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RESEARCH ARTICLE (Open Access)

Genetic analysis reveals spatial structure in an expanding introduced rusa deer population

Scarlett Li-Williams https://orcid.org/0000-0002-6967-544X A # , Katarina C. Stuart A # , Sebastien Comte https://orcid.org/0000-0001-7984-8159 B , David M. Forsyth https://orcid.org/0000-0001-5356-9573 B , Michelle Dawson https://orcid.org/0009-0001-9313-3333 C D , William B. Sherwin https://orcid.org/0000-0002-1578-8473 A and Lee A. Rollins https://orcid.org/0000-0002-3279-7005 A *
+ Author Affiliations
- Author Affiliations

A Evolution & Ecology Research Centre, School of Biological, Earth and Environmental Sciences, The University of New South Wales, Sydney, NSW 2052, Australia.

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

C South-East Local Land Services, 84 Crown Street, Wollongong East, NSW 2520, Australia.

D Cultural and Ecosystem Adaptation Team, NSW Office of Energy and Climate Change, 12 Darcy Street, Parramatta, NSW 2124, Australia.

* Correspondence to: l.rollins@unsw.edu.au
# These authors contributed equally to this paper

Handling Editor: Graham Nugent

Wildlife Research 50(9) 757-769 https://doi.org/10.1071/WR22128
Submitted: 14 July 2022  Accepted: 11 June 2023   Published: 10 July 2023

© 2023 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: Rusa deer (Cervus timorensis), originally introduced in the 1860s, are still spreading in eastern Australia. The expanding peri-urban rusa deer population in the Illawarra region of New South Wales, Australia is having undesirable impacts on human and ecological communities, but the spatial structure of this population has not been investigated. Genetic information on invasive species is potentially useful in identifying management units to mitigate undesirable impacts.

Aims: The aim of this study was to investigate population structure, characterise dispersal, and determine if natural and human-made landscape features affected gene flow in rusa deer invading the Illawarra region of New South Wales.

Methods: We used reduced representation sequencing (DArT-Seq) to analyse single nucleotide polymorphisms distributed throughout the genomic DNA of rusa deer culled during a management program. We used admixture and Principal Component Analyses to investigate population structure with respect to natural and human-made landscape features, and we investigated whether our genetic data supported the presence of sex-biased dispersal.

Key results: Genetic diversity was highest in the north, near the original introduction site. A railway line demarcated restricted gene flow. Surprisingly, the Illawarra escarpment, a prominent landscape feature, did not restrict gene flow. There was no evidence of sex-biased dispersal and seven individuals were identified as genetic outliers.

Conclusions: The genetic structure of the Illawarra rusa deer population is consistent with individuals spreading south from their introduction site in Royal National Park. The population is not panmictic, and a landscape feature associated with urbanisation was associated with increased spatial genetic structure. Outliers could indicate hybridisation or secondary incursion events.

Implications: Rusa deer can be expected to continue invading southwards in the Illawarra region, but landscape features associated with urbanisation might reduce dispersal across the landscape. The genetic structuring of the population identified three potential management units on which to prioritise ground shooting operations.

Keywords: geographical range, invasive species, molecular ecology, population biology, population management, rusa deer, spatial structure, wildlife management.


References

AdaptNSW NSW Government (2022) Climate change in the Illawarra. Available at https://www.climatechange.environment.nsw.gov.au/illawarra#:~:text=The%20Illawarra%20region%20covers%20an,km2%20south%20of%20Sydney [Accessed 29 June 2022]

Alexander, DH, Novembre, J, and Lange, K (2009). Fast model-based estimation of ancestry in unrelated individuals. Genome Research 19, 1655–1664.
Fast model-based estimation of ancestry in unrelated individuals.Crossref | GoogleScholarGoogle Scholar |

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 |

Amos, M, Baxter, G, Finch, N, Lisle, A, and Murray, P (2014). I just want to count them! Considerations when choosing a deer population monitoring method. Wildlife Biology 20, 362–370.
I just want to count them! Considerations when choosing a deer population monitoring method.Crossref | GoogleScholarGoogle Scholar |

