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RESEARCH ARTICLE (Open Access)

Seasonal and daily activity of non-native sambar deer in and around high-elevation peatlands, south-eastern Australia

Sebastien Comte https://orcid.org/0000-0001-7984-8159 A * , Elaine Thomas B , Andrew J. Bengsen https://orcid.org/0000-0003-2205-4416 A , Ami Bennett https://orcid.org/0000-0002-1908-1475 C , Naomi E. Davis https://orcid.org/0000-0002-5551-8822 C D , Sean Freney A G , Stephen M. Jackson https://orcid.org/0000-0002-7252-0799 A H , Matt White E , David M. Forsyth https://orcid.org/0000-0001-5356-9573 A and Daniel Brown F
+ Author Affiliations
- Author Affiliations

A Vertebrate Pest Research Unit, NSW Department of Primary Industries, 1447 Forest Road, Orange, NSW 2800, Australia.

B Parks Victoria, Mt Beauty, Vic. 3699, Australia.

C School of BioSciences, The University of Melbourne, Melbourne, Vic. 3010, Australia.

D Parks Victoria, 535 Bourke Street, Melbourne, Vic. 3000, Australia.

E Department of Environment, Land, Water and Planning, Arthur Rylah Institute for Environmental Research, 123 Brown Street, Heidelberg, Vic. 3084, Australia.

F Parks Victoria, Bright, Vic. 3741, Australia.

G Present address: North Coast Local Land Services, 24–26 Mulgi Drive, South Grafton, NSW 2460, Australia.

H Present address: Australian Museum Research Institute, 1 William Street, Sydney, NSW 2010, Australia.

* Correspondence to: sebastien.comte@dpi.nsw.gov.au

Handling Editor: Andrea Taylor

Wildlife Research 49(7) 659-672 https://doi.org/10.1071/WR21147
Submitted: 15 October 2021  Accepted: 7 March 2022   Published: 26 May 2022

© 2022 The Author(s) (or their employer(s)). Published by CSIRO Publishing. This is an open access article distributed under the Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License (CC BY-NC-ND)

Abstract

Context: Of the six species of non-native deer present in Australia, the sambar deer is the largest and has been identified as a major threat to high-elevation peatlands in south-eastern Australia. However, little is known about sambar deer activity in high-elevation peatlands.

Aims: The aims of this study were to quantify sambar deer activity (including wallowing) seasonally and daily in response to biotic and abiotic variables, and how activity was impacted by ground-based shooting.

Methods: To estimate sambar deer activity, camera traps were continuously deployed for 4 years in two ~4300-ha areas in Alpine National Park, Victoria, south-eastern Australia. One area was subject to management operations using ground-based shooting to target deer and the other was not. Monthly activity of sambar deer was modelled using biotic (woody vegetation cover), abiotic (snow depth, aspect, slope, distance to water, road and peatland) and management (treatment versus non-treatment) covariates. Additional camera traps were deployed to monitor sambar deer activity at wallows.

Key results: Sambar deer activity decreased when snow depth increased (between July and September), and was highest in easterly and northerly aspects with dense woody vegetation close to high-elevation peatlands and roads. During our 4-year study, sambar deer activity decreased in the treatment area but increased in the non-treatment area. Sambar deer exhibited a crepuscular diel cycle, with greatest activity around sunset. Only male sambar deer were observed to wallow, with most wallowing occurring in the afternoon during October–June.

Conclusions: Sambar deer utilised high-elevation peatlands during October–June. Daily activity was crepuscular and was greatest in dense tree cover close to roads. Ground-based shooting reduced sambar deer activity in and around high-elevation peatlands.

Implications: Control operations targeting sambar deer at high elevations in south-eastern Australia should be conducted during October–June. Outside this period sambar deer appear to use lower-elevation habitats. The effectiveness of ground-based shooting could be improved by focusing this control action around sunset (when sambar deer are most active) and in places with dense vegetation close to roads and high-elevation peatlands.

