Mammal Study 30: S25–S30 (2005) © the Mammalogical Society of Japan

A brief review of the biology on moles in Japan

Yasushi Yokohata

Laboratory of Wildlife Conservation, Department of Environmental Biology and Chemistry, Faculty of Science, University of Toyama, Gofuku 3190, Toyama City, Toyama 930-8555, Japan

Abstract. There are six species of moles, mizura, uchidai, M. tokudae, M. echigo, M. imaizumii and M. wogura, in in Japan, and the diverse talpid fauna has been interested in for a long time. Aspects on the , morphology, ecology and conservation of them were reviewed briefly.

Key words: Euroscaptor, Japan, Mogera, moles, Talpidae.

Japanese Archipelago is a small region of only 378 many turns and twists. Imaizumi (1955) proposed three million km2, but it is distributed by eight species of subspecies of E. mizura, E. mizura mizura, E. mizura Talpidae, including six species in the genera of ohtai and E. mizura hiwaensis (also see Imaizumi 1960), Euroscaptor (1) and Mogera (5) (Abe 2005), and the but some biologists are suspicious to their distinctness. diverse talpid fauna has been interested in for a long time. For example, E. mizura hiwaensis were described based In this paper, I review the aspects on taxonomy, mor- on obscure differences of body size and pelage color, phology, ecology and conservation of the moles in Japan with only one specimen, which were captured western briefly. Bibliography of Japanese Insectivora (Motokawa Honshu, about 300 km apart from western limit of E. et al. 1999) includes total 808 papers, of which 104 are ohtai. But some new localities were reported in western on the moles (23 on Euroscaptor, one on Nesoscaptor, Japan after then (e.g. Sagara et al. 1989; Kanaizuka et (= Mogera) and 80 on Mogera (papers on multiple al. 1991), so that the apparent distribution of E. mizura genera were counted repeatedly). Recent papers since in western Honshu became more continuous. 1999 are not included in these numbers, however, the Taxonomical status of the Japanese moles in the genus researchers in this field is increasing recently in Japan, Mogera was basically established by Abe (1967), which and some excellent studies are now being conducted. recognized three distinct species, M. tokudae (= present M. tokudae and M. etigo), M. wogura (= present M. Taxonomy imaizumii) and M. kobeae (= present M. wogura in Japan and M. robusta in the Continent). M. tokudae had ever In Japan, we can find six species of moles; Japanese been treated as a subspecies of M. kobeae (e.g. Imaizumi mountain mole Euroscaptor mizura (Gunther 1880), 1960), but its distinctness has become to be undoubted in Senkaku mole Mogera uchidai (Abe et al. 1991), Sado Abe’s thorough analysis of morphological variations of mole M. tokudae Kuroda, 1940, Etigo mole M. etigo many skull specimens of the moles (also see Yoshiyuki Yoshiyuki and Imaizumi, 1991, lesser M. 1986). Smaller individuals of M. imaizumii in some imaizumii (Kuroda 1957) and large Japanese mole M. mountainous and insular habitats have often been treated wogura Temminck, 1842, and all of them are endemic in as a subspecies M. wogura (= imaizumii) imaizumii or M. this country (Abe 2005). Their distribution map with wogura minor in some literatures (e.g. Imaizumii 1960), recent information is shown in Fig. 1. General features but it is thought as local phenotypical variations of M. of them are compiled in Abe (2005). imaizumii (see 2. morphology). The scientific names of The taxonomical treatment of them has been with the lesser Japanese mole and the large Japanese mole are

E-mail: [email protected] S26 Study 30 (2005)

since Abe (1995). Further, moles in Echigo Plain in central Honshu, which have been treated as a part of M. tokudae since Abe (1967), showed different chromo- some constitution from that of M. tokudae in Sado Island (type locality of this species) off the Echigo Plain. In both two cases, those differences are thought to be enough to establish reproductive isolation. Hence, M. wogura was recognized as an endemic species of Japan, and the moles in Echigo Plain were did as distinct spe- cies, M. etigo, based on the nomenclature by Yoshiyuki and Imaizumi (1991), although they did not conduct chromosomal analysis. The establishing history of fauna of Mogera spp. in Japan is shown in Shinohara et al. (2005) in this proceeding.