Amos, M, Pople, A, Brennan, M, Sheil, D, Kimber, M, and Cathcart, A (2023). Home ranges of rusa deer (Cervus timorensis) in a subtropical peri-urban environment in South East Queensland. Australian Mammalogy 45, 116–120.
Home ranges of rusa deer (Cervus timorensis) in a subtropical peri-urban environment in South East Queensland.Crossref | GoogleScholarGoogle Scholar |

Ando, C (2003). The relationship between deer-train collisions and daily activity of the sika deer, Cervus nippon. Mammal Study 28, 135–143.
The relationship between deer-train collisions and daily activity of the sika deer, Cervus nippon.Crossref | GoogleScholarGoogle Scholar |

Andrews S (2010) FastQC: a quality control tool for high throughput sequence data. Babraham Institute. Available at http://www.bioinformatics.babraham.ac.uk/projects/fastqc [Accessed 20 May 2021]

Anton, V, Hartley, S, and Wittmer, HU (2018). Evaluation of remote cameras for monitoring multiple invasive mammals in New Zealand. New Zealand Journal of Ecology 42, 74–79.
Evaluation of remote cameras for monitoring multiple invasive mammals in New Zealand.Crossref | GoogleScholarGoogle Scholar |

Ashcroft MB (2009) The spatial variation of environmental factors on the Illawarra escarpment and their influence on vegetation patterns. PhD thesis, University of Wollongong.

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 |

Bentley A (1998) ‘An introduction to the deer of Australia, with special reference to Victoria.’ (The Koetong Trust Fund and the Forest Commission of Victoria: Melbourne)

Bhardwaj, M, Olsson, M, Håkansson, E, Söderström, P, and Seiler, A (2022). Ungulates and trains–factors influencing flight responses and detectability. Journal of Environmental Management 313, 114992.
Ungulates and trains–factors influencing flight responses and detectability.Crossref | GoogleScholarGoogle Scholar |

Borda-de-Água L, Barrientos R, Beja P, Pereira HM (2017) ‘Railway ecology.’ (Springer Nature)

Brussard, PF (1984). Geographic patterns and environmental gradients: the central-marginal model in Drosophila revisited. Annual Review of Ecology and Systematics 15, 25–64.
Geographic patterns and environmental gradients: the central-marginal model in Drosophila revisited.Crossref | GoogleScholarGoogle Scholar |

Burgin, S, Mattila, M, McPhee, D, and Hundloe, T (2015). Feral deer in the suburbs: an emerging issue for Australia? Human Dimensions of Wildlife 20, 65–80.
Feral deer in the suburbs: an emerging issue for Australia?Crossref | GoogleScholarGoogle Scholar |

Catchen, J, Hohenlohe, PA, Bassham, S, Amores, A, and Cresko, WA (2013). Stacks: an analysis tool set for population genomics. Molecular Ecology 22, 3124–3140.
Stacks: an analysis tool set for population genomics.Crossref | GoogleScholarGoogle Scholar |

Chalmers PRS (2018) ‘New Zealand’s sambar and rusa deer. The full story.’ (Philip RS Chalmers: Whakatane)

Chhatre VE (2018) Distruct v2.3. A modified cluster membership plotting script. Available at http://distruct2.popgen.org

Ciach, M, and Fröhlich, A (2019). Ungulates in the city: light pollution and open habitats predict the probability of roe deer occurring in an urban environment. Urban Ecosystems 22, 513–523.
Ungulates in the city: light pollution and open habitats predict the probability of roe deer occurring in an urban environment.Crossref | GoogleScholarGoogle Scholar |

Coulon, A, Guillot, G, Cosson, J-F, Angibault, JMA, Aulagnier, S, Cargnelutti, B, Galan, M, and Hewison, AJM (2006). Genetic structure is influenced by landscape features: empirical evidence from a roe deer population. Molecular Ecology 15, 1669–1679.
Genetic structure is influenced by landscape features: empirical evidence from a roe deer population.Crossref | GoogleScholarGoogle Scholar |

Cripps, JK, Pacioni, C, Scroggie, MP, Woolnough, AP, and Ramsey, DSL (2019). Introduced deer and their potential role in disease transmission to livestock in Australia. Mammal Review 49, 60–77.
Introduced deer and their potential role in disease transmission to livestock in Australia.Crossref | GoogleScholarGoogle Scholar |

Crittle T, Millynn B (2020) Pest animal mapping 2020 final report. NSW Department of Primary Industries, Orange, NSW, Australia.