Keywords: Alpine National Park, biological invasions, camera trap, Cervus unicolor, diel cycle, invasive species, population dynamics, ungulates, wallowing.


References

Agostinelli C, Lund U (2017) R package ‘circular’: Circular statistics. version 0.4-93. Available at https://CRAN.R-project.org/package=circular [verified 22 September 2021].

Allen, RB, Forsyth, DM, Allen, RKJ, Affeld, K, and MacKenzie, DI (2015). Solar radiation determines site occupancy of coexisting tropical and temperate deer species introduced to New Zealand forests. PLoS One 10, e0128924.
Solar radiation determines site occupancy of coexisting tropical and temperate deer species introduced to New Zealand forests.Crossref | GoogleScholarGoogle Scholar | 26061426PubMed |

Asher, GW (2011). Reproductive cycles of deer. Animal Reproduction Science 124, 170–175.
Reproductive cycles of deer.Crossref | GoogleScholarGoogle Scholar | 20884138PubMed |

Bengsen, AJ, Forsyth, DM, Harris, S, Latham, ADM, McLeod, SR, and Pople, A (2020). A systematic review of ground-based shooting to control overabundant mammal populations. Wildlife Research 47, 197–207.
A systematic review of ground-based shooting to control overabundant mammal populations.Crossref | GoogleScholarGoogle Scholar |

Bengsen, AJ, Forsyth, DM, Ramsey, DSL, Amos, M, Brennan, M, Pople, AR, Comte, S, and Crittle, T (2022). Estimating deer density and abundance using spatial mark-resight models with camera trap data. Journal of Mammalogy , gyac016.
Estimating deer density and abundance using spatial mark-resight models with camera trap data.Crossref | GoogleScholarGoogle Scholar |

Benhaiem, S, Delon, M, Lourtet, B, Cargnelutti, B, Aulagnier, S, Hewison, AJM, Morellet, N, and Verheyden, H (2008). Hunting increases vigilance levels in roe deer and modifies feeding site selection. Animal Behaviour 76, 611–618.
Hunting increases vigilance levels in roe deer and modifies feeding site selection.Crossref | GoogleScholarGoogle Scholar |

Bennett, A, and Coulson, G (2010). The impacts of sambar cervus unicolor on the threatened shiny nematolepis Nematolepis wilsonii. Pacific Conservation Biology 16, 251–260.
The impacts of sambar cervus unicolor on the threatened shiny nematolepis Nematolepis wilsonii.Crossref | GoogleScholarGoogle Scholar |

Bennett, A, Haydon, S, Stevens, M, and Coulson, G (2015). Culling reduces fecal pellet deposition by introduced sambar (Rusa unicolor) in a protected water catchment. Wildlife Society Bulletin 39, 268–275.
Culling reduces fecal pellet deposition by introduced sambar (Rusa unicolor) in a protected water catchment.Crossref | GoogleScholarGoogle Scholar |

Bentley A (1998) ‘An introduction to the deer of Australia, with special reference to Victoria.’ (The Australian Deer Research Foundation Ltd: Melbourne)

Bilney, RJ (2013). Antler rubbing of yellow-wood by sambar in east Gippsland, Victoria. The Victorian Naturalist 130, 68–74.

Bischof, R, Ali, H, Kabir, M, Hameed, S, and Nawaz, MA (2014). Being the underdog: an elusive small carnivore uses space with prey and time without enemies. Journal of Zoology 293, 40–48.
Being the underdog: an elusive small carnivore uses space with prey and time without enemies.Crossref | GoogleScholarGoogle Scholar |

Bivand R, Rundel C (2020) Rgeos: Interface to geometry engine - open source (‘geos’). R package version 0.5-3. Available at https://CRAN.R-project.org/package=rgeos [verified 22 September 2021]