Morphology

Abe (1967) found remarkable geographical body size variation of the moles of the genus Mogera. In general, the moles of this genus in suitable habitats, such as large alluvial plains with soft and fertile soil and abundant soil vertebrates, are obviously larger than those in unsuitable habitats such as small islands and narrow valleys. Abe Fig. 1. Distribution map of moles in Japan with recent information. It is mainly based on Abe (1997, 2005) and Biodiversity Center of (1997) used multivariate analytical methods to detect Japan (2002), with some information on Euroscaptor mizura in west- significant environmental factors determining body ern Honshu (Sagara 1999), Mogera imaizumii (a: Tottori Prefectural size of the moles. In this analysis, habitat area (= popu- Government 2002 (to be reexamined); b: Hikone City Goverment lation size of the moles) was most significant among 2005; c: Dr. Nishigaki, personal communication) and M. wogura (d: Kawaguchi 2003). Distribution of the continental Mogera robusta is various factors (temperature, precipitation, etc.). Fur- partly shown. Mogera echigo (E), M. tokudae (T), M. uchidai (U) and ther, Yoshiyuki and Imaizumi (1992) described the size Euroscaptor mizura (others), M. imaizumii, M. wogura, M. robusta. variation in one local population of M. imaizumii in the central area of Honshu. changed from M. wogura and M. kobeae to M. imaizumii There are several cases of albino moles in Japan (M. and M. wogura, respectively, based on surveillances of imaizumii: Fujiwara 1960a, 1961; M. wogura: Fujiwara type specimens (Abe 1995; Motokawa and Abe 1996). 1960b; Yokohata 1997a). Imaizumi and Yoshiyuki The Senkaku mole was described by Abe et al. (1991) (1960) found higher occurrence of albinism in Japanese as a new species Nesoscaptor uchidai from Uotsuri-jima moles than in larger Japanese -moles, Urotrichus in Senkaku Islands, with proposal of the new genus talpoides and pointed out that this difference might be Nesoscaptor, mainly based on difference on total teeth related to low predation pressure because of the subterra- number (38) from that of the genus Mogera (42). But nean habit of the moles. Motokawa et al. (2001) included this species to Mogera, Dentition of Japanese moles has been studied on E. because of shape similarity of the skull of this species mizura (Sakai and Hanamura 1973), M. imaizumii to that of Mogera insularis from Taiwan, which is near (Sakai and Hanamura 1973) and M. wogura (Sakai and the Senkaku Islands, and because of frequent variation Hanamura 1973). Yokohata (1999) estimated species in the dental number of M. insularis. of a fossil skull specimen of the genus Mogera from Kawada et al. (2001) conducted detailed analyses on Shizuoka Prefecture, central Japan as M. wogura using the karyotypes of many moles from Japan and Korea. discriminant functions. Tsuchiya (1988) described karyo- They found remarkable differences of chromosome con- types of Japanese moles, as well as Kawada et al. (2001), stitution between M. wogura in Japan and continental which was cited above. moles, which have been treated as a part of M. wogura The description of hair types of E. mizura, M. Yokohata, Brief review on moles in Japan S27 imaizumii and M. wogura was provided by Sagara and Imaizumi 1966; Yokohata 2000; M. wogura: (1986), with special reference to the presence of straight Yokohata 1994, 1997b). All of these studies are based guard hairs in these talpids. on the wearing pattern of upper molars. Yokohata (1998) showed a case of six yr-old individual of M. wogura with Ecology observation of annual rings in its upper canine teeth. There is no precise study on the social system and activ- Ecological aspects of the Japanese talpids are ity pattern of Mogera species in Japan, but they have reviewed in Yokohata (1998) in detail. This chapter is a experimentally been thought to be solitary and active summarized a review with some recent findings on the both in day and night (e.g., Abe 1968), as is also possibly moles since 1999. Ecological information on E. mizura true on E. mizura. Some observations on the thigmotaxis and M. uchidai had been very few and limited only to of Mogera species in Japan have ever been performed fragmental records at capture (E. mizura: Fujiwara 1955, (M. imaizumii: Tezuka 1958; M. wogura: Makita 1980). 