Register      Login
Wildlife Research Wildlife Research Society
Ecology, management and conservation in natural and modified habitats
RESEARCH ARTICLE

Short-term impacts of large re-stocking on European hare population dynamics in a grassland landscape

Charles P. Henriot https://orcid.org/0000-0002-1724-2351 A * , Léopold Berger A , Thibaut Powolny B , Régis Renaude B , Christophe Bonenfant C # and Mickaël Sage A #
+ Author Affiliations
- Author Affiliations

A Faune INNOV’ R&D – Wildlife INNOVATION, Besançon, France.

B Fédération Départementale des Chasseurs du Doubs, Gonsans, France.

C Université Claude Bernard Lyon 1, UMR 5558 CNRS, Laboratoire Biométrie et Biologie Évolutive, Cedex, Villeurbanne 69622, France.

* Correspondence to: charles.henriot@faune-innov.fr

# These authors contributed equally to this paper

Handling Editor: Aaron Wirsing

Wildlife Research 52, WR25069 https://doi.org/10.1071/WR25069
Submitted: 29 April 2025  Accepted: 23 September 2025  Published: 13 October 2025

© 2025 The Author(s) (or their employer(s)). Published by CSIRO Publishing

Abstract

Context

The current rate of biodiversity loss increasingly motivates population reinforcement efforts. If reinforcement aims at increasing population abundance, the factors that influence the success of such operations remain poorly understood. Moreover, unintended side effects may lead to the opposite outcome such as a decrease in abundance following reinforcement. The abundance of European hare (Lepus europaeus) in France has been repeatedly reported to be in decline since the 1960s, especially in grassland landscapes. Although past studies have investigated the effects of hare translocation operations (i.e. reintroduction and re-stocking) on post-release survival rates, whether these wildlife management actions have led to an increase in the population abundance of European hare remains unclear.

Aims

Our objectives were to assess the impact of re-stocking operations on the population dynamics of European hare in a grassland landscape in the Jura Mountains (France), where hare populations are declining in numbers, and to determine whether these actions effectively enhance local abundance.

Methods

We investigated here the effects of hare re-stocking on (1) the rate and causes of mortality of 103 released individuals (51 in 2021 and 52 in 2022) and (2) the overall spatial and temporal impact of releases on local population dynamics by analysing the variation of hare abundance estimated from nocturnal point counts.

Key results

Despite the large number of released individuals, we report a limited short-term temporal impact of re-stocking on local population abundance of European hare. Overall survival rates of released hares were low (<10% after 1 year) owing to predation (57%), roadkill (20%), and diseases (12%), although the relative contribution of each cause varied over time after release.

Conclusion

Our results suggest that re-stockings had limited benefits for the mitigation of European hare decline, primarily owing to high post-release mortality, which is likely being driven by predator attraction to released individuals, especially in the days following release.

Implications

Reinforcement operations, especially for prey species, should perhaps consider strategies such as acclimatisation, soft-release, and anti-predator training, to reduce predation risk and improve the success of these interventions.

Keywords: demographic rates, grassland, Lepus europaeus, population management, population monitoring, reinforcement, re-stocking, spatially explicit models.

References

Agresti A (2013) ‘Categorical data analysis.’ 3rd edn. (Wiley: Hoboken, NJ, USA)

Anderson SC, Ward EJ (2019) Black swans in space: modeling spatiotemporal processes with extremes. Ecology 100(1), e02403.
| Crossref | Google Scholar | PubMed |

Angelici FM, Riga F, Boitani L, Luiselli L (2000) Fate of captive-reared brown hares Lepus europaeus released at a mountain site in central Italy. Wildlife Biology 6(3), 173-178.
| Crossref | Google Scholar |

Armstrong DP, Seddon PJ (2008) Directions in reintroduction biology. Trends in Ecology & Evolution 23(1), 20-25.
| Crossref | Google Scholar | PubMed |

Arnold TW (2010) Uninformative parameters and model selection using Akaike’s information criterion. Journal of Wildlife Management 74(6), 1175-1178.
| Crossref | Google Scholar |

Avril A, Léonard Y, Letty J, Péroux R, Guitton J-S, Pontier D (2011) Natal dispersal of European hare in a high-density population. Mammalian Biology 76(2), 148-156.
| Crossref | Google Scholar |

Báldi A, Faragó S (2007) Long-term changes of farmland game populations in a post-socialist country (Hungary). Agriculture, Ecosystems & Environment 118(1–4), 307-311.
| Crossref | Google Scholar |

Banasiak NM, Hayward MW, Kerley GIH (2021) Ten years on: have large carnivore reintroductions to the Eastern Cape Province, South Africa, worked? African Journal of Wildlife Research 51(1), 111-126.
| Crossref | Google Scholar |