Bivand R, Keitt T, Rowlingson B (2020) Rgdal: Bindings for the ‘geospatial’ data abstraction library. R package version 1.5-10. Available at https://CRAN.R-project.org/package=rgdal [verified 22 September 2021]

Brodie, JF, and Brockelman, WY (2009). Bed site selection of red muntjac (Muntiacus muntjak) and sambar (Rusa unicolor) in a tropical seasonal forest. Ecological Research 24, 1251–1256.
Bed site selection of red muntjac (Muntiacus muntjak) and sambar (Rusa unicolor) in a tropical seasonal forest.Crossref | GoogleScholarGoogle Scholar |

Cairns S, Robertson G (2014) ‘Feral horses in the Australian alps national parks: The design and analysis of surveys conducted in April–may 2014.’ (Australian Alps Liaison Committee)

Chalmers PRS (2018) ‘New Zealand’s sambar and rusa deer.’ (Philip R. S. Chalmers: Whakatane)

Coe, PK, Clark, DA, Nielson, RM, Gregory, SC, Cupples, JB, Hedrick, MJ, Johnson, BK, and Jackson, DH (2018). Multiscale models of habitat use by mule deer in winter. The Journal of Wildlife Management 82, 1285–1299.
Multiscale models of habitat use by mule deer in winter.Crossref | GoogleScholarGoogle Scholar |

Conn BJ (1993) Natural regions and vegetation of Victoria. In ‘Flora of Victoria’. Vol. 1. pp. 79–158. (Inkata Press)

Côté, SD, Rooney, TP, Tremblay, J-P, Dussault, C, and Waller, DM (2004). Ecological impacts of deer overabundance. Annual Review of Ecology, Evolution, and Systematics 35, 113–147.
Ecological impacts of deer overabundance.Crossref | GoogleScholarGoogle Scholar |

Cromsigt, JPGM, Kuijper, DPJ, Adam, M, Beschta, RL, Churski, M, Eycott, A, Kerley, GIH, Mysterud, A, Schmidt, K, and West, K (2013). Hunting for fear: Innovating management of human–wildlife conflicts. Journal of Applied Ecology 50, 544–549.
Hunting for fear: Innovating management of human–wildlife conflicts.Crossref | GoogleScholarGoogle Scholar |

Dahlan, I, and Dawend, J (2013). Growth and reproductive performance of sambar deer in Sabal forest reserve of Sarawak, Malaysia. Tropical Animal Health and Production 45, 1469–1476.
Growth and reproductive performance of sambar deer in Sabal forest reserve of Sarawak, Malaysia.Crossref | GoogleScholarGoogle Scholar | 23475732PubMed |

Davies, C, Wright, W, Hogan, FE, and Davies, H (2020). Detectability and activity patterns of sambar deer (Rusa unicolor) in Baw Baw National Park, Victoria. Australian Mammalogy 42, 312–320.
Detectability and activity patterns of sambar deer (Rusa unicolor) in Baw Baw National Park, Victoria.Crossref | GoogleScholarGoogle Scholar |

Davis NE, Bennett A, Forsyth DM (2015a) ‘Monitoring changes in deer density/abundance and habitat use associated with the Parks Victoria deer control trial in the Alpine National Park: Field survey manual.’ Report prepared for Parks Victoria, Melbourne. p. 51.

Davis NE, Bennett A, Forsyth DM (2015b) ‘Monitoring changes in deer density/abundance and habitat use associated with the Parks Victoria deer control trial in the Alpine National Park: Survey design and rationale.’ Report prepared for Parks Victoria, Melbourne. p. 35.