1957; Abe 1968; Yukawa 1968; M. uchidai: Shiraishi Several studies have been performed on a synbiotic and Arai 1980). relationship in forest environment between some Japa- Diets of the moles of genus Mogera have been studied nese talpids, mainly moles, and mashrooms, Hebeloma by some researchers (Fujiwara 1956a; Ondo and Fukuda radicosum and H. spoliatum (Euroscaptor: Sagara 1998; 1961; Abe 1964, 1968). These studies show that main Sagara et al. 1989, 1993a; Mogera: Sagara 1978, 1980, diets of the moles are earthworms and no apparent differ- 1998; Sagara and Abe 1993; Sagara et al. 1993a; also ence is observed among Mogera species, so that strong Urotrichus: Sagara 1998; Sagara et al. 1981, Apodemus interspecific competition is suggested to take place. (rare): Sagara et al. 1993b, soricids (probably ) Imaizumi (1978, 1979) described the earthworm forag- (rare): Sagara, personal communication). These mash- ing behavior of M. imaizumii. He described two charac- rooms grow from the latrines of the moles and other teristic hunting tactics of the mole to the earthworms, i.e. insectivorous as a nitrogen source, develop back-with-grip and bite-and retreat. The former tactic is their ectomycorrhiza around the roots of trees of some retreating with the earthworms bit in its mouth in its genera, such as Fagus and Quercus, as a carbon source, tunnel to pull the preys into the tunnel, and the latter is and form fruit bodies on the ground surface. The latrines repeating of bite, release and retreating the struggling the of moles of genus Mogera are usually made near their preys to immobilize them. Further, the mole frequently nests, so that we can find the nests under the fruit bodies. ate the preys in order from their heads to their tails. This Before this finding, the reports on the Japanese mole’s feeding behavior was observed also in U. talpoides, nests were fragmental (M. imaizumii in Shizuoka and however, the frequency is lower in the mole than in the Ibaragi Prefectures: Tashiro 1958; Horikoshi et al. shrew-mole. 1967, respectively; M. wogura in Hiroshima Prefecture: Based on observations of many specimens, litter size Fujiwara 1956b). Sagara (1995, 1999) stated this phe- of both M. tokudae (including M. etigo) and M. imaizumii nomenon as a “habitat cleanig synbiosis”. is 2–6 (Abe 1968), and that of M. wogura is 3–6 Interspecific ecological relationships among the (Fujiwara 1960c, 1962). These studies and Hoslett and Mogera species in Japan have been much interested and Imaizumi (1966, 1967) showed that the reproductive repeatedly studied by some researchers. Strong interspe- season of Mogera species in Japan is usually limited in cific competitive exclusion of smaller M. imaizumii by spring with a slight difference among regions. However, larger M. wogura, and the subsequent parapatric distribu- Sagara and Abe (1993) found three youngs of M. wogura tion of these two species, has been observed in several in their nest in October in western Japan, and Kawaguchi regions in Japan (e.g. Abe 1967, 1968, 1974, 1985, (2004) obtained a sexually active male and a lactating 2001a, b; Kushihashi 1982). Abe (1974, 1985) observed female of M. wogura in October and September, respec- the distribution area of M. wogura expanded 3 km in 14 tively. Further, Hashimoto and Abe (2001) reported years (1959–1973) and 0.6 km in 6 years (1973–1979) usual autumn breeding of M. imaizumii in Echigo Plain, with displacing M. imaizumii in Kiso Valley in Nagano central Japan where this species and M. etigo inhabit Prefecture, central Honshu. In many cases, M. wogura parapatrically. Age structure of Mogera species in Japan immediately excludes M. imaizumii in soft and rich soil has been studied repeatedly (M. tokudae (including M. in fertile alluvial plains, and some isolated relic popu- etigo): Abe 1968; M. imaizumi: Abe 1964, 1968; Hoslett lations of M. imaizumii exist in Shikoku and western S28 Mammal Study 30 (2005)