Cunningham, CX, Perry, GLW, Bowman, DMJS, Forsyth, DM, Driessen, MM, Appleby, M, Brook, BW, Hocking, G, Buettel, JC, French, BJ, Hamer, R, Bryant, SL, Taylor, M, Gardiner, R, Proft, K, Scoleri, VP, Chiu-Werner, A, Travers, T, Thompson, L, Guy, T, and Johnson, CN (2022). Dynamics and predicted distribution of an irrupting ‘sleeper’ population: fallow deer in Tasmania. Biological Invasions 24, 1131–1147.
Dynamics and predicted distribution of an irrupting ‘sleeper’ population: fallow deer in Tasmania.Crossref | GoogleScholarGoogle Scholar |

Danecek, P, Auton, A, Abecasis, G, Albers, CA, Banks, E, DePristo, MA, Handsaker, RE, Lunter, G, Marth, GT, Sherry, ST, and McVean, G (2011). The variant call format and VCFtools. Bioinformatics 27, 2156–2158.
The variant call format and VCFtools.Crossref | GoogleScholarGoogle Scholar |

Davies, C, Wright, W, Wedrowicz, F, Pacioni, C, and Hogan, FE (2022). Delineating genetic management units of sambar deer (Rusa unicolor) in south-eastern Australia, using opportunistic tissue sampling and targeted scat collection. Wildlife Research 49, 147–157.
Delineating genetic management units of sambar deer (Rusa unicolor) in south-eastern Australia, using opportunistic tissue sampling and targeted scat collection.Crossref | GoogleScholarGoogle Scholar |

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 |

Dawson M (2017) Tools for managing wild deer: ground shooting in the Northern Illawarra Wild Deer Management Program. In ‘2016 National Wild Deer Management Workshop Proceedings, Canberra, Australia, 2017’. (Eds DM Forsyth, T Pople, B Page, A Moriarty, D Ramsey, J Parkes, A Wiebkin, C Lane) pp. 22–23. (Invasive Animals Cooperative Research Centre)

Department of Agriculture Water and the Environment (2011) Feral deer. Available at https://www.environment.gov.au/biodiversity/invasive-species/publications/factsheet-feral-deer [Accessed 20 March 2021]

Diefenbach, DR, Long, ES, Rosenberry, CS, Wallingford, BD, and Smith, DR (2008). Modeling distribution of dispersal distances in male white-tailed deer. Journal of Wildlife Management 72, 1296–1303.
Modeling distribution of dispersal distances in male white-tailed deer.Crossref | GoogleScholarGoogle Scholar |

Drolet, A, Dussault, C, and Côté, SD (2016). Simulated drilling noise affects the space use of a large terrestrial mammal. Wildlife Biology 22, 284–293.
Simulated drilling noise affects the space use of a large terrestrial mammal.Crossref | GoogleScholarGoogle Scholar |

Dryden, GM (2000). Biology and production attributes of the farmed rusa deer. Asian-Australasian Journal of Animal Sciences 13, 69–72.

Forsyth, DM, Duncan, RP, Bomford, M, and Moore, G (2004). Climatic suitability, life-history traits, introduction effort, and the establishment and spread of introduced mammals in Australia. Conservation Biology 18, 557–569.
Climatic suitability, life-history traits, introduction effort, and the establishment and spread of introduced mammals in Australia.Crossref | GoogleScholarGoogle Scholar |

Forsyth DM, Pople T, Page B, Moriarty T, Ramsey D, Parkes J, Wiebkin A, Lane C (2017) 2016 National Wild Deer Management Workshop Proceedings, 17–18 November 2016, Adelaide, Australia’. (Invasive Animals Cooperative Research Centre: Canberra, Australia)

Forsyth, DM, Comte, S, Davis, NE, Bengsen, AJ, Côté, SD, Hewitt, DG, Morellet, N, and Mysterud, A (2022). Methodology matters when estimating deer abundance: a global systematic review and recommendations for improvements. The Journal of Wildlife Management 86, e22207.
Methodology matters when estimating deer abundance: a global systematic review and recommendations for improvements.Crossref | GoogleScholarGoogle Scholar |