Davis, NE, Bennett, A, Forsyth, DM, Bowman, DMJS, Lefroy, EC, Wood, SW, Woolnough, AP, West, P, Hampton, JO, and Johnson, CN (2016). A systematic review of the impacts and management of introduced deer (family Cervidae) in Australia. Wildlife Research 43, 515–532.
A systematic review of the impacts and management of introduced deer (family Cervidae) in Australia.Crossref | GoogleScholarGoogle Scholar |

Denwood, MJ (2016). Runjags: An r package providing interface utilities, model templates, parallel computing methods and additional distributions for mcmc models in jags. Journal of Statistical Software 71, 1–25.
Runjags: An r package providing interface utilities, model templates, parallel computing methods and additional distributions for mcmc models in jags.Crossref | GoogleScholarGoogle Scholar |

Diagne, C, Leroy, B, Vaissière, A-C, Gozlan, RE, Roiz, D, Jarić, I, Salles, J-M, Bradshaw, CJA, and Courchamp, F (2021). High and rising economic costs of biological invasions worldwide. Nature 592, 571–576.
High and rising economic costs of biological invasions worldwide.Crossref | GoogleScholarGoogle Scholar | 33790468PubMed |

Didham, RK, Tylianakis, JM, Hutchison, MA, Ewers, RM, and Gemmell, NJ (2005). Are invasive species the drivers of ecological change? Trends in Ecology and Evolution 20, 470–474.
Are invasive species the drivers of ecological change?Crossref | GoogleScholarGoogle Scholar | 16701420PubMed |

Dolman, PM, and Wäber, K (2008). Ecosystem and competition impacts of introduced deer. Wildlife Research 35, 202–214.
Ecosystem and competition impacts of introduced deer.Crossref | GoogleScholarGoogle Scholar |

Forman, RTT, and Deblinger, RD (2000). The ecological road-effect zone of a Massachusetts (U.S.A.) suburban highway. Conservation Biology 14, 36–46.
The ecological road-effect zone of a Massachusetts (U.S.A.) suburban highway.Crossref | GoogleScholarGoogle Scholar |

Forsyth, DM, McLeod, SR, Scroggie, MP, and White, MD (2009). Modelling the abundance of wildlife using field surveys and GIS: Non-native sambar deer (Cervus unicolor) in the Yarra Ranges, south-eastern Australia. Wildlife Research 36, 231–241.
Modelling the abundance of wildlife using field surveys and GIS: Non-native sambar deer (Cervus unicolor) in the Yarra Ranges, south-eastern Australia.Crossref | GoogleScholarGoogle Scholar |

Forsyth, DM, Woodford, L, Moloney, PD, Hampton, JO, Woolnough, AP, and Tucker, M (2014). How does a carnivore guild utilise a substantial but unpredictable anthropogenic food source? Scavenging on hunter-shot ungulate carcasses by wild dogs/dingoes, red foxes and feral cats in south-eastern Australia revealed by camera traps. PLoS One 9, e97937.
How does a carnivore guild utilise a substantial but unpredictable anthropogenic food source? Scavenging on hunter-shot ungulate carcasses by wild dogs/dingoes, red foxes and feral cats in south-eastern Australia revealed by camera traps.Crossref | GoogleScholarGoogle Scholar | 24918425PubMed |

Forsyth, DM, Caley, P, Davis, NE, Latham, ADM, Woolnough, AP, Woodford, LP, Stamation, KA, Moloney, PD, and Pascoe, C (2018). Functional responses of an apex predator and a mesopredator to an invading ungulate: dingoes, red foxes and sambar deer in south-east Australia. Austral Ecology 43, 375–384.
Functional responses of an apex predator and a mesopredator to an invading ungulate: dingoes, red foxes and sambar deer in south-east Australia.Crossref | GoogleScholarGoogle Scholar |

Forsyth, DM, Latham, ADM, Davis, NE, Caley, P, Letnic, M, Moloney, PD, Woodford, LP, and Woolnough, AP (2019). Interactions between dingoes and introduced wild ungulates: concepts, evidence and knowledge gaps. Australian Mammalogy 41, 12–26.
Interactions between dingoes and introduced wild ungulates: concepts, evidence and knowledge gaps.Crossref | GoogleScholarGoogle Scholar |

