Cookies Notification

We use cookies to improve your website experience. To learn about our use of cookies and how you can manage your cookie settings, please see our Cookie Policy.

Overlap in habitat use and activity patterns between sika deer (Cervus nippon) and Japanese serows (Capricornis crispus) in subalpine habitats: exploitative competition rather than direct interference?

Publication: Canadian Journal of Zoology
12 July 2023


Examining multiple niche relationships, such as habitat use and activity patterns, between sympatric species contributes to an understanding of the mechanisms of coexistence and competition. Sika deer (Cervus nippon Temminck, 1838) and Japanese serows (Capricornis crispus (Temminck, 1836)) often show different habitat use, and diversity in vegetation and topography seems to facilitate their coexistence. Conversely, their habitat use may overlap in areas with low-diversity habitats; however, this remains unstudied. Moreover, whether they spatiotemporally avoid each other to reduce direct interference is unclear. We examined the fine-scale patterns of habitat use and activity of deer and serows in the subalpine forests of Mount Fuji, which have low-diversity habitats, using camera trapping data over 3 years. Deer and serow habitat niches largely overlapped, especially in summer (92%–94%) and autumn (89%–91%), suggesting that low-diversity habitats facilitate overlapping habitat use. Both species selected areas frequently used by the other during spring to autumn and their temporal niches largely overlapped, especially in summer (88%) and autumn (83%), suggesting that they do not spatiotemporally avoid each other (i.e., there is no direct interference). Rapid range expansion of deer into these subalpine habitats may exclude native serows through resource exploitative competition rather than interference competition.

Get full access to this article

View all available purchase options and get full access to this article.