Frantz, AC, Bertouille, S, Eloy, MC, Licoppe, A, Chaumont, F, and Flamand, MC (2012). Comparative landscape genetic analyses show a Belgian motorway to be a gene flow barrier for red deer (Cervus elaphus), but not wild boars (Sus scrofa). Molecular Ecology 21, 3445–3457.
Comparative landscape genetic analyses show a Belgian motorway to be a gene flow barrier for red deer (Cervus elaphus), but not wild boars (Sus scrofa).Crossref | GoogleScholarGoogle Scholar |

Fraser, D, Ironside, KE, Wayne, RK, and Boydston, EE (2019). Connectivity of mule deer (Odocoileus hemionus) populations in southern California: a genetic survey of a mobile ungulate in a highly fragmented urban landscape. Landscape Ecology 34, 1–19.
Connectivity of mule deer (Odocoileus hemionus) populations in southern California: a genetic survey of a mobile ungulate in a highly fragmented urban landscape.Crossref | GoogleScholarGoogle Scholar |

Garcia-Gonzalez, C, Campo, D, Pola, IG, and Garcia-Vazquez, E (2012). Rural road networks as barriers to gene flow for amphibians: Species-dependent mitigation by traffic calming. Landscape and Urban Planning 104, 171–180.
Rural road networks as barriers to gene flow for amphibians: Species-dependent mitigation by traffic calming.Crossref | GoogleScholarGoogle Scholar |

Georges A, Gruber B (2019) SNP analysis using dartR: guide to preparatory analysis. Session 1: introduction to DArTSeq (short course — A Primer on dartR). University of Canberra (B. biomatix.org.au: The Institute for Applied Ecology)

Gill, RMA, Thomas, ML, and Stocker, D (1997). The use of portable thermal imaging for estimating deer population density in forest habitats. The Journal of Applied Ecology 34, 1273–1286.
The use of portable thermal imaging for estimating deer population density in forest habitats.Crossref | GoogleScholarGoogle Scholar |

Gilmour P, Bowden R, Denniss R (2016) Evaluation of the Northern Illawarra wild deer management program – report. First Person Consulting Pty Ltd. Available at https://www.aph.gov.au/DocumentStore.ashx?id=20a0bb4c-7c92-4a91-b159-5fe01ec7e512&subId=665019#:~:text=The%20NIWDMP%20appears%20to%20be,or%20downward%20trend%20in%20impacts [Accessed 26 June 2023]

Goudet, J, Perrin, N, and Waser, P (2002). Tests for sex-biased dispersal using bi-parentally inherited genetic markers. Molecular Ecology 11, 1103–1114.
Tests for sex-biased dispersal using bi-parentally inherited genetic markers.Crossref | GoogleScholarGoogle Scholar |

Gruber, B, Unmack, PJ, Berry, OF, and Georges, A (2018). dartr: an r package to facilitate analysis of SNP data generated from reduced representation genome sequencing. Molecular Ecology Resources 18, 691–699.
dartr: an r package to facilitate analysis of SNP data generated from reduced representation genome sequencing.Crossref | GoogleScholarGoogle Scholar |

Hall, GP, and Gill, KP (2005). Management of wild deer in Australia. Journal of Wildlife Management 69, 837–844.
Management of wild deer in Australia.Crossref | GoogleScholarGoogle Scholar |

Hampton, JO, MacKenzie, DI, and Forsyth, DM (2022). Animal welfare outcomes of professional vehicle-based shooting of peri-urban rusa deer in Australia. Wildlife Research , .
Animal welfare outcomes of professional vehicle-based shooting of peri-urban rusa deer in Australia.Crossref | GoogleScholarGoogle Scholar |

Hedges S, Duckworth JW, Timmins R, Semiadi G, Dryden G (2015) Rusa timorensis. The IUCN red list of threatened species 2015: e.T41789A22156866. Available at http://dx.doi.org/10.2305/IUCN.UK.2015-2.RLTS.T41789A22156866.en

Hepenstrick, D, Thiel, D, Holderegger, R, and Gugerli, F (2012). Genetic discontinuities in roe deer (Capreolus capreolus) coincide with fenced transportation infrastructure. Basic and Applied Ecology 13, 631–638.
Genetic discontinuities in roe deer (Capreolus capreolus) coincide with fenced transportation infrastructure.Crossref | GoogleScholarGoogle Scholar |

Hill, E, Murphy, N, Li-Williams, S, Davies, C, Forsyth, D, Comte, S, Rollins, L, Hogan, F, Wedrowicz, F, Crittle, T, Thomas, E, Woodford, L, and Pacioni, C (2023). Hybridisation rates, population structure, and dispersal of sambar deer (Cervus unicolor) and rusa deer (Cervus timorensis) in south-eastern Australia. Wildlife Research , .