Gaynor, KM, Hojnowski, CE, Carter, NH, and Brashares, JS (2018). The influence of human disturbance on wildlife nocturnality. Science 360, 1232–1235.
The influence of human disturbance on wildlife nocturnality.Crossref | GoogleScholarGoogle Scholar | 29903973PubMed |

Gelman, A, and Rubin, DB (1992). Inference from iterative simulation using multiple sequences. Statistical Science 7, 457–511.
Inference from iterative simulation using multiple sequences.Crossref | GoogleScholarGoogle Scholar |

Gormley, AM, Forsyth, DM, Griffioen, P, Lindeman, M, Ramsey, DSL, Scroggie, MP, and Woodford, L (2011). Using presence-only and presence–absence data to estimate the current and potential distributions of established invasive species. Journal of Applied Ecology 48, 25–34.
Using presence-only and presence–absence data to estimate the current and potential distributions of established invasive species.Crossref | GoogleScholarGoogle Scholar | 21339812PubMed |

Green, MJB (1987). Ecological separation in Himalayan ungulates. Journal of Zoology 1, 693–719.
Ecological separation in Himalayan ungulates.Crossref | GoogleScholarGoogle Scholar |

Harrel Jr FE (2020) Hmisc: Harrell miscellaneous. R package version 4.4-0. Available at https://CRAN.R-project.org/packageHmisc [verified 22 September 2021]

Harrison, M (2010) ‘Sambar the magnificent deer.’ (The Australian Deer Research Foundation Ltd: Melbourne)

Hijmans RJ (2020) Raster: Geographic data analysis and modeling. R package version 3.1-5. Available at https://CRAN.R-project.org/package=raster [verified 22 September 2021]

Ikeda, T, Takahashi, H, Igota, H, Matsuura, Y, Azumaya, M, Yoshida, T, and Kaji, K (2019). Effects of culling intensity on diel and seasonal activity patterns of sika deer (Cervus nippon). Scientific Reports 9, 17205.
Effects of culling intensity on diel and seasonal activity patterns of sika deer (Cervus nippon).Crossref | GoogleScholarGoogle Scholar | 31748671PubMed |

Jackson SM, Groves C (2015) ‘Taxonomy of Australian Mammals.’ (CSIRO Publishing: Melbourne, Vic., Australia)

Johnsingh, AJT (1983). Large mammalian prey and predators in Bandipur India. The Journal of the Bombay Natural History Society 80, 1–57.

Kawanishi, K, and Sunquist, ME (2004). Conservation status of tigers in a primary rainforest of peninsular Malaysia. Biological Conservation 120, 329–344.
Conservation status of tigers in a primary rainforest of peninsular Malaysia.Crossref | GoogleScholarGoogle Scholar |

Kie, JG (1999). Optimal foraging and risk of predation: Effects on behavior and social structure in ungulates. Journal of Mammalogy 80, 1114–1129.
Optimal foraging and risk of predation: Effects on behavior and social structure in ungulates.Crossref | GoogleScholarGoogle Scholar |

Kowalski M, Kowalski M (2013) Exifpro photo browser v 2.1.0. Available at https://github.com/mikekov/ExifPro [verified 22 September 2021]

Laguna, E, Carpio, AJ, Vicente, J, Barasona, JA, Triguero-Ocaña, R, Jiménez-Ruiz, S, Gómez-Manzaneque, Á, and Acevedo, P (2021). The spatial ecology of red deer under different land use and management scenarios: protected areas, mixed farms and fenced hunting estates. Science of The Total Environment 786, 147124.
The spatial ecology of red deer under different land use and management scenarios: protected areas, mixed farms and fenced hunting estates.Crossref | GoogleScholarGoogle Scholar | 33965822PubMed |

Laundre, JW, Hernandez, L, and Ripple, WJ (2010). The landscape of fear: Ecological implications of being afraid. The Open Ecology Journal 3, 1–7.
The landscape of fear: Ecological implications of being afraid.Crossref | GoogleScholarGoogle Scholar |