Honshu, where M. wogura dominates. The expansion of bare areas were recognized using artificial satellite M. wogura is often inhibited where the soft, fertile soil images (Yokohata et al. 2003). Environmental Ministry is lacked (Abe 2001a, b). Further, the Echigo Plain in of Japan (2002) and Okinawa Prefectural Government central Honshu, smaller M. imaizumi is excluding larger (2005) appointed this species to Critically endangered. M. etigo, and the isolated small populations of the latter Three academic societies in Japan adopted urgent species are surrounded by the former species (Imaizumi appeals in 2002 and 2003 (Yokohata 2003), but unfortu- and Imaizumi 1970). In this plain, the difference in the nately, countermeasures of this problem is very difficult, number of reproductive periods per year between the two because of political conflicts on the sovereignty of the species stated above may be attributable to the domi- Senkaku Islands among China, Japan and Taiwan. nance of M. imaizumii (Hashimoto and Abe 2001). Parasitological studies have been conducted on the Acknowledgments: I thank Dr. N. Sagara, Emeritus Pro- endoparasites of E. mizura (Duszynski and Wattam fessor of Kyoto University, for providing the information 1988; Yokohata and Sagara 1995) and of Mogera spp. in on the nest of soricids under the fruit body of Hebeloma Japan (Yamaguti 1941; Duszynski and Wattam 1988; radicosum. Drs. M. Nishigaki, Fukui Prefectural Coastal Yokohata et al. 1988a, b, 1989; Yokohata and Abe 1989; Nature Center and Y. Abe, Taga Town Museum are also Sawada and Koyasu 1991; Yokohata and Sagara 1995; acknowledged for providing information on the distribu- Yokohata and Onodera 1997; Sakata and Asakawa tion of M. imaizumii. 2003). In these studies, there have been reported 2 pro- tozoan and 4 nematode parasites from E. mizura. From Mogera spp., 3 protozoan, 1 trematode, 2 cestode and 18 Reference nematode species. The endoparasite fauna of the moles Abe, H. 1964. of Japan (1), Insectivora (genus Mogera), of Mogera apparently similar among each species on this Mogera wogura. Honyurui Kagaku [Mammalian Science] 7: 1– genus, though no information exists on that on M. 10 (in Japanese). Abe, H. 1967. Classification and biology of Japanese Insectivora uchidai. The information on the ectoparasites of Japa- (Mammalia) I. Studies on variation and classification. Journal of nese moles is scarce. Faculty of Agriculture, Hokkaido University 55: 191–265. Abe, H. 1968. Classification and biology of Japanese Insectivora (Mammalia) II. Biological aspects. Journal of Faculty of Agricul- Conservation ture, Hokkaido University 55: 429–458. Abe, H. 1974. Change of the boundary-line of two mole’s distribu- In Japan, M. imaizumii and M. wogura are under low tions in a period of 14 years. Journal of Mammalogical Society of Japan 6: 13–23 (in Japanese with English summary). risk of extinction, at least in the level of species. M. Abe, H. 1985. Changing mole distributions in Japan. In (T. Kawamichi, etigo in the Echigo Plain and some isolated populations ed.) Contemporary Mammalogy in China and Japan. Pp. 108– of M. imaizumii in the western Japan are affected by 112. Mammalogical Society of Japan, Tokyo. Abe, H. 1995. Revision of the Asian moles of the genus Mogera. other dominant species and decreasing their areas (Abe Journal of Mammalogical Society of Japan 20: 51–58. 