Hindrikson, M, Remm, J, Pilot, M, Godinho, R, Stronen, AV, Baltrūnaité, L, Czarnomska, SD, Leonard, JA, Randi, E, Nowak, C, Åkesson, M, López-Bao Jé, V, Álvares, F, Llaneza, L, Echegaray, J, Vilà, C, Ozolins, J, Rungis, D, Aspi, J, Paule, L, Skrbinšek, T, and Saarma, U (2017). Wolf population genetics in Europe: a systematic review, meta-analysis and suggestions for conservation and management. Biological Reviews 92, 1601–1629.
Wolf population genetics in Europe: a systematic review, meta-analysis and suggestions for conservation and management.Crossref | GoogleScholarGoogle Scholar |

Hodnett E (2005) Thermal imaging applications in urban deer control. In ‘Proceedings of the 11th Wildlife Damage Management Conference’. (Eds DL Nolte, KA Fagerstone) pp. 141–148. (UNL Digital Commons)

Huijser, MP, and Begley, JS (2022). Implementing wildlife fences along highways at the appropriate spatial scale: a case study of reducing road mortality of Florida Key deer. Nature Conservation 47, 283–302.
Implementing wildlife fences along highways at the appropriate spatial scale: a case study of reducing road mortality of Florida Key deer.Crossref | GoogleScholarGoogle Scholar |

Ingleby S (2022) Rusa deer. Australian Museum.

Kilian A, Wenzl P, Huttner E, Carling J, Xia L, Blois H, Caig V, Heller-Uszynska K, Jaccoud D, Hopper C (2012) Diversity arrays technology: a generic genome profiling technology on open platforms. In ‘Data production and analysis in population genomics’. (Eds F Pompanon, A Bonin) pp. 67–89. (Springer)

King CM, Forsyth DM (2021) ‘The handbook of New Zealand mammals.’ (CSIRO Publishing)

Kuehn, R, Hindenlang, KE, Holzgang, O, Senn, J, Stoeckle, B, and Sperisen, C (2006). Genetic effect of transportation infrastructure on roe deer populations (Capreolus capreolus). Journal of Heredity 98, 13–22.
Genetic effect of transportation infrastructure on roe deer populations (Capreolus capreolus).Crossref | GoogleScholarGoogle Scholar |

Laikre, L, Palm, S, and Ryman, N (2005). Genetic population structure of fishes: implications for coastal zone management. AMBIO: A Journal of the Human Environment 34, 111–119.
Genetic population structure of fishes: implications for coastal zone management.Crossref | GoogleScholarGoogle Scholar |

Langin, KM, Sillett, TS, Funk, WC, Morrison, SA, and Ghalambor, CK (2017). Partial support for the central–marginal hypothesis within a population: reduced genetic diversity but not increased differentiation at the range edge of an island endemic bird. Heredity 119, 8–15.
Partial support for the central–marginal hypothesis within a population: reduced genetic diversity but not increased differentiation at the range edge of an island endemic bird.Crossref | GoogleScholarGoogle Scholar |

Latch, EK, Gee, KL, Webb, SL, Honeycutt, RL, DeYoung, RW, Gonzales, RA, Demarais, S, and Toby, R (2021). Genetic consequences of fence confinement in a population of white-tailed deer. Diversity 13, 126.
Genetic consequences of fence confinement in a population of white-tailed deer.Crossref | GoogleScholarGoogle Scholar |

Lawson Handley, L-J, Estoup, A, Evans, DM, Thomas, CE, Lombaert, E, Facon, B, Aebi, A, and Roy, HE (2011). Ecological genetics of invasive alien species. BioControl 56, 409–428.
Ecological genetics of invasive alien species.Crossref | GoogleScholarGoogle Scholar |