Banks PB, Norrdahl K, Korpimäki E (2002) Mobility decisions and the predation risks of reintroduction. Biological Conservation 103(2), 133-138.
| Crossref | Google Scholar |

Basalamah F, Atmoko SSU, Perwitasari-Farajallah D, Qayim I, Sihite J, Noordwijk MV, Willems E, Schaik CPV (2018) Monitoring orangutan reintroduction: results of activity budgets, diets, vertical use and associations during the first year post-release in Kehje Sewen Forest, East Kalimantan, Indonesia. Biodiversitas Journal of Biological Diversity 19(2), 639-650.
| Crossref | Google Scholar |

Benmergui M, Reitz F, Fiechter A (1990) Taux de reprise et dispersion de lièvres (Lepus europaeus) sauvages d’Europe centrale relâchés dans l’Est de la France. Gibier Faune Sauvage 7, 255-274.
| Google Scholar |

Boucher S, Nouaille L (2013) ‘Maladies des lapins.’ 3rd edn. [mise à jour et enrichie]. (Éd. France Agricole: Paris, France)

Bray Y, Devillard S, Marboutin E, Mauvy B, Péroux R (2007) Natal dispersal of European hare in France. Journal of Zoology 273(4), 426-434.
| Crossref | Google Scholar |

Bubac CM, Johnson AC, Fox JA, Cullingham CI (2019) Conservation translocations and post-release monitoring: identifying trends in failures, biases, and challenges from around the world. Biological Conservation 238, 108239.
| Crossref | Google Scholar |

Burnham KP, Anderson DR (Eds) (2002) Advanced issues and deeper insights. In ‘Model selection and multimodel inference: a practical information-theoretic approach’. pp. 267–351. (Springer: New York, NY, USA) doi:10.1007/978-0-387-22456-5_6

Catalán I, Rodríguez-Hidalgo P, Tortosa FS (2008) Is habitat management an effective tool for wild rabbit (Oryctolagus cuniculus) population reinforcement? European Journal of Wildlife Research 54, 449-453.
| Crossref | Google Scholar |

Champagnon J, Elmberg J, Guillemain M, Gauthier-Clerc M, Lebreton J-D (2012) Conspecifics can be aliens too: a review of effects of restocking practices in vertebrates. Journal for Nature Conservation 20(4), 231-241.
| Crossref | Google Scholar |

Costantini D, Casagrande S, Di Lieto G, Fanfani A, Dell’Omo G (2005) Consistent differences in feeding habits between neighbouring breeding kestrels. Behaviour 142(9/10), 1403-1415.
| Google Scholar |

Čučković Z (2016) Advanced viewshed analysis: a Quantum GIS plug-in for the analysis of visual landscapes. The Journal of Open Source Software. Plugin available at https://plugins.qgis.org/plugins/ViewshedAnalysis/#plugin-about

Cukor J, Havránek F, Linda R, Bukovjan K, Painter MS, Hart V (2018) First findings of brown hare (Lepus europaeus) reintroduction in relation to seasonal impact. PLoS ONE 13(10), e0205078.
| Crossref | Google Scholar | PubMed |

Delibes-Mateos M, Fernandez De Simon J, Villafuerte R, Ferreras P (2008) Feeding responses of the red fox (Vulpes vulpes) to different wild rabbit (Oryctolagus cuniculus) densities: a regional approach. European Journal of Wildlife Research 54, 71-78.
| Crossref | Google Scholar |

Dickens MJ, Delehanty DJ, Michael Romero L (2010) Stress: an inevitable component of animal translocation. Biological Conservation 143(6), 1329-1341.
| Crossref | Google Scholar |

Esposito L, Auletta L, Ciani F, Pelagalli A, Pasolini MP, Lamagna B, Piscopo N, Amici A (2017) Hair cortisol levels in captive brown hare (Lepus europaeus): potential effect of sex, age, and breeding technology. European Journal of Wildlife Research 63, 62.
| Crossref | Google Scholar |

Ewen JG, Sainsbury AW, Jackson B, Canessa S (2015) Disease risk management in reintroduction. In ‘Advances in reintroduction biology of Australian and New Zealand fauna’. (Eds DP Armstrong, MW Hayward, D Moro, PJ Seddon) pp. 43–57. (CSIRO Publishing)

Fickel J, Schmidt A, Putze M, Spittler H, Ludwig A, Streich WJ, Pitra C (2005) Genetic structure of populations of European brown hare: implications for management. Journal of Wildlife Management 69(2), 760-770.
| Crossref | Google Scholar |

Fischer C, Tagand R (2012) Spatial behaviour and survival of translocated wild brown hares. Animal Biodiversity and Conservation 35, 189-196.
| Crossref | Google Scholar |

Gross IP, Wilson AE, Wolak ME (2024) The fitness consequences of wildlife conservation translocations: a meta-analysis. Biological Reviews 99(2), 348-371.
| Crossref | Google Scholar | PubMed |