2001b). The distribution areas of M. etigo and M. Abe, H. 1997. Habitat factors affecting the geographic size variation tokudae are very much limited, and Ministry of the Envi- of Japanese moles. Journal of Mammalogical Society of Japan 21: 71–87. ronment of Japan (2002) assigned these two species to Abe, H. 2001a. Soil hardness, a factor affecting the range expansion Critically endangered and Endangered, respectively. E. of Mogera wogura in Japan. Journal of Mammalogical Society of mizura are rare species, and it may be susceptive under Japan 26: 45–52. Abe, H. 2001b. Isolated relic populations and their keeping mecha- local extinction with habitat destruction and competition nisms in moles. Honyurui Kagaku [Mammalian Science] 41: 35– with other mole species in some areas in Japan. Many 52 (in Japanese with English summary). prefectural red lists in Japan appoint this species Vulner- Abe, H. (ed.) 2005. A Guide to the Mammals of Japan. Revised edi- tion. Tokai University Press, Hatano. able or Rare. Abe, H., Shiraishi, S. and Arai, S. 1991. A new mole from Uotsuri- M. uchidai is severely under the risk of extinction, jima, the Ryukyu Islands. Journal of Mammalogical Society of because of goats, Capra aegarus, introduced in 1978 into Japan 15: 47–60. Biodiversity Center of Japan. 2002. The National Survey on the Natu- Uotsuri-jima in Senkaku Islands, Okinawa Prefecture, ral Environment Report of the Distributional Survey of Japanese the only one distribution area of this species. This island Animals (Mammals). Biodiversity Center of Japan, Fujiyoshida is very small (3.8 km2), but the goats have increased to (in Japanese). Duszynski, D. W. and Wattam, A. R. 1988. Coccidian parasites more than 300 (Yokohata and Yokota 2000). The eco- (Apicomplexa: Eimeriidae) from insectivores V. ten forms from system of this island is obviously damaged, and some the moles of Japan (Euroscaptor, Mogera spp.). Journal of Proto- Yokohata, Brief review on moles in Japan S29

zoology 35: 55–57. 47–51 (in Japanese with English summary). Environmental Ministry of Japan. 2002. Threatened Wildlife of Japan. Kawada, S., Harada, M., Obara, Y., Kobayashi, S., Koyasu, K. and Red Data Book 2nd ed. Japan Wildlife Research Center, Tokyo Oda, S. 2001. Karyosystematic analysis of Japanese talpine (in Japanese with English summary). moles in the genera Euroscaptor and Mogera (Insectivora, Fujiwara, M. 1955. [A Japanese mountain mole was captured.] Jour- Talpidae). Zoological Science 18: 1003–1010. nal of Hiba Society of Natural History 35: 14 (in Japanese). Kawaguchi, S. 2004. A trapping method by tube traps and observation Fujiwara, M. 1956a. [Does mole eat plants?] Journal of Hiba Society of reproductive organs of Mogera wogura. Honyurui Kagaku of Natural History 42: 15–16 (in Japanese). [Mammalian Science] 43: 121–126 (in Japanese). Fujiwara, M. 1956b. [On the nest of moles.] Journal of Hiba Society Kushihashi, H. 1982. Occurrence of two species in Mogera from the of Natural History 41: 16 (in Japanese). Is. of Shodo-shima, Kagawa Prefecture, Japan. Kagawa Seibutsu Fujiwara, M. 1957. Notes on micrura minor Kuroda. Journal 10: 43–51 (in Japanese). of Mammalian Society of Japan 1: 65 (in Japanese). Makita Y. 1980. Notes on Mogera kobeae (4). Journal of the Mam- Fujiwara, M. 1960a. [White Mogera wogura imaizumii was cap- malogical Society of Japan 8: 113–116 (in Japanese with English tured.] Journal of Hiba Society of Natural History 54: 19 (in summary). Japanese). Ministry of the Environment of Japan. 