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Context

Populations of red deer (Cervus elaphus), roe deer (Capreolus capreolus) and wild boar (Sus scrofa) have been expanding rapidly in France since the 1970s. After a long period of population recovery with minor and generally very localised effects on the forest environment, these increases raise new questions.
•    What effects do these populations have on the biodiversity of the undergrowth and the renewal of forest species in particular?
•    What means should be used to adapt the management of these populations to different territorial entities?

Forest renewal with wild ungulates

After decades of capitalising on wood resources in the forest, the current economic challenges and climatic hazards are prompting the forestry and wood industry to intensify forestry operations. This process, accompanied by a proactive forestry policy (National Forestry and Wood Plan 2016-2026), recommends an increase in wood harvests and early forest renewal.

In this context, the herbivory pressure exerted by wild ungulates on forest species leads to an increasing risk of disturbance. This hindrance primarily affects the renewal phase of the forest and then has an impact on the course of the silvicultural itineraries.

Our work concerns the study of demographic parameters (survival, growth) of various forest species in order to characterise their capacity for resistance and resilience to the effects of herbivores. This knowledge is intended to feed growth dynamics models for forest stands, but also to build tools to help diagnose the silvicultural impact of ungulates at multiple scales (stand, forest, region). The aim is to propose the elements necessary to support forest renewal in line with silvicultural objectives, the environmental functions of the forest and also climate issues.

For more information:

• Mårell, A., Hamard, J.-P., Pérot, T., Perret, S., Korboulewsky, N. 2018. The effect of deer browsing and understory light availability on stump mortality and sprout growth capacity in sessile oak. Forest Ecology and Management 430:134-142. https://doi.org/10.1016/j.foreco.2018.08.015
• Laurent, L., Mårell, A., Balandier, P., Holveck, H., Saïd, S., 2017. Understory vegetation dynamics and tree regeneration as affected by deer herbivory in temperate hardwood forests. iForest - Biogeosciences and Forestry 10:837-844. https://doi.org/10.3832/ifor2186-010
• Laurent, L., Mårell, A., Korboulewsky, N., Saïd, S., Balandier, P. 2017. How does disturbance affect the intensity and importance of plant competition along resource gradients? Forest Ecology and Management 391:239-245. https://doi.org/10.1016/j.foreco.2017.02.003
• Bideau, E., Maublanc, M.L. Picot, D., Hamard, J.P., Ballon, P., Gerard, J.F. (2015). Short-term browsing by roe deer has little effect on survival and growth of sessile oak seedlings. Scandinavian Journal of Forest Research 31(1) : 40-45. https://doi.org/10.1080/02827581.2015.1054873

The effects of wild ungulates on the biodiversity of forest ecosystems

This theme addresses the effects of wild ungulates on the functioning and biodiversity of diverse terrestrial ecosystems (grasslands, forests or tundra). We study the direct effects of herbivory on plant demography (e.g. anemone, bramble, oaks), plant community dynamics (vascular plants and understorey bryophytes) and forestry (forest regeneration). We use experimental enclosures, spatial and temporal herbivory gradients (natural and simulated) and modelling. We address the direct and indirect effects of herbivory on plant-plant interactions (competition, facilitation) and on other ecosystem components in the context of changing silvicultural practices, climate change or animal overabundance.

For more information:

• Boulanger, V., Dupouey, J.-L., Archaux, F., Badeau, V., Baltzinger, C., Chevalier, R., Corcket, E., Dumas, Y., Forgeard, F., Mårell, A., Montpied, P., Paillet, Y., Picard, J.-F., Saïd, S., Ulrich, E. 2018. Ungulates increase forest plant species richness to the benefit of non-forest specialists. Global Change Biology, 24:e485–e495. https://doi.org/10.1111/gcb.13899
• Laurent, L., Mårell, A., Balandier, P., Holveck, H., Saïd, S., 2017. Understory vegetation dynamics and tree regeneration as affected by deer herbivory in temperate hardwood forests. iForest - Biogeosciences and Forestry 10:837-844. https://doi.org/10.3832/ifor2186-010
• Boulanger, V., Baltzinger, C., Said, S., Ballon, P., Picard, J.F., Dupouey, J.L. (2015). Decreasing deer browsing pressure influenced understory vegetation dynamics over 30 years. Annals of Forest Science 72:367-378. https://doi.org/10.1007/s13595-014-0431-z

Functional role of wild ungulates

This theme addresses ecological processes associated with wild ungulates such as trophic interactions (herbivory, predation) and their cascading effects, zoochory and physical engineering. These processes affect the spatial and temporal dynamics of the flora and the functioning of ecosystems. Both selective consumers of plants and prey for predators, wild ungulates create heterogeneity (e.g. directed flows of nutrients and diaspores, physico-chemical modifications of the soil) within their home ranges and on a population scale. Their impacts are all the more important as these species have high movement capacities, consume a wide range of plants, and are extremely flexible, with rapid adaptation capacities to their environment. Indeed, while the role of ungulates in the functioning of ecosystems is largely determined by their spatial and feeding behaviour, the latter are themselves greatly influenced by human activities, landscape structure and heterogeneity, with cascading consequences on ecological processes and on the spatial distribution of associated services and disservices. This is why we are also interested in the movement ecology of these large herbivores, in order to better understand their roles and impacts on ecosystems. Furthermore, in order to better predict the spatial and temporal distribution of impacts, it is also important to take into account the individual heterogeneity observed in these patterns within ungulate populations.

