Seed contents of sika deer (Cervus nippon) dung and the fate of seeds in a temperate short grassland in an urban park in Japan

다국어 초록 (Multilingual Abstract)

Many studies have suggested the positive effects of grazing by large herbivorous mammals on seed dispersal, but little is known about how herbivores could affect the fate of ingested seeds. This study examined the effects of seed ingestion by sika deer (Cervus nippon) on seed fate in a temperate grassland established in an urban park long resided by high densities of sika deer. I compared species composition and seasonal traits of seed abundance and maturity in the grassland community with those in deer fecal pellets. In total, 27 herbaceous species were observed, including the predominant Zoysia japonica. Seed phenology and production differed among the three dominant species (Z. japonica, Digitaria violascens, and Hydrocotyle maritima). Pellets contained at least 26 species of herbaceous seeds, and their abundance differed among species. Of the 26 species, 15 were observed in the vegetation at the study site. The peak of seed abundance in pellets for the dominant species appeared 1 month after the peak of inflorescence production (but most of the inflorescences were immature and susceptible to digestion) and consequently corresponded to the peak of mature inflorescence. Because sika deer are likely to ingest seeds at any maturity stage in the grassland and immature seeds are less hardened, ingested immature seeds can suffer great losses. The results suggested that the survival of germable seeds with great losses of immature seeds may be a factor determining which plant species can be successfully dispersed by herbivores.

참고문헌 (Reference)

1 Kasahara Y, "Weeds of Japan Illustrated: Seeds, Seedlings and Plants. 2nd ed" Yokendo 1968

2 Numata M, "Weed flora of Japan: Illustrated in Colour" Zenkoku Noson Kyoiku Kyokai 1971

3 Takatsuki S, "The vegetation of Nara Park with reference to the grazing effects of Sika deer (Cervus nippon). In: Annual Reports on Sika Deer in Nara Park" Kasuga Kenshokai, Nara 113-131, 1980

4 Shimoda K, "The regeneration of pioneer tree species under browsing pressure of Sika deer in an evergreen oak forest" 9 : 85-92, 1994

5 Bruun HH, "The past impact of livestock husbandry on dispersal of plant seeds in the landscape of Denmark" 31 : 425-431, 2002

6 Gill RMA, "The impact of deer on woodlands: the effects of browsing and seed dispersal on vegetation structure and composition" 74 : 209-218, 2001

7 Welch D, "Studies in the grazing of heather moorland in north-east Scotland. IV. Seed dispersal and plant establishment in dung" 22 : 461-472, 1985

8 Maesako Y, "Spatial distribution of two invasive alien species, Podocarpus nagi and Sapium sebiferum, spreading in a warm-temperate evergreen forest of the Kasugayama Forest Reserve, Japan" 24 : 103-112, 2007

9 Miura S, "Sociological studies on Sika deer in Nara park with reference to individual distribution and behavior. In: Annual Reports on Sika Deer in Nara Park" Kasuga Kenshokai, Nara 3-41, 1977

10 Torii H, "Sika deer in Nara Park: unique human-wildlife relations. In: Sika Deer: Biology and Management of Native and Introduced Populations" Springer 347-363, 2009

11 Nakayama S, "Seeds of Wild Plants in Japan" Tohoku University Press 2000

12 Kitagawa N, "Seed dispersal of Zoysia japonica by sika deer in the ecosystem of Nara Park" 36 : 9-12, 1988

13 Myers JA, "Seed dispersal by white-tailed deer: implications for long-distance dispersal, invasion, and migration of plants in eastern North America" 139 : 35-44, 2004

14 Yamashiro A, "Seed dispersal by Kerama Deer (Cervus nippon keramae) on Aka Island, the Ryukyu Archipelago, Japan" 38 : 405-413, 2006

15 Takatsuki S, "Plants and plant communities on Kinkazan Island, northern Japan, in relation to sika deer herbivory. In: Sika Deer: Biology and Management of Native and Introduced Populations" Springer 125-143, 2009

16 Couvreur M, "Large herbivores as mobile links between isolated nature reserves through adhesive seed dispersal" 7 : 229-236, 2004

