramus. Journal of the Faculty of Agriculture of Kyushu Univer- feeding is unproductive in this season. As stated by sity 23: 95–115. Anthony et al. (1981), long night roosting periods and Jones, G. 1990. Prey selection by the (Rhinolo- phus ferrumequinum): optimal foraging by echolocation? Journal short foraging periods are associated with cool nights of Ecology 59: 587–602. and low prey density. In the mating season, a pair of Jones, G., Duvergé, P. L. and Ransome, R. D. 1995. Conservation adult male and female was found in October (Fig. 12). It biology of an endangered species: field studies of greater horse- shoe bats. Symposium of the Zoological Society of London 67: is probable that night roosts may be used as a mating 309–324. place late in autumn. In November, the bats only used Jones, G. and Morton, M. 1992. Radio-tracking studies on habitat use the night roosts for a short period and returned to their by greater horseshoe bats (Rhinolophus ferrumequinum). In (I. G. Prede and S. H. Swift, eds.) Wildlife Telemetry. Pp. 521–537. day roosts about two hrs after sunset. They considerably Ellis Horwood, New York and London. restrained their activity in this season, as in M. fuligin- Jones, G. and Rayner, J. M. V. 1989. Foraging behavior and echoloca- osus (Funakoshi and Uchida 1975) and P. abramus tion of wild horseshoe bats Rhinolophus ferrumequinum and R. (Funakoshi and Uchida 1978b). hipposideros (Chiroptera, Rhinolophidae). Behavioral Ecology and Sociobiolgy 25: 183–191. Jones, G. and Rydell, J. 1994. Foraging strategy and predation risk as Acknowledgements: We would like to thank Tomonori factors influencing emergence time in echolocating bats. Philo- Masuyama, Takumi Nagata, Yusuke Ito, Hiromi Naka- sophical Transactions of the Royal Society of London B346: 445–455. mura, Mariya Shioiri and Seiji Wakiyama for their help Kunz, T. H. 1982. Roosting ecology of bats. In (T. H. Kunz, ed.) with the field work, and Hirotoshi Waki for allowing us Ecology of Bats. Pp. 1–55. Plenum Press, New York and London. to use Satsuma-cho General Playground. We are also Kuramoto, T. 1972. 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