Li, H, and Durbin, R (2009). Fast and accurate short read alignment with Burrows–Wheeler transform. Bioinformatics 25, 1754–1760.
Fast and accurate short read alignment with Burrows–Wheeler transform.Crossref | GoogleScholarGoogle Scholar |

Lizarralde, MS, Bailliet, G, Poljak, S, Fasanella, M, and Giulivi, C (2008). Assessing genetic variation and population structure of invasive North American beaver (Castor Canadensis Kuhl, 1820) in Tierra Del Fuego (Argentina). Biological Invasions 10, 673–683.
Assessing genetic variation and population structure of invasive North American beaver (Castor Canadensis Kuhl, 1820) in Tierra Del Fuego (Argentina).Crossref | GoogleScholarGoogle Scholar |

Long, ES, Diefenbach, DR, Rosenberry, CS, Wallingford, BD, and Grund, MD (2005). Forest cover influences dispersal distance of white-tailed deer. Journal of Mammalogy 86, 623–629.
Forest cover influences dispersal distance of white-tailed deer.Crossref | GoogleScholarGoogle Scholar |

Long, ES, Diefenbach, DR, Wallingford, BD, and Rosenberry, CS (2010). Influence of roads, rivers, and mountains on natal dispersal of white-tailed deer. The Journal of Wildlife Management 74, 1242–1249.
Influence of roads, rivers, and mountains on natal dispersal of white-tailed deer.Crossref | GoogleScholarGoogle Scholar |

Loving SC (2022) Integrating telemetry, remote sensing and landscape genetics for assessing feral pig (Sus scrofa) movement ecology. PhD thesis, Charles Darwin University, Australia.

Martins, RF, Schmidt, A, Lenz, D, Wilting, A, and Fickel, J (2018). Human-mediated introduction of introgressed deer across Wallace’s line: historical biogeography of Rusa unicolor and R. timorensis. Ecology and Evolution 8, 1465–1479.
Human-mediated introduction of introgressed deer across Wallace’s line: historical biogeography of Rusa unicolor and R. timorensis.Crossref | GoogleScholarGoogle Scholar |

Meek, PD, Ballard, GA, Sparkes, J, Robinson, M, Nesbitt, B, and Fleming, PJS (2019). Camera trap theft and vandalism: occurrence, cost, prevention and implications for wildlife research and management. Remote Sensing in Ecology and Conservation 5, 160–168.
Camera trap theft and vandalism: occurrence, cost, prevention and implications for wildlife research and management.Crossref | GoogleScholarGoogle Scholar |

Meisingset, EL, Loe, LE, Brekkum, Ø, Van Moorter, B, and Mysterud, A (2013). Red deer habitat selection and movements in relation to roads. The Journal of Wildlife Management 77, 181–191.
Red deer habitat selection and movements in relation to roads.Crossref | GoogleScholarGoogle Scholar |

Moriarty, A (2004a). 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 |

Moriarty AJ (2004b) Ecology and environmental impact of Javan rusa deer (Cervus timorensis russa) in the Royal National Park. PhD thesis, University of Western Sydney, Sydney.

Murtagh, F, and Legendre, P (2014). Ward’s hierarchical agglomerative clustering method: which algorithms implement Ward’s criterion? Journal of classification 31, 274–295.
Ward’s hierarchical agglomerative clustering method: which algorithms implement Ward’s criterion?Crossref | GoogleScholarGoogle Scholar |

NPWS (2002) Wollongong LGA bioregional assessment (part I): native vegetation of the Illawarra Escarpment and coastal plain. NSW National Parks and Wildlife Service, Sydney.

NSW Department of Planning Industry and Environment (2021) Feral deer. Available at https://www.environment.nsw.gov.au/topics/animals-and-plants/pest-animals-and-weeds/pest-animals/feral-deer [Accessed 12 March 2021]

NSW Government, Office of Environment and Heritage (2014) Illawarra Climate change snapshot report. Office of Environment and Heritage.

Olsson M (2007) The use of highway crossings to maintain landscape connectivity for moose and roe deer. PhD thesis, Fakulteten för samhälls-och livsvetenskaper.

