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A. J. Mark Hewison CEFS (Comportement et Ecologie de la Faune Sauvage) U.R. I.N.R.A. 0035 24 Chemin de Borde Rouge, Auzeville, CS 52627 31326 Castanet-Tolosan Cedex, France. Tél.: 33(0) 5 61 28 51 23 E-mail: mark.hewison@inra.fr ProdINRA

Training

Ph.D (1993) University of Southampton, UK

HDR (1999) Université Paul Sabatier, Toulouse, France

Current position: Research Director (DR1) at the CEFS (INRA, Toulouse)

Research questions

 I am a behavioural ecologist interested in the interplay between behaviour and demographic performance in large wild herbivores, with a particular focus on understanding inter-individual variation in life history tactics. In most of my research projects, I use the roe deer as the biological model species, but I’m more interested in testing novel questions in behavioural ecology than on describing a new natural history anecdote. The best answers are generally provided by long-term monitoring of individual animals, so I supervise a large-scale capture-mark-recapture programme of roe deer in a rural agricultural setting in south-west France. I also collaborate with others doing similar things on large herbivores in France, as well as across Europe more widely (see http://www.eurodeer.org), so as to i/ be sure that our findings are repeatable, and ii/ explore how large-scale environmental gradients and, by extension, global change impact the behaviour and ecology of wild herbivores.

There are three main components to my research programmes:

 1. Behavioural plasticity in heterogeneous fragmented landscapes:

Initially, the main focus of the Vallons et Coteaux de Gascogne roe deer study was to understand how large wild herbivores adjust to the marked human footprint in fragmented agricultural landscapes. Roe deer are eminently variable in their behaviour and ecology across the wide variety of landscapes they inhabit, which is without doubt one of the reasons they have so successfully recolonised much of Europe in recent years. We have focused on documenting behavioural flexibility at the population level in key behaviours such as habitat selection (Morellet et al. 2011), feeding behaviour (Abbas et al. 2011) and dispersal (Debeffe et al. 2012). Furthermore, we have attempted to link this within-population variation in behaviour to landscape heterogeneity (Hewison et al. 2001, Couriot et al. 2018) and human activity, including disturbance (Padie et al. 2015a, Martin et al. 2018), the road network (Coulon et al. 2008) and hunting (Bonnot et al. 2013, Padie et al. 2015b). Finally, we have investigated the link between landscape heterogeneity, demographic traits (Hewison et al. 2009, Bonnot et al. 2018) and population structure (Coulon et al. 2004, 2006). Based on this research and our comparative work with European colleagues (e.g. Tucker et al. 2018, Peters et al. 2019), we aim to construct models that predict how roe deer populations will respond to ongoing global change.

Some key papers on this subject (see full list):

Abbas et al. (2011) Landscape fragmentation generates spatial variation of diet composition and quality in a generalist herbivore. Oecologia 167, 401-411.
Bonnot et al. (2013) Habitat use under predation risk: hunting, roads and human dwellings influence the spatial behaviour of roe deer. Eur J Wildl Res 59, 185-193.
Coulon et al. (2008) Inferring the effects of landscape structure on roe deer (Capreolus capreolus) movements using a step selection function. Landsc Ecol 23, 603-614.
Coulon et al. (2004) Landscape connectivity influences gene flow in a roe deer population inhabiting a fragmented landscape : an individual-based approach. Mol Ecol, 13, 2841-2850.
Couriot, O. et al. (2018) Truly sedentary? The multi-range tactic as a response to resource heterogeneity and unpredictability in a large herbivore. Oecologia 187, 47–60.
Debeffe et al.. (2012) Condition-dependent natal dispersal in a large herbivore: heavier animals show a greater propensity to disperse and travel further. J Anim Ecol 81, 1327-1337.
Hewison et al (2009) Landscape fragmentation influences winter body mass of roe deer. Ecography 32, 1062-1070.
Hewison, A.J.M. et al (2001) The effects of woodland fragmentation and human activity on roe deer distribution in agricultural landscapes. Can J Zool, 79, 679-689.
Martin et al. (2018) Temporal shifts in landscape connectivity for an ecosystem engineer, the roe deer, across a multiple-use landscape. Landsc Ecol 33, 937–954.
Morellet et al. (2011) Landscape composition influences roe deer habitat selection at both home range and landscape scales. Landsc Ecol 26, 999-1010.
Padié et al. (2015a) Time to leave? Immediate response of roe deer to experimental disturbances using playbacks. Eur J Wildl Res 61, 871-879.
Padié et al. (2015b) Roe deer at risk: teasing apart habitat selection and landscape constraints in risk exposure at multiple scales. Oikos 124, 1536-1546.
Peters et al. (2019) Large herbivore migration plasticity along environmental gradients in Europe: life-history traits modulate forage effects. Oikos 128, 416–429.
Tucker et al. (2018) Moving in the Anthropocene: Global reductions in terrestrial mammalian movements. Science 359, 466-469.

 2. Individual behavioural tactics underlie life history variation

 However, plasticity is not without costs so that an individual often does not display the full population repertoire of behaviours. Much of our recent work has centred on the notion that natural selection is unlikely to favour a single optimum behavioural solution to an ecological problem, but rather that alternative life history tactics can be maintained over evolutionary time due to temporal or spatial variation in fitness payoffs (Monestier et al. 2015). In other words, while it may pay to be bold and aggressive in some circumstances, for instance, when competition for food is fierce, you are better off taking a cautious approach when predators, or hunters, are abundant in your home range (Bonnot et al. 2015). However, limited behavioural plasticity means that an individual cannot be both, so that alternative tactics are favoured depending on the relative strength of the selection pressures for acquiring high quality resources and avoiding predation risk (Benhaiem et al. 2008, Bonnot et al. 2017). These tactics (aka behavioural syndromes, personalities, temperaments, coping styles, or whatever else you want to call them...) are