2002. Threatened wildlife of Fujiwara, M. 1960b. On the albino of Talpa wogura kobeae. Collect- Japan. — Red Data Book 2nd ed. —. Japan Wildlife Research ing and Breeding 22: 216 (in Japanese). Center, Tokyo (in Japanese with English summary). Fujiwara, M. 1960c. The seasonal change of testes in two species of Motokawa, M. and Abe, H. 1996. On the specific names of the Japa- Japanese moles (primary report). Miscellaneous Report of Hiwa nese moles of the genus Mogera. Mammal Study 21: 115–123. Museum for Natural History 3: 18–23 (in Japanese). Motokawa, M., Koyasu, K., Yokohata, Y. and Abe, H. 1999. Bibli- Fujiwara, M. 1961. The albino and the partial albino of some animals. ography of the Japanese Insectivora. In (Y. Yokohata and S. Miscellaneous Report of Hiwa Museum for Natural History 4: Nakamura, eds.) Recent Advances in the Biology of Japanese 28–29 (in Japanese). Insectivora — Proceedings on the Symposium on the Biology of Fujiwara, M. 1962. Some examples of pregnant female and the breed- Insectivores in Japan and on the Wildlife Conservation. Pp. 105– ing season of Japanese mole, Mogera kobeae kobeae Thomas, in 150. Hiba Society of Natural History and Hiwa Museum for Hiroshima Prefecture. Miscellaneous Report of Hiwa Museum Natural History, Shobara and Hiwa. for Natural History 5: 28–29 (in Japanese). Motokawa, M., Lin, L.-K. Cheng, H.-C. and Harada, M. 2001. Taxo- Hashimoto, T. and Abe, M. 2001. Body size and reproductive sched- nomic status of the Senkaku mole, Nesoscaptor uchidai, with ules in two parapatric moles, Mogera tokudae and Mogera special reference to variation in Mogera insularis from Taiwan imaizumii, in the Etigo Plain. Journal of Mammalogical Society (Mammalia: Insectivora). Zoological Science 18: 733–740. of Japan 26: 35–44. Okinawa Prefectural Government. 2005. Threatened wildlife in Hikone City Government. 2005. Red Data Book Hikone. Hikone City Okinawa 2nd ed. (Animals) — Red Data Okinawa —. Nature Government, Hikone. Conservation Division, Department of Cultural and Environmen- Horikoshi, I., Imaizumi, Y.-N. and Tsuchiya, K. 1967. On a nest of tal Affairs Okinawa Prefectural Government, Naha (in Japanese). Mogera wogura wogura at Mitsukaido-shi, Ibaragi Prefecture. Ondo, Y. and Fukuda, K. 1961. Ecological studies on the Temminck’s Journal of Mamamlogical Society of Japan 3: 137–139 (in Japa- mole, Talpa micrura kobeae. (1) an outline of ecological infor- nese). mations. Liberal Arts Journal — the Liberal Arts Department, Hoslett, S. A. and Imaizumi, Y.-H. 1966. Age structure of a Japanese Tottori University, Natural Science 12: 50–60 (in Japanese with mole population. Journal of Mamamlogical Society of Japan 2: English summary). 151–156. Sagara, N. 1978. The occurrence of fungi in association with wood Hoslett, S. A. and Imaizumi, Y.-H. 1967. Seasonal weight changes in mouse nests. Transactions of the Mycological Society of Japan a Japanese mole population. Journal of Mamamlogical Society of 19: 201–214. Japan 3: 53–36. Sagara, N. 1980. Not mouse but mole. Transactions of the Mycologi- Imaizumi, Y.-H. 1978. Hunting methods in relation to hunting situa- cal Society of Japan 21: 519. tions in Japanese shrew-mole, Urotrichus talpoides. Annotations Sagara, N. 1986. Hair types of Japanese Insectivora, with special ref- Zoologicae Japonensis 51: 245–253. erence to the presence of straight guard hairs. Journal of Mam- Imaizumi, Y.-H. 1979. Hunting methods in relation to hunting situa- malogical Society of Japan 11: 57–64 (in Japanese with English tions in Japanese shrew-mole, Urotrichus talpoides. II. Detection summary). of the earthworm “Head”. Annotations Zoologicae Japonensis Sagara, N. 1995. Association of ectomycorrhizal fungi with decom- 52: 212–224. posed wastes in forest habitats: a cleaning symbiosis? Imaizumi, Y.-H. and Imaizumi, T. 1970. Interspecific relationship in Canadian Journal of Botany 73 Suppl. 1: S1423–S1433. two mole species in the plains of Niigata, Honshu. 1. Geographic Sagara, N. 1998. Methods for studying the nesting ecology of moles distribution. Journal of Mammalogical Society of Japan 5: 15–18 through observation of mushroom fruiting. Honyurui Kagaku (in Japanese with English summary). [Mammalian Science] 38: 271–292 (in Japanese with English Imaizumi, Y.-N. 1955. Taxonomic studies on the Japanese mountain summary). mole (Talpa mizura), with descriptions of two new subspecies. Sagara, N. 1999. Mycological approach to the natural history of talpid Bulletin of the National Science Museum, Tokyo, 1: 26–38. moles — A review with new data and proposal of “habitat-clean- Imaizumi, Y.