Le Saout, S, Padié, S, Chamaillé-Jammes, S, Chollet, S, Côté, S, Morellet, N, Pattison, J, Harris, E, and Martin, J-L (2014). Short-term effects of hunting on naïve black-tailed deer (Odocoileus hemionus sitkensis): Behavioural response and consequences on vegetation growth. Canadian Journal of Zoology 92, 915–925.
Short-term effects of hunting on naïve black-tailed deer (Odocoileus hemionus sitkensis): Behavioural response and consequences on vegetation growth.Crossref | GoogleScholarGoogle Scholar |

Linkie, M, and Ridout, MS (2011). Assessing tiger–prey interactions in Sumatran rainforests. Journal of Zoology 284, 224–229.
Assessing tiger–prey interactions in Sumatran rainforests.Crossref | GoogleScholarGoogle Scholar |

Little, AR, Demarais, S, Gee, KL, Webb, SL, Riffell, SK, Gaskamp, JA, and Belant, JL (2014). Does human predation risk affect harvest susceptibility of white-tailed deer during hunting season? Wildlife Society Bulletin 38, 797–805.
Does human predation risk affect harvest susceptibility of white-tailed deer during hunting season?Crossref | GoogleScholarGoogle Scholar |

Long JL (2003) ‘Introduced mammals of the world: their history, distribution and influence.’ (CSIRO Publishing: Melbourne, Victoria, Australia, and CABI Publishing: Wallingford, UK)

Luccarini, S, Mauri, L, Ciuti, S, Lamberti, P, and Apollonio, M (2006). Red deer (‘Cervus elaphus’) spatial use in the Italian alps: home range patterns, seasonal migrations, and effect of snow and winter feeding. Ethology Ecology & Evolution 18, 127–145.
Red deer (‘Cervus elaphus’) spatial use in the Italian alps: home range patterns, seasonal migrations, and effect of snow and winter feeding.Crossref | GoogleScholarGoogle Scholar |

Matsubayashi, H, Lagan, P, Sukor, JRA, and Kitayama, K (2007). Seasonal and daily use of natural licks by sambar deer (Cervus unicolor) in a Bornean tropical rain forest. Tropics 17, 81–86.
Seasonal and daily use of natural licks by sambar deer (Cervus unicolor) in a Bornean tropical rain forest.Crossref | GoogleScholarGoogle Scholar |

Menichetti, L, Touzot, L, Elofsson, K, Hyvönen, R, Kätterer, T, and Kjellander, P (2019). Interactions between a population of fallow deer (Dama dama), humans and crops in a managed composite temperate landscape in southern Sweden: conflict or opportunity? PLoS One 14, e0215594.
Interactions between a population of fallow deer (Dama dama), humans and crops in a managed composite temperate landscape in southern Sweden: conflict or opportunity?Crossref | GoogleScholarGoogle Scholar | 31013322PubMed |

Moloney, PD, Gormley, AM, Toop, SD, Flesch, JS, Forsyth, DM, Ramsey, DSL, and Hampton, JO (2022). Bayesian modelling reveals differences in long-term trends in the harvest of native and introduced species by recreational hunters in Australia. Wildlife Research. , .
Bayesian modelling reveals differences in long-term trends in the harvest of native and introduced species by recreational hunters in Australia.Crossref | GoogleScholarGoogle Scholar |

Moriarty, A (2004). The liberation, distribution, abundance and management of wild deer in Australia. Wildlife Research 31, 291–299.
The liberation, distribution, abundance and management of wild deer in Australia.Crossref | GoogleScholarGoogle Scholar |