Ariefiandy A., Purwandana D., Azmi M., Panggur M.R., Mardani J., Parra D.P., Jessop T.S. 2021. Invasive water buffalo population trends and competition-related consequences for native rusa deer in eastern Indonesian protected areas. Mamm. Biol. 101(6): 917–931.
Bagchi S., Goyal S.P., Sankar K. 2003. Niche relationships of an ungulate assemblage in a dry tropical forest. J. Mammal. 84(3): 981–988.
Bartón K. 2019. MuMIn: multi-model inference. R package version 1.43.6. Available from [accessed May 2023].
Bates D., Maechler M., Bolker B., Walker S. 2015. Fitting linear mixed effects models using lme4. J. Stat. Softw. 67: 1–48.
Belovsky G.E. 1986. Generalist herbivore foraging and its role in competitive interactions. Am. Zool. 26(1): 51–69.
Burnham K.P., Anderson D. 2002. Model selection and multimodel inference. Springer, New York.
Carothers J.H., Jaksić F.M. 1984. Time as a niche difference: the role of interference competition. Oikos, 42(3): 403–406.
Chen Y., Xiao Z., Zhang L., Wang X., Li M., Xiang Z. 2019. Activity rhythms of coexisting red serow and Chinese serow at Mt. Gaoligong as identified by camera traps. Animals, 9: 1071.
Chirichella R., Ciuti S., Apollonio M. 2013. Effects of livestock and non-native mouflon on use of high-elevation pastures by Alpine chamois. Mamm. Biol. 78(5): 344–350.
Connor E.F., Simberloff D. 1979. The assembly of species communities: chance or competition? Ecology, 60(6): 1132–1140.
Côté S.D., Festa-Bianchet M. 2001. Birthdate, mass and survival in mountain goat kids: effects of maternal characteristics and forage quality. Oecologia, 127: 230–238.
Ferretti F., Sforzi A., Lovari S. 2012. Avoidance of fallow deer by roe deer may not be habitat-dependent. Hystrix, 23(2): 28–34.
Gause G.F. 1934. The struggle for existence. Williams and Wilkins, Philadelphia
Gordon I.J., Illius A.W. 1989. Resource partitioning by ungulates on the Isle of Rhum. Oecologia, 79(3): 383–389.
Hairston N.G. 1980. The experimental test of an analysis of field distributions: competition in terrestrial salamanders. Ecology, 61(4): 817–826.
Hashimoto Y., Mori Y. 2017. Transition in the distribution of the Japanese serow in the Suzuka Mountains Japanese Serow Protection Area. Wildl. Hum. Soc. 5(2): 1–8.
Hiruma M., Takada H., Washida A., Koike K. 2023. Dietary partitioning and competition between sika deer and Japanese serows in high elevation habitats. Mamm. Res.
Hutchinson G.E. 1957. Concluding remarks. Cold Spring Harb. Symp. Quant. Biol. 22: 415–427.
Ikeda T., Nakamori S., Ando M., Shirakawa T., Okamoto T., Suzuki M. 2021. Seasonal diel activity patterns of three sympatric ungulates in forested area in central Japan. Mamm. Study, 47(1): 47–56.
Ikeda T., Takahashi H., Igota H., Matsuura Y., Azumaya M., Yoshida T., Kaji K. 2019. Effects of culling intensity on diel and seasonal activity patterns of sika deer (Cervus nippon). Sci. Rep. 9(1): 17205.
Ikeda T., Takahashi H., Yoshida T., Igota H., Matsuura Y., Takeshita K., Kaji K. 2015. Seasonal variation of activity pattern in sika deer (Cervus nippon) as assessed by camera trap survey. Mamm. Study, 40(4): 199–205.
Jenkins K.J., Wright R.G. 1988. Resource partitioning and competition among cervids in the northern Rocky Mountains. J. Appl. Ecol. 25(1): 11–24.
Jiang Z., Torii H., Takatsuki S., Ohba T. 2008. Local variation in diet composition of the Japanese serow during winter. Zool. Sci. 25(12): 1220–1226.
Latham J. 1999. Interspecific interactions of ungulates in European forests: an overview. For. Ecol. Manag. 120(1–3): 13–21.
Lysy M., Stasko A., Swanson H. 2021. nicheROVER: niche region and niche overlap metrics for multidimensional ecological niches. R package version 1.1.0. Available from [accessed May 2023].
Minami M. 2008. Sika deer: reproductive success. In Mammalogy in Japan 2 (middle-, and large sized mammals including primates). Edited by Takatsuki S., Yamagiwa J. University of Tokyo Press, Tokyo. pp. 123–148  [In Japanese].
Namgail T., Fox J.L., Bhatnagar Y.V. 2004. Habitat segregation between sympatric Tibetan argali Ovis ammon hodgsoni and blue sheep Pseudois nayaur in the Indian Trans-Himalaya. J. Zool. 262(1): 57–63.
Nawa A. 2009. Mori no Kenjya Kamoshika. Sun-Rise Press, Hikone  [In Japanese].
Ochiai K. 1999. Diet of the Japanese serow (Capricornis crispus) on the Shimokita Peninsula, northern Japan, in reference to variations with a 16-year interval. Mamm. Study, 24(2): 91–102.
Oka T., Iijima H., Kamata A., Ishida A., Eguchi N., Aikawa T., et al. 2022. The process of population expansion of sika deer. In Sika deer: life history plasticity and management. Edited by Kaji K., Uno H., Iijima H., Springer, Singapore. pp. 11–23.
Pacala S., Roughgarden J. 1982. Resource partitioning and interspecific competition in two two-species insular Anolis lizard communities. Science, 217(4558): 444–446.
Putman R.J. 1996. Competition and resource partitioning in temperate ungulate assemblies. Chapman & Hall, London.
R Core Team. 2022. R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Available from [accessed May 2023].
Ridout M., Linkiem M. 2009. Estimating overlap of daily activity patterns from camera trap data. J. Agric. Biol. Environ. Stat. 14(3): 322–337.
Rowcliffe M. 2023. activity: Animal Activity Statistics. R package version 1.3.3. Available from [accessed May 2023].
Schoener T.W. 1974. Resource partitioning in ecological communities: research on how similar species divide resources helps reveal the natural regulation of species diversity. Science, 185(4145): 27–39.
Seki Y., Hayama S. 2021. Habitat selection and activity patterns of Japanese serows and sika deer with currently sympatric distributions. Animals, 11(12): 3398.
Takada H. 2020. The summer spatial distribution of Japanese serows (Capricornis crispus) in an area without predation risk. Mamm. Biol. 100(1): 63–71.
Takada H. 2022. Interspecific relationships and behavioral traits of three sympatric herbivores, Japanese serows, sika deer, and Japanese hares, in the northern slope of Mt. Fuji. In Mount Fuji Research Institute Research Report 18. Mount Fuji Research Institute, Yamanashi, Japan. pp 1–23. Available from [accessed May 2023].
Takada H. 2023. Unique spatial behavior of the Japanese serow (Capricornis crispus) in the open mountains of Mt. Fuji. Acta Ethol. .
Takada H., Minami M. 2019. Do differences in ecological conditions influence grouping behaviour in a solitary ungulate, the Japanese serow?. Behaviour 156: 245–264.
Takada H., Minami M. 2021. A preliminary study on habitat selection of the Japanese serow (Capricornis crispus) at two temporal scales, season and time of day, in a montane forest. J. Ethol. 40(1): 91–95.
Takada H., Washida A. 2020. Ecological drivers of group size variation in sika deer: habitat structure, population density, or both? Mamm. Biol. 100(5): 445–452.
Takada H., Hiruma M., Washida A., Katsumata E. 2020b. Present status of Japanese serow (Capricornis crispus) and sika deer (Cervus nippon) in the alpine habitat of Mt. Fuji. Mt Fuji Res. 14: 1–10.
Takada H., Nakamura K., Minami M. 2019. Effects of the physical and social environment on flight response and habitat use in a solitary ungulate, the Japanese serow (Capricornis crispus). Behav. Processes, 158: 228–233.
Takada H., Nakamura K., Takatsuki S., Minami M. 2018. Freezing behavior of the Japanese serow (Capricornis crispus) in relation to habitat and group size. Mamm. Res. 63(1): 107–112.
Takada H., Ohuchi R., Watanabe H., Yano R., Miyaoka R., Nakagawa T., et al. 2020a. Habitat use and the coexistence of the sika deer and the Japanese serow, sympatric ungulates from Mt Asama, central Japan. Mammalia, 84(6): 503–511.
Takada H., Yano R., Katsumata A., Takatsuki S., Minami M. 2021. Diet compositions of two sympatric ungulates, the Japanese serow (Capricornis crispus) and the sika deer (Cervus nippon), in a montane forest and an alpine grassland of Mt Asama, central Japan. Mamm. Biol. 101(5): 681–694.
Takatsuki S. 2009. Geographical variations in food habits of sika deer: the northern grazer vs. the southern browser. In Sika deer. Edited by McCullough D.R., Takatsuki S., Kaji K. Springer, New York. pp. 231–237.
Takatsuki S., Kobayashi-Hori Y., Ito T. 1995. Food habits of Japanese serow (Capricornis crispus) in the western foothills of Mt. Zao, with reference to snow cover. J. Mamm. Soc. Jpn. 20(2): 151–155.
van Doormaal N., Ohashi H., Koike S., Kaji K. 2015. Influence of human activities on the activity patterns of Japanese sika deer (Cervus nippon) and wild boar (Sus scrofa) in Central Japan. Eur. J. Wildl. Res. 61(4): 517–527.
Voeten M.M., Prins H.H. 1999. Resource partitioning between sympatric wild and domestic herbivores in the Tarangire region of Tanzania. Oecologia, 120(2): 287–294.
Wilson D.E., Mittermeier R.A. 2011. Hand book of the mammals of the world: 2. Hoofed mammals. Lynx, Barcelona.
Wrobell D.J., Gergits W.F., Jaeger R.G. 1980. An experimental study of interference competition. among terrestrial salamanders. Ecology, 61(5): 1034–1039.
Yamashiro A., Kaneshiro Y., Kawaguchi Y., Yamashiro T. 2019. Dietary overlap but spatial gap between sympatric Japanese serow (Capricornis crispus) and sika deer (Cervus nippon) on Eastern Shikoku, Japan. Mamm. Study, 44(4): 261–267.