For more information:

Garrido, P., Mårell, A., Öckinger, E., Skarin, A., Jansson, A., Thulin, C.-G. 2019. Experimental rewilding enhances grassland functional composition and pollinator habitat use. Journal of Applied Ecology 56:946-955. https://doi.org/10.1111/1365-2664.13338

Milotić, T., Baltzinger, C., Eichberg, C., Eycott, A.E., Heurich, M., Müller, J., Noriega, J.A., Menendez, R., Stadler, J., Ádám, R., Bargmann, T., Bilger, I., Buse, J., Calatayud, J., Ciubuc, .C., Boros, G., Jay‐Robert, P., Kruus, M., Merivee, E., Miessen, G., Must, A., Ardali, E., Preda, E., Rahimi, I., Rohwedder, D., Rose, R., Slade, E.M., Somay, L., Tahmasebi, P., Ziani, S., Hoffmann, M. 2019. Functionally richer communities improve ecosystem functioning: Dung removal and secondary seed dispersal by dung beetles in the Western Palaearctic. Journal of Biogeography 46:70-82. https://doi.org/10.1111/jbi.13452

Baltzinger, C., Karimi, S., Shukla, U. 2019. Plants on the move: hitch-hiking with ungulates distributes diaspores across landscapes Frontiers in Ecology and Evolution 7:38. https://doi.org/10.3389/fevo.2019.00038
Cocquelet, A., Mårell, A., Bonthoux, S., Baltzinger, C. & Archaux, F. 2019. Direct and indirect effects of ungulates on forest birds' nesting failure? An experimental test with artificial nests. Forest Ecology and Management 437:148-155. https://doi.org/10.1016/j.foreco.2019.01.025

Karimi, S., Hemami, M.R., Esfahani, M.T., Baltzinger, C. 2018. Complementary endozoochorous seed dispersal by large mammals in the Golestan National Park, Iran. Seed Science Research 8:294-302. https://doi.org/10.1017/S0960258518000351

Karimi, S., Hemami, M.R., Esfahani, M.T., Baltzinger, C. 2018. The role of Brown bear (Ursus arctos) in the plant seed dispersal of Golestan National Park. Iranian Journal of Applied Ecology 7:29-42. http://ijae.iut.ac.ir/article-1-870-en.html
Liehrmann, O., Jégoux, F., Guilbert, M.A., Isselin-Nondedeu, F., Saïd, S., Locatelli, Y. & Baltzinger, C. 2018. Epizoochorous dispersal by ungulates depends on fur, grooming and social interactions. Ecology and Evolution 8(3): 1582-1594. https://doi.org/10.1002/ece3.3768

Milotić, T., Baltzinger, C., Eichberg, C., Eycott, A. E, Heurich, M., Müller, J., et al. 2018. Dung beetle assemblages, dung removal and secondary seed dispersal: data from a large-scale, multi-site experiment in the Western Palaearctic. Frontiers of Biogeography 10 :e37829. https://doi.org/10.21425/F5101-237289

Pellerin, M., Picard, M., Saïd, S., Baubet, E. & Baltzinger, C. 2016. Complementary endozoochorous long- distance seed dispersal by three native herbivorous ungulates. Basic and Applied Ecology 17(4): 321-332. https://doi.org/10.1016/j.baae.2016.01.005

Picard, M., Chevalier, R., Barrier, R., Boscardin, Y. & Baltzinger, C. 2016. Functional traits of seeds dispersed through endozoochory by native forest ungulates. Journal of Vegetation Science 27(5): 987–998. https://doi.org/10.1111/jvs.12418

Albert, A., Auffret, A., Cosyns, E., Cousins, S., D'hondt, B., Eichberg, C., Eycott, A., Heinken, T., Hoffmann, M., Jaroszewicz, B., Malo, J., Mårell, A., Mouissie, M., Pakeman, R., Picard, M., Plue, J., Poschlod, P., Provoost, S., Schulze, K.A., Baltzinger, C. (2015). Seed dispersal by ungulates as an ecological filter: a trait-based meta- analysis. Oikos 124(9) : 1109-1120. https://doi.org/10.1111/oik.02512
Albert, A., Mårell, A., Picard, M., Baltzinger, C. (2015). Using basic plant traits to predict ungulate seed dispersal potential. Ecography 38:440-449. https://doi.org/10.1111/ecog.00709

Picard, M., Papaïx, J., Gosselin, F., Picot, D., Bideau, E., Baltzinger, C. (2015). Temporal dynamics of seed excretion by wild ungulates: implications for plant dispersal. Ecology and Evolution 5(13): 2621-2632. https://doi.org/10.1002/ece3.1512

Integrated management of wild ungulate populations and their effects

The search for a forest-game balance is part of the objective of sustainable and multifunctional forest management. Based on the concept of adaptive management, we engage in participatory research-action projects involving forest managers, wildlife managers, administrations and scientists. The acquisition and sharing of knowledge, experimentation and dialogue form the basis for a collective evaluation of know-how, which is essential for taking concerted decisions and for the emergence of shared actions. The work carried out is based on indicators for monitoring animal populations, forestry and silviculture indicators, but also on consultation between stakeholders.