Peterson, BE, Storm, DJ, Norton, AS, and Van Deelen, TR (2017). Landscape influence on dispersal of yearling male white-tailed deer. The Journal of Wildlife Management 81, 1449–1456.
Landscape influence on dispersal of yearling male white-tailed deer.Crossref | GoogleScholarGoogle Scholar |

Phillips, BL, Brown, GP, Travis, JMJ, and Shine, R (2008). Reid’s paradox revisited: the evolution of dispersal kernels during range expansion. The American Naturalist 172, S34–S48.
Reid’s paradox revisited: the evolution of dispersal kernels during range expansion.Crossref | GoogleScholarGoogle Scholar |

Pinkston R (2009) Military thermal technology adapted to wildlife control and management. In ‘Proceedings of the 13th Wildlife Damage Management Conference’. (Ed. JR Boulanger) pp. 121–125. (UNL Digital Commons) https://doi.org/10.13140/RG.2.2.22383.02727

Pitra, C, Fickel, J, Meijaard, E, and Groves, C (2004). Evolution and phylogeny of old world deer. Molecular Phylogenetics and Evolution 33, 880–895.
Evolution and phylogeny of old world deer.Crossref | GoogleScholarGoogle Scholar |

PVC (Research Infrastructure), UNSW Sydney (2010) Katana. UNSW, Sydney.

Pérez-Espona, S, Pérez-Barbería, FJ, McLeod, JE, Jiggins, CD, Gordon, IJ, and Pemberton, JM (2008). Landscape features affect gene flow of Scottish Highland red deer (Cervus elaphus). Molecular Ecology 17, 981–996.
Landscape features affect gene flow of Scottish Highland red deer (Cervus elaphus).Crossref | GoogleScholarGoogle Scholar |

Pérez-Espona, S, Pérez-Barbería, FJ, Jiggins, CD, Gordon, IJ, and Pemberton, JM (2010). Variable extent of sex-biased dispersal in a strongly polygynous mammal. Molecular Ecology 19, 3101–3113.
Variable extent of sex-biased dispersal in a strongly polygynous mammal.Crossref | GoogleScholarGoogle Scholar |

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

Reddy, PA, Cushman, SA, Srivastava, A, Sarkar, MS, and Shivaji, S (2017). Tiger abundance and gene flow in Central India are driven by disparate combinations of topography and land cover. Diversity and Distributions 23, 863–874.
Tiger abundance and gene flow in Central India are driven by disparate combinations of topography and land cover.Crossref | GoogleScholarGoogle Scholar |

Reddy, PA, Puyravaud, J-P, Cushman, SA, and Segu, H (2019). Spatial variation in the response of tiger gene flow to landscape features and limiting factors. Animal Conservation 22, 472–480.
Spatial variation in the response of tiger gene flow to landscape features and limiting factors.Crossref | GoogleScholarGoogle Scholar |

Riva, F, Acorn, JH, and Nielsen, SE (2018). Narrow anthropogenic corridors direct the movement of a generalist boreal butterfly. Biology Letters 14, 20170770.
Narrow anthropogenic corridors direct the movement of a generalist boreal butterfly.Crossref | GoogleScholarGoogle Scholar |

Robinson, SJ, Samuel, MD, Lopez, DL, and Shelton, P (2012). The walk is never random: subtle landscape effects shape gene flow in a continuous white-tailed deer population in the Midwestern United States. Molecular Ecology 21, 4190–4205.
The walk is never random: subtle landscape effects shape gene flow in a continuous white-tailed deer population in the Midwestern United States.Crossref | GoogleScholarGoogle Scholar |

Rollins, LA, Woolnough, AP, and Sherwin, WB (2006). Population genetic tools for pest management: a review. Wildlife Research 33, 251–261.
Population genetic tools for pest management: a review.Crossref | GoogleScholarGoogle Scholar |

Rollins, LA, Woolnough, AP, Wilton, AN, Sinclair, R, and Sherwin, WB (2009). Invasive species can’t cover their tracks: using microsatellites to assist management of starling (Sturnus vulgaris) populations in Western Australia. Molecular Ecology 18, 1560–1573.
Invasive species can’t cover their tracks: using microsatellites to assist management of starling (Sturnus vulgaris) populations in Western Australia.Crossref | GoogleScholarGoogle Scholar |