-N. 1960. Coloured illustrations of the mammals of ing symbiosis”. In (Y. Yokohata and S. Nakamura, eds.) Recent Japan. Hoikusha, Osaka, 196 p. + 68 pls (in Japanese). Advances in the Biology of Japanese Insectivora — Proceedings Imaizumi, Y.-N. and Yoshiyuki, M. 1960. [On the albinism of of the Symposium on the Biology of Insectivores in Japan and on talpids]. Shizenkagaku to Hakubutsukan [Natural Science and the Wildlife Conservation. Pp. 33–55. Hiba Society of Natural Museum] 27: 20–28 (in Japanese). History and Hiwa Museum for Natural History, Shobara and Kanaizuka, T., Shiramizu, T. and Hoshino, K. 1991. Some notes on Hiwa. Japanese mountain moles (Euroscaptor mizura) in Hiroshima. Sagara, N. and Abe, H. 1993. A case of late breeding in the mole Miscellaneous Report of Hiwa Museum for Natural History 29: Mogera kobeae and its nest. Journal of Mammalogical Society of S30 Mammal Study 30 (2005)

Japan 18: 53–59. Faculty of Education, Toyama University, Series B. 49: 47–54 Sagara, N., Honda, S., Kuroyanagi, E. and Takayama, S. 1981. The (in Japanese with English summary). occurrence of Hebeloma spoliatum and Hebeloma radicosum Yokohata, Y. 1998. [The ecology of Talpidae.] In (H. Abe and Y. on the dung-deposited burrows of Urotrichus talpoides (shrew Yokohata, eds.) The Natural History of Insectivora (Mammalia) mole). Transactions of the Mycological Society of Japan 22: 441– in Japan. Pp. 67–187. Hiba Society of Natural History, Shobara 455. (in Japanese). Sagara, N., Kobayashi, S., Ota, H., Toshiaki, I. and Okabe, H. 1989. Yokohata, Y. 1999. A case of estimation of species of a fossil of the Finding Euroscaptor mizura (Mammalia: Insectivora) and its nest genus Mogera from Japan using discriminant functions. Memoirs from under Hebeloma radicosum (Fungi: Agaricales) in Ashiu, of the Faculty of Education, Toyama University 53: 37–44 (in Kyoto, with data of possible contiguous occurrences of three Japanese with English summary). talpine species in this region. Contributions from the Biological Yokohata, Y. 2000. Age structure of the lesser Japanese moles Laboratory, Kyoto University 27: 261–272. Mogera imaizumii from Sagamihara Golf Club in Kanagawa Sagara, N., Abe, H. and Okabe, H. 1993a. The persistence of moles Prefecture, Japan. Memoirs of the Faculty of Education, Toyama in nesting at the same site as indicated by mushroom fruiting University 54: 161–169 (in Japanese with English summary). and nest reconstruction. Canadian Journal of Zoology 71: 1690– Yokohata, Y. 2003. The problem of feral goats on Uotsuri-jima in the 1693. Senkaku Islands and appeals for countermeasures to resolve the Sagara, N., Okabe, H. and Kikuchi, J. 1993b. Occurrence of an agaric problem. Japanese Journal of Conservation Ecology 8: 87–96 (in fungus Hebeloma radicosum on the underground nest of wood Japanese with English summary). mouse. Transactions of the Mycological Society of Japan 34: Yokohata, Y. and Abe, H. 1989. Two new spirurid nematodes in 315–322. Japanese Moles, Mogera spp. Japanese Journal of Parasitology Sakai, T. and Hanamura, H. 1973. A morphological study on the 38: 92–99. dentition of Insectivora. II. Talpidae. Japanese Journal of Oral Yokohata, Y. and Sagara, N. 1995. Some parasitic nematodes of the Biology 15: 333–346 (in Japanese). Japanese mountain mole, Euroscaptor mizura. Memoirs of the Sakata, K. and Asakawa, M. 2003. Parastic nemtodes of Sado Faculty of Education, Toyama University, Series B. 47: 19–25. moles (Mogera tokudae) with the first geographical record of Yokohata, Y. and Onodera, A. 1997. Small mammals and parasitic Tricholinstowia talpae (Morgan, 1928) from Sado I. and a brief nematode fauna in their alimentary tracts in Nakaikemi-marsh, description of the species. Journal of Rakuno Gakuen University Turuga City, Fukui Prefecture, Japan. Memoirs of the Faculty of 27: 211–214 (in Japanese with English summary). Education, Toyama University 50: 41–46 (in Japanese with Sawada, I. and Koyasu, K. 1991. Hymenolepis mogerae sp. nov. English summary). (Cestoda: Hymenolepididae) from the large Japanese mole, Yokohata, Y. and Yokota, M. 2000. The problem of introduced goats Mogera kobeae Thomas of Aichi Prefecture. Japanese Journal on Uotsuri-jima in the Senkaku Islands. Wildlife Conservation 5: of Parasitology 40: 267–269. 1–12 (in Japanese with English summary). Shinohara, A., Campbell, K. L. and Suzuki, H. 2005. An evolutionary Yokohata, Y., Abe, H. and Kamiya, M. 1988a. Redescription view on the Japanese talpids based on nucleotide sequences. and multivariate morphometrics of Moguranema nipponicum Mammal Study 30: S19–S24. Yamaguti. Japanese Journal of Veterinary Research 36: 223–233. Shiraishi, S. and Arai, S. 1980. Terrestrial zoological survey (2) Yokohata, Y., Jiang, Y.-P., Abe, H. and Ohbayash, M. 1988b. Mainly mammals. In (Okinawa Development Agency, ed.) The Pseudoparasitism by thelastomatid nematodes in moles, Mogera Report of the Senkaku Islands Survey (Scientific Survey). Pp. spp., in Japan. Japanese Journal of Veterinary Research 36: 53– 47–86. Okinawa Development Agency, Tokyo (in Japanese). 67. Tashiro, M. 1958. On a nest of Talpa wogura. Journal of Mammalog- Yokohata, Y., Abe, H., Jiang, Y.-P. and Kamiya, M. 1989. Gastro- ical Society of Japan 1: 104 (in Japanese). intestinal helminth fauna of Japanese moles, Mogera spp. Japa- Tezuka, H. 1958. Notes on thigmotaxis of Talpa wogura wogura. nese Journal of Veterinary Research 37: 1–13. Journal of Mammalogical Society of Japan 1: 84–86 (in Japanese Yokohata, Y., Ikeda, Y., Yokota, M. and Ishizaki, H. 2003. The with English summary). effects of introduced goats on the ecosystem of Uotsuri-Jima, Tottori Prefectural Government. 2002. Threatened Wildlife of Tottori Senkaku Islands, Japan, as assessed by remote-sensing tech- Prefecture. — Red Data Book: Animals. Tottori Prefectural niques. Biosphere Conservation 5: 39–46. Government, Tottori. Yoshiyuki, M. 1986. The phylogenetic status of Mogera tokudae Tsuchiya, K. 1988. Cytotaxonomic studies of the Family Talpidae Kuroda, 1940 on the basis of body skeletons. Journal of Mamma- from Japan. Honyurui Kagaku [Mammalian Science] 28: 49–61 logical Society of Japan 19: 203–213 (in Japanese with English (in Japanese). summary). Yamaguti, S. 1941. Studies on the helminth fauna of Japan. Part 35. Yoshiyuki, M. and Imaizumi, Y.-N. 1991. Taxonomic status of the Mammalian nematodes. II. Japanese Journal of Zoology 9: 409– from the Etigo Plain, central Japan, with description of 438. a new species (Mammalia, Insectivora, Talpidae). Bulletin of Yokohata, Y. 1994. Age determination of Mogera robusta (Mam- National Science Museum, ser. A 17: 101–110. malia; Talpidae) and age structure of its population in Hiwa, Yoshiyuki, M. and Imaizumi, Y.-N. 1992. Analysis on size variations Hiroshima Prefecture, Japan. Memoirs of the Faculty of Educa- of Mogera wogura wogura (Temminck, 1843) from Sagamihara tion, Toyama University, Series B. 45: 63–74 (in Japanese with Golf Club Garden in Kanagawa Prefecture, Japan. Bulletin of the English summary). Kanagawa Prefectural Museum (Natural Science) 21: 81–98 (in Yokohata, Y. 1997a. A case of pelage-color variant in the large Japanese with English summary). Japanese mole, Mogera wogura, in Hiwa, Hiroshima Prefecture, Yukawa, M. 1968. Two mizura moles (Euroscaptor mizura Gunther) Japan. Miscellaneous Report of Hiwa Museum for Natural collected in Hiwa cho, Hiroshima Pref. Miscellaneous Report of History 35: 195–197 (in Japanese with English summary). Hiwa Museum for Natural History 12: 18–19 (in Japanese). Yokohata, Y. 1997b. Analysis of the population structure of large Japanese moles, Mogera wogura (Mammalia; Talpidae) in Hiwa, Received 29 November 2005. Accepted 27 January 2006. Hiroshima Prefecture, using cohort analysis. Memoirs of the