Pal, R, Thakur, S, Arya, S, Bhattacharya, T, and Sathyakumar, S (2020). Mammals of the Bhagirathi basin, western Himalaya: understanding distribution along spatial gradients of habitats and disturbances. Oryx 55, 657–667.
Mammals of the Bhagirathi basin, western Himalaya: understanding distribution along spatial gradients of habitats and disturbances.Crossref | GoogleScholarGoogle Scholar |

Parsons, AW, Forrester, T, McShea, WJ, Baker-Whatton, MC, Millspaugh, JJ, and Kays, R (2017). Do occupancy or detection rates from camera traps reflect deer density? Journal of Mammalogy 98, 1547–1557.
Do occupancy or detection rates from camera traps reflect deer density?Crossref | GoogleScholarGoogle Scholar |

Pfeiffer, MB, Iglay, RB, Seamans, TW, Blackwell, BF, and DeVault, TL (2020). Deciphering interactions between white-tailed deer and approaching vehicles. Transportation Research Part D: Transport and Environment 79, 102251.
Deciphering interactions between white-tailed deer and approaching vehicles.Crossref | GoogleScholarGoogle Scholar |

Plummer, M, Best, N, Cowles, K, and Vines, K (2006). Coda: Convergence diagnosis and output analysis for MCMC. R News 6, 7–11.

R Core Team (2020) ‘R: a language and environment for statistical computing.’ (R Foundation for Statistical Computing: Vienna, Austria)

Ridout, MS, and Linkie, M (2009). Estimating overlap of daily activity patterns from camera trap data. Journal of Agricultural, Biological, and Environmental Statistics 14, 322–337.
Estimating overlap of daily activity patterns from camera trap data.Crossref | GoogleScholarGoogle Scholar |

Robertson, G, Wright, J, Brown, D, Yuen, K, and Tongway, D (2019). An assessment of feral horse impacts on treeless drainage lines in the Australian Alps. Ecological Management & Restoration 20, 21–30.
An assessment of feral horse impacts on treeless drainage lines in the Australian Alps.Crossref | GoogleScholarGoogle Scholar |

Ross, J, Hearn, AJ, Johnson, PJ, and Macdonald, DW (2013). Activity patterns and temporal avoidance by prey in response to Sunda clouded leopard predation risk. Journal of Zoology 290, 96–106.
Activity patterns and temporal avoidance by prey in response to Sunda clouded leopard predation risk.Crossref | GoogleScholarGoogle Scholar |

Semiadi, G, Muir, PD, and Barry, TN (1994). General biology of sambar deer (Cervus unicolor) in captivity. New Zealand Journal of Agricultural Research 37, 79–85.
General biology of sambar deer (Cervus unicolor) in captivity.Crossref | GoogleScholarGoogle Scholar |

Sih, A, Ziemba, R, and Harding, KC (2000). New insights on how temporal variation in predation risk shapes prey behavior. Trends in Ecology and Evolution 15, 3–4.
New insights on how temporal variation in predation risk shapes prey behavior.Crossref | GoogleScholarGoogle Scholar |

Simberloff, D, Parker, IM, and Windle, PN (2005). Introduced species policy, management, and future research needs. Frontiers in Ecology and the Environment 3, 12–20.
Introduced species policy, management, and future research needs.Crossref | GoogleScholarGoogle Scholar |

Simcharoen, A, Savini, T, Gale, G, Roche, E, Chimchome, V, and Smith, JL (2014). Ecological factors that influence sambar (Rusa unicolor) distribution and abundance in western Thailand: implications for tiger conservation. Raffles Bulletin of Zoology 62, 100–106.