Rousset, F (1997). Genetic differentiation and estimation of gene flow from F-statistics under isolation by distance. Genetics 145, 1219–1228.
Genetic differentiation and estimation of gene flow from F-statistics under isolation by distance.Crossref | GoogleScholarGoogle Scholar |

Sakai, AK, Allendorf, FW, Holt, JS, Lodge, DM, Molofsky, J, With, KA, Baughman, S, Cabin, RJ, Cohen, JE, Ellstrand, NC, McCauley, DE, O’Neil, P, Parker, IM, Thompson, JN, and Weller, SG (2001). The population biology of invasive species. Annual Review of Ecology and Systematics 32, 305–332.
The population biology of invasive species.Crossref | GoogleScholarGoogle Scholar |

Sexton, JP, Hangartner, SB, and Hoffmann, AA (2014). Genetic isolation by environment or distance: which pattern of gene flow is most common? Evolution 68, 1–15.
Genetic isolation by environment or distance: which pattern of gene flow is most common?Crossref | GoogleScholarGoogle Scholar |

Shimatani, Y, Fukue, Y, Kishimoto, R, and Masuda, R (2010). Genetic variation and population structure of the feral American mink (Neovison vison) in Nagano, Japan, revealed by microsatellite analysis. Mammal Study 35, 1–7.
Genetic variation and population structure of the feral American mink (Neovison vison) in Nagano, Japan, revealed by microsatellite analysis.Crossref | GoogleScholarGoogle Scholar |

Soulsbury, CD, and White, PCL (2015). Human–wildlife interactions in urban areas: a review of conflicts, benefits and opportunities. Wildlife Research 42, 541–553.
Human–wildlife interactions in urban areas: a review of conflicts, benefits and opportunities.Crossref | GoogleScholarGoogle Scholar |

Spaggiari, J, and de Garine-Wichatitsky, M (2006). Home range and habitat use of introduced rusa deer (Cervus timorensis russa) in a mosaic of savannah and native sclerophyll forest of New Caledonia. New Zealand Journal of Zoology 33, 175–183.
Home range and habitat use of introduced rusa deer (Cervus timorensis russa) in a mosaic of savannah and native sclerophyll forest of New Caledonia.Crossref | GoogleScholarGoogle Scholar |

Stout, RJ, Knuth, BA, and Curtis, PD (1997). Preferences of suburban landowners for deer management techniques: a step towards better communication. Wildlife Society Bulletin 25, 348–359.

The Australian Bureau of Statistics (2020) Region summary: Illawarra. Available at &https://dbr.abs.gov.au/region.html?lyr=sa4&rgn=107 [Accessed 20 June 2022]

Webley, LS, Zenger, KR, English, AW, and Cooper, DW (2004). Low levels of genetic variation within introduced Javan rusa deer (Cervus timorensis russa) in Australia. European Journal of Wildlife Research 50, 137–140.
Low levels of genetic variation within introduced Javan rusa deer (Cervus timorensis russa) in Australia.Crossref | GoogleScholarGoogle Scholar |

Wickham H (2016) ‘ggplot2: elegant graphics for data analysis.’ (Springer-Verlag: New York)

Wollongong City Council (2020) Deer. Available at https://www.wollongong.nsw.gov.au/about/environment/pest-animals/deer [Accessed 6 June 2021]

Woodford KB, Dunning A (1992) Production cycles and characteristics of rusa deer in Queensland, Australia. In ‘The biology of deer’. (Ed. RD Brown) pp. 197–202. (Springer-Verlag: New York)

Young RW (1979) The Illawarra Escarpment. Wollongong Studies in Geography. University of Wollongong.

Youngson, AF, Jordan, WC, Verspoor, E, McGinnity, P, Cross, T, and Ferguson, A (2003). Management of salmonid fisheries in the British Isles: towards a practical approach based on population genetics. Fisheries Research 62, 193–209.
Management of salmonid fisheries in the British Isles: towards a practical approach based on population genetics.Crossref | GoogleScholarGoogle Scholar |

Zheng, X, Levine, D, Shen, J, Gogarten, SM, Laurie, C, and Weir, BS (2012). A high-performance computing toolset for relatedness and principal component analysis of SNP data. Bioinformatics 28, 3326–3328.
A high-performance computing toolset for relatedness and principal component analysis of SNP data.Crossref | GoogleScholarGoogle Scholar |