Supplementary material

Supplementary Material 1 (XLSX / 42.9 KB).
Supplementary Material 2 (XLSX / 222 KB).

Information & Authors


Published In

cover image Canadian Journal of Zoology
Canadian Journal of Zoology
Volume 101Number 11November 2023
Pages: 980 - 990


Received: 21 January 2023
Accepted: 23 May 2023
Accepted manuscript online: 9 June 2023
Version of record online: 12 July 2023

Data Availability Statement

All data generated or analyzed during this study are included in this published article (and its supplementary files).


Request permissions for this article.

Key Words

  1. activity pattern
  2. Capricornis crispus
  3. Cervus nippon
  4. competition
  5. habitat use
  6. sika deer
  7. Japanese serow



Wildlife Management Center, Tokyo University of Agriculture and Technology, 3-5-8, Saiwai-Cho, Fuchu, Tokyo 183-8509, Japan
Mount Fuji Research Institute, Yamanashi Prefecture Government, 5597-1 Kenmarubi, Kamiyoshida, Fujiyoshida, Yamanashi 403-0005, Japan
Author Contributions: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Project administration, Resources, Software, Supervision, Validation, Visualization, and Writing – original draft.
Keita Nakamura
Mount Fuji Research Institute, Yamanashi Prefecture Government, 5597-1 Kenmarubi, Kamiyoshida, Fujiyoshida, Yamanashi 403-0005, Japan
Author Contributions: Data curation, Investigation, Methodology, Resources, and Writing – review & editing.

Author Contributions

Conceptualization: HT
Data curation: HT, KN
Formal analysis: HT
Investigation: HT, KN
Methodology: HT, KN
Project administration: HT
Resources: HT, KN
Software: HT
Supervision: HT
Validation: HT
Visualization: HT
Writing – original draft: HT
Writing – review & editing: KN

Competing Interests

The authors declare there are no competing interests.

Funding Information

The authors declare no specific funding for this work.

Metrics & Citations


Other Metrics


Cite As

Export Citations

If you have the appropriate software installed, you can download article citation data to the citation manager of your choice. Simply select your manager software from the list below and click Download.

There are no citations for this item

View Options

Get Access

Login options

Check if you access through your login credentials or your institution to get full access on this article.


Click on the button below to subscribe to Canadian Journal of Zoology

Purchase options

Purchase this article to get full access to it.

Restore your content access

Enter your email address to restore your content access:

Note: This functionality works only for purchases done as a guest. If you already have an account, log in to access the content to which you are entitled.

View options


View PDF

Full Text

View Full Text





Share Options


Share the article link

Share on social media