Sotorra, S, Blair, D, Blanchard, W, and Lindenmayer, D (2021). Modelling the factors influencing Sambar Deer (Rusa unicolor) occurrence in the wet eucalypt forests of south-eastern Australia. Australian Zoologist 41, 241–253.
Modelling the factors influencing Sambar Deer (Rusa unicolor) occurrence in the wet eucalypt forests of south-eastern Australia.Crossref | GoogleScholarGoogle Scholar |

Stewart, KM, Bowyer, RT, Kie, JG, Cimon, NJ, and Johnson, BK (2002). Temporospatial distributions of elk, mule deer, and cattle: Resource partitioning and competitive displacement. Journal of Mammalogy 83, 229–244.
Temporospatial distributions of elk, mule deer, and cattle: Resource partitioning and competitive displacement.Crossref | GoogleScholarGoogle Scholar |

Tolsma A (2009) An assessment of mossbeds across the Victorian Alps, 2004–2009. Report to Parks Victoria. Arthur Rylah Institute for Environmental Research, Department of Sustainability and Environment, Melbourne. Copy of the report was provided by the author on 11 February 2022.

Wardle, DA, Barker, GM, Yeates, GW, Bonner, KI, and Ghani, A (2001). Introduced browsing mammals in New Zealand natural forests: Aboveground and belowground consequences. Ecological Monographs 71, 587–614.
Introduced browsing mammals in New Zealand natural forests: Aboveground and belowground consequences.Crossref | GoogleScholarGoogle Scholar |

Watter, K, Thomas, E, White, N, Finch, N, and Murray, PJ (2020). Reproductive seasonality and rate of increase of wild sambar deer (Rusa unicolor) in a new environment, Victoria, Australia. Animal Reproduction Science 223, 106630.
Reproductive seasonality and rate of increase of wild sambar deer (Rusa unicolor) in a new environment, Victoria, Australia.Crossref | GoogleScholarGoogle Scholar | 33166829PubMed |

White, KS, Pendleton, GW, and Hood, E (2009). Effects of snow on sitka black-tailed deer browse availability and nutritional carrying capacity in southeastern Alaska. Journal of Wildlife Management 73, 481–487.
Effects of snow on sitka black-tailed deer browse availability and nutritional carrying capacity in southeastern Alaska.Crossref | GoogleScholarGoogle Scholar |

Whitehead GK (1993) ‘The whitehead encyclopedia of deer.’ (Swan Hill Press: Shrewsbury, UK)

Wisdom MJ, Ager AA, Preisler HK, Cimon NJ, Johnson BK (2004) Effects of off-road recreation on mule deer and elk. In ‘Transactions of the 69th North American Wildlife and Natural Resources Conference’. pp. 531–550.

Wood, SN (2011). Fast stable restricted maximum likelihood and marginal likelihood estimation of semiparametric generalized linear models. Journal of the Royal Statistical Society: Series B (Statistical Methodology) 73, 3–36.
Fast stable restricted maximum likelihood and marginal likelihood estimation of semiparametric generalized linear models.Crossref | GoogleScholarGoogle Scholar |

Yamada, K, Elith, J, McCarthy, M, and Zerger, A (2003). Eliciting and integrating expert knowledge for wildlife habitat modelling. Ecological Modelling 165, 251–264.
Eliciting and integrating expert knowledge for wildlife habitat modelling.Crossref | GoogleScholarGoogle Scholar |

Yen, S-C, Wang, Y, Yu, P-H, Kuan, Y-P, Liao, Y-C, Chen, K-H, and Weng, G-J (2019). Seasonal space use and habitat selection of sambar in Taiwan. The Journal of Wildlife Management 83, 22–31.
Seasonal space use and habitat selection of sambar in Taiwan.Crossref | GoogleScholarGoogle Scholar |

Zar JH (1996) ‘Biostatistical analysis’. 3rd edn. (Prentice Hall: New Jersey)

Zhang, J, Hull, V, Ouyang, Z, Li, R, Connor, T, Yang, H, Zhang, Z, Silet, B, Zhang, H, and Liu, J (2017). Divergent responses of sympatric species to livestock encroachment at fine spatiotemporal scales. Biological Conservation 209, 119–129.
Divergent responses of sympatric species to livestock encroachment at fine spatiotemporal scales.Crossref | GoogleScholarGoogle Scholar |