Gryz J, Krauze-Gryz D (2022) Why did brown hare Lepus europaeus disappear from some areas in central Poland? Diversity 14(6), 465.
| Crossref | Google Scholar |

Hackländer K (2022) European Hare Lepus europaeus Pallas, 1778. In ‘Handbook of the Mammals of Europe’. (Eds K Hackländer, FE Zachos) pp. 1–36. (Springer International Publishing: Cham, Switzerland) doi:10.1007/978-3-319-65038-8_9-1

Hackländer K, Schai-Braun S (2018) Lepus europaeus; European hare. The IUCN Red List of Threatened Species. Available at https://dx.doi.org/10.2305/IUCN.UK.2019-1.RLTS.T41280A45187424.en

Henriot CP, Bonenfant C, Berger L, Powolny T, Renaude R, Sage M (2025) Survival, habitat selection, and success of wildlife reinforcement: a case study of European brown hares in a grassland-dominated landscape. In ‘Book of abstracts, 14th European vertebrate management conference’. (Eds B Pokorny, K Flajšman, J Jacob) p. 41. (Slovenian Forestry Institute) doi:10.20315/evmc.2025.021

Hušek J, Panek M (2024) Demographic and external drivers of European hare (Lepus europaeus) population dynamics in western Poland from 1960 to 2009. European Journal of Wildlife Research 70, 14.
| Crossref | Google Scholar |

IPBES (2019) Global assessment report on biodiversity and ecosystem services of the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services. Zenodo. Available at https://dx.doi.org/10.5281/ZENODO.3831673

IUCN (1987) The IUCN position statement on translocation of living organisms: introductions, re-introductions and re-stocking. Species Survival Commission in collaboration with the Commission on Ecology and the Commission on Environmental Policy, Law and Administration, Gland, Switzerland.

IUCN/SSC (2013) ‘Guidelines for reintroductions and other conservation translocations. Version 1.0.’ (IUCN Species Survival Commission: Gland, Switzerland)

Karmiris I (2006) Releasing captive brown hare (Lepus europaeus) to the wild – the role of predators. In ‘Management and development of mountainous and Island areas’. (Ed. E Manolas) pp. 205–208.

Kassambara A, Kosinski M, Biecek P (2021) Survminer: drawing survival curves using ‘ggplot2’. Available at https://CRAN.R-project.org/package=survminer

Kletty F, Pelé M, Capber F, Habold C (2020) Are all conservation measures for endangered species legitimate? Lines of thinking with the European hamster. Frontiers in Ecology and Evolution 8, 536937.
| Crossref | Google Scholar |

Letty J, Aubineau J, Marchandeau S (2008) Improving rabbit restocking success: a review of field experiments in France. In ‘Lagomorph biology’. (Eds PC Alves, N Ferrand, K Hackländer) pp. 327–348. (Springer: Heidelberg, Germany) doi:10.1007/978-3-540-72446-9_22

Linklater WL, Adcock K, Du Preez P, Swaisgood RR, Law PR, Knight MH, Gedir JV, Kerley GIH (2011) Guidelines for large herbivore translocation simplified: black rhinoceros case study. Journal of Applied Ecology 48(2), 493-502.
| Crossref | Google Scholar |

Machado RD, Magalhães P, Godinho S, Santos P (2017) Wild rabbit restocking: suitable acclimation conditions foster adaptive behaviour and improve survival of captive reared rabbits. World Rabbit Science 25(4), 407-414.
| Crossref | Google Scholar |

Marboutin E, Benmergui M, Pradel R, Fiechter A (1990) Survival patterns in wild and captive-reared leverets (Lepus europaeus Pallas) determined by telemetry. Gibier Faune Sauvage 7, 325-342.
| Google Scholar |

Misiorowska M, Wasilewski M (2012) Survival and causes of death among released brown hares (Lepus europaeus Pallas, 1778) in Central Poland. Acta Theriologica 57, 305-312.
| Crossref | Google Scholar | PubMed |

Mori E, Carbone R, Viviano A, Calosi M, Fattorini N (2022) Factors affecting spatiotemporal behaviour in the European brown hare Lepus europaeus: a meta-analysis. Mammal Review 52(3), 454-470.
| Crossref | Google Scholar |

Morris SD, Brook BW, Moseby KE, Johnson CN (2021) Factors affecting success of conservation translocations of terrestrial vertebrates: a global systematic review. Global Ecology and Conservation 28, e01630.
| Crossref | Google Scholar |

Moseby KE, Carthey A, Schroeder T (2015) The influence of predators and prey naivety on reintroduction success: current and future directions. In ‘Advances in reintroduction biology of Australian and New Zealand fauna’. (Eds D Armstrong, M Hayward, D Moro, PJ Seddon) pp. 29–42. (CSIRO Publishing)

Nelder JA (1974) Log linear models for contingency tables: a generalization of classical least squares. Applied Statistics 23(3), 323-329.
| Crossref | Google Scholar |

Paci G, Bagliacca M, Lavazza A (2006) Stress evaluation in hares (Lepus europaeus Pallas) captured for traslocation. Italian Journal of Animal Science 5(2), 175-181.
| Crossref | Google Scholar |

Panek M (2013) Long-term changes in the feeding pattern of red foxes Vulpes vulpes and their predation on brown hares Lepus europaeus in western Poland. European Journal of Wildlife Research 59, 581-586.
| Crossref | Google Scholar |

Panek M (2018) Habitat factors associated with the decline in brown hare abundance in Poland in the beginning of the 21st century. Ecological Indicators 85, 915-920.
| Crossref | Google Scholar |

Panek M (2023) Predation of young brown hares (Lepus europaeus) by common buzzards (Buteo buteo) in western Poland. European Journal of Wildlife Research 69, 110.
| Crossref | Google Scholar |

Péroux R, Mauvy B, Lartiges A, Bray Y, Marboutin E (1997) Point transect sampling: a new approach to estimate densities or abundances of European hare (Lepus europaeus) from spotlight counts. Game and Wildlife 14, 525-529.
| Google Scholar |

QGIS.org (2024) QGIS Geographic Information System. Open source geospatial foundation project. Available at http://qgis.org

R Core Team (2024) R: a language and environment for statistical computing. Available at https://www.R-project.org/

Reading RP, Miller B, Shepherdson D (2013) The value of enrichment to reintroduction success. Zoo Biology 32(3), 332-341.
| Crossref | Google Scholar | PubMed |

Réseau Lièvre (2020) Lièvres infos. Lettre d’information du Réseau Lièvre. No. 6. Office Français de la Biodiversité, (OFB).

Resende PS, Viana-Junior AB, Young RJ, Azevedo CSd (2020) A global review of animal translocation programs. Animal Biodiversity and Conservation 43, 221-232.
| Crossref | Google Scholar |

Roedenbeck IA, Voser P (2008) Effects of roads on spatial distribution, abundance and mortality of brown hare (Lepus europaeus) in Switzerland. European Journal of Wildlife Research 54, 425-437.
| Crossref | Google Scholar |

Savouré-Soubelet A, Arthur C-P, Aulagnier S, Body G, Callou C, Marchandeau S, Moutou F, Saint-Andrieux C (2021) ‘Atlas des mammifères sauvages de France – Ongulés et Lagomorphes’. (Muséum National d’Histoire Naturelle: Paris, France)

Sikes RS, the Animal Care and Use Committee of the American Society of Mammalogists (2016) 2016 Guidelines of the American Society of Mammalogists for the use of wild mammals in research and education. Journal of Mammalogy 97(3), 663-688.
| Crossref | Google Scholar |

Smith RK, Vaughan Jennings N, Harris S (2005) A quantitative analysis of the abundance and demography of European hares Lepus europaeus in relation to habitat type, intensity of agriculture and climate. Mammal Review 35(1), 1-24.
| Crossref | Google Scholar |

Sokos C, Birtsas P, Papaspyropoulos KG, Giannakopoulos A, Athanasiou LV, Manolakou K, Spyrou V, Billinis C (2015) Conservation considerations for a management measure: an integrated approach to hare rearing and release. Environmental Management 55, 19-30.
| Crossref | Google Scholar | PubMed |

Teixeira CP, De Azevedo CS, Mendl M, Cipreste CF, Young RJ (2007) Revisiting translocation and reintroduction programmes: the importance of considering stress. Animal Behaviour 73(1), 1-13.
| Crossref | Google Scholar |

Therneau T (2024) A package for survival analysis in R. Available at https://CRAN.R-project.org/package=survival

UNEP (2022) Decision adopted by the conference of the parties to the convention on biological diversity. 15/4. Kunming–Montreal global biodiversity framework. 15th Convention on Biological Diversity. UNEP – United Nations Environment Programme, Montreal, Canada.

Vaughan N, Lucas E-A, Harris S, White PCL (2003) Habitat associations of European hares Lepus europaeus in England and Wales: implications for farmland management. Journal of Applied Ecology 40(1), 163-175.
| Crossref | Google Scholar |

Ver Hoef JM, Boveng PL (2007) Quasi-poisson vs. negative binomial regression: how should we model overdispersed count data? Ecology 88(11), 2766-2772.
| Crossref | Google Scholar | PubMed |

Zellweger-Fischer J, Kéry M, Pasinelli G (2011) Population trends of brown hares in Switzerland: the role of land-use and ecological compensation areas. Biological Conservation 144(5), 1364-1373.
| Crossref | Google Scholar |