In terms of experimentation, new approaches need to be tested. In fact, the hunting management of wild ungulate populations is no longer always sufficient to contain their growth and their impact on the environment. It is therefore necessary to consider complementary solutions with a view to the sustainable management of populations and territories. Approaches based on modifying the behavioural patterns of animals, in the context of a landscape of fear, must now be evaluated.

For more information:

• Bonnot, N. C., Couriot, O., Berger, A., Cagnacci, F., Ciuti, S., De Groeve, J. E., Gehr, B., Heurich, M., Kjellander, P., Kroschel, M., Morellet, N., Sonnichsen, L., Hewison, A. J. M. 2020. Fear of the dark? Contrasting impacts of humans versus lynx on diel activity of roe deer across Europe. Journal of Animal Ecology 89:132-145. https://doi.org/10.1111/1365-2656.13161
• Gehr, B., Bonnot, N.C., Heurich, M., Cagnacci, F., Ciuti, S., Hewison, A.J.M., Gaillard, J.M., Ranc, N., Premier, J., Vogt, K., Hofer, E., Ryser, A., Vimercati, E., Keller, L. 2020. Stay home, stay safe-Site familiarity reduces predation risk in a large herbivore in two contrasting study sites. Journal of Animal Ecology. https://doi.org/10.1111/1365-2656.13202
• Baltzinger et al. (Eds.) 2019. Habitats forestiers et forêts Habitées. Actes de colloques « Habitats forestiers et forêts Habitées » 26-27 mars 2019, Chambord, France, 17 articles, numéro spécial de la Revue Forestière Française, 4-5, p. 329-541. http://documents.irevues.inist.fr/handle/2042/70807
• De Groeve, J., Cagnacci, F., Ranc, N., Bonnot, N.C., Gehr, B., Heurich, M., Hewison, A.J.M., Kroeschel, M., Linnell, J.D.C., Morellet, N., Mysterud, A., Sandfort, R., Van De Weghe, N. 2019. Individual Movement - Sequence Analysis Method (IM-SAM): characterizing spatio-temporal patterns of animal habitat use across landscapes. International Journal of Geographical Information Science 1-22. https://doi.or/10.1080/13658816.2019.1594822
• Baltzinger, M., Mårell, A., Deconchat, M., Barrier, R. 2018. Open game fences and their socio-spatial effects: Placing red deer, placing humans, managing territories. TRACE ∴ Finnish Journal for Human-Animal Studies 4:28-61. https://doi.org/10.23984/fjhas.67671
• Baltzinger, M., Mouche, J., Blondet, M., Hautdidier, B. 2016. Political ecology de l’engrillagement forestier privé en Sologne : quels sont les enjeux socioenvironnementaux au cœur du conflit ? Natures Sciences Sociétés 24:97-108. https://doi.org/10.1051/nss/2016017

Figure 1. Schematic presentation of the interactions studied

The challenge

We study the direct and indirect effects of wild ungulates, but also rodents, on forest and peri-forest ecosystems. Our work provides the scientific basis for the management of wild ungulate populations and their habitats, taking into account the socio-economic consequences of the impacts and services provided by their presence.
We are consolidating our approach to the ecological footprint of wild ungulates within landscape mosaics, integrating societal expectations at the scale of coherent territorial entities and determining how wild ungulates can contribute to the functioning and biodiversity of terrestrial ecosystems as well as to their restoration.

We address the following 4 themes:
•    Forest renewal: How to adapt forest renewal in the presence of wild ungulates in a context of climate change?
•    Biodiversity: What are the effects of wild ungulates on the functioning and biodiversity of forest ecosystems?
•    Functional role: How do wild ungulates change the dynamics of ecosystems?
•    Integrated management: What tools should be proposed to facilitate the management of wild ungulate populations and control their effects on the environment?

Team composition (permanent staff)

The projects (ongoing and last 5 years)

•    SyCoMore (2021-2024): Precision and connected silviculture
•    ReForest (2016-2021): Guarantee the renewal of the forest resource by ensuring the forest-game balance
•    COSTAUD (2016-2018): Contribution of ungulates to ecosystem functioning and services at Chambord
•    EquiForCe 76 (2015-2016): Analysis of the steering process of the forest-cervid balance in Seine-Maritime
•    DYSPERSE (2012-2015): Spatial and temporal dynamics of grazing in Sologne and ecosystem services
•    DIPLO 1 (2009-2011) et DIPLO 2 (2012-2014): Plant dispersal by wild ungulates: a comparative approach to three vectors - deer, roe deer and wild boar
•    OPTMix (2011-) : Oak Pine Tree Mixture experiment