17 Cosyns E, "Horse dung germinable seed content in relation to plant species abundance, diet composition and seed characteristics" 6 : 11-24, 2005

18 Crawley MJ, "Herbivory: The Dynamics of Animal-Plant Interactions" University of California Press 1983

19 Malo JE, "Herbivorous mammals as seed dispersers in a Mediterranean dehesa" 104 : 246-255, 1995

20 Takatsuki S, "Food habits of Sika deer in Nara Park, assessed by fecal analysis (II). In: Annual Reports on Sika Deer in Nara Park" Kasuga Kenshokai, Nara 25-37, 1978

21 Takatsuki S, "Food habits of Sika deer in Nara Park assessed by fecal analysis (I). In: Annual Reports on Sika Deer in Nara Park" Kasuga Kenshoukai, Nara 129-141, 1977

22 Takatsuki S, "Effects of sika deer on vegetation in Japan: a review" 142 : 1922-1929, 2009

23 Takatsuki S, "Effects of sika deer on the regeneration of a Fagus crenata forest on Kinkazan Island, northern Japan" 9 : 115-120, 1994

24 Ishikawa H, "Effects of ingestion of seeds by sika deer (Cervus nippon) and dung presence on their germination in a herbaceous community" 25 : 591-598, 2010

25 Ishikawa H, "Effects of dung beetles on seedling emergence from herbaceous seeds in the dung of sika deer (Cervus nippon) in a temperate Japanese grassland ecosystem" 26 : 725-734, 2011

26 Okuda T, "Effects of deer browsing on the early stage of pyrogenic succession on Miyajima Island, Southeastern Japan" 5 : 353-366, 1990

27 Akashi N, "Effects of bark-stripping by Sika deer (Cervus nippon) on population dynamics of a mixed forest in Japan" 113 : 75-82, 1999

28 Mouissie AM, "Ecological correlates of seed survival after ingestion by Fallow Deer" 19 : 284-290, 2005

29 Pakeman RJ, "Ecological correlates of endozoochory by herbivores" 16 : 296-304, 2002

30 Tagawa H, "Ecological Lecture 11-a: Ecological Succession I" Kyoritsu Shuppan 1973

31 Takatsuki S, "Ecological History of Sika Deer" University of Tokyo Press 2006

32 Maesako Y, "Disturbance by white-whiskered boars (Sus scrofa leucomystax) on the grassland of Tobihino, Nara Park, Nara, Central Japan" 24 : 19-23, 2002

33 von Oheimb G, "Dispersal of vascular plants by game in northern Germany. Part II: Red deer (Cervus elaphus)" 124 : 55-65, 2005

34 Heinken T, "Dispersal of vascular plants by four species of wild mammals in a deciduous forest in NE Germany" 32 : 627-643, 2002

35 Janzen DH, "Dispersal of small seeds by big herbivores: foliage is the fruit" 123 : 338-353, 1984

36 Hofmann RR, "Digestive physiology of the deer: their morphophysiological specialization and adaptation" 22 : 393-407, 1985

37 Couvreur M, "Complementarity of epi- and endozoochory of plant seeds by free ranging donkeys" 28 : 37-48, 2005

38 Grime JP, "Comparative Plant Ecology: A Functional Approach to Common British Species" Unwin Hyman 1988

39 Japan Meteorological Agency, "Climate statistics in Japan"

40 Fukunaga H, "Behavior of sika deer (Cervus nippon) in Nara Park (I). Habitat use and daily activity. In: Annual Reports on Sika Deer in Nara Park" Kasuga Kenshokai, Nara 3-13, 1975

41 Fukunaga H, "Behavior of sika deer (Cervus nippon) in Nara Park (I). Daily activity and habitat use (2). In: Annual Reports on Sika Deer in Nara Park" Kasuga Kenshokai, Nara 63-69, 1976

42 Foundation for the Protection of Deer in Nara Park, "Assessment of total population of sika deer in Nara Park, Japan"

43 Miura S, "Artiodactyla. In: A Guide to the Mammals of Japan" Tokai University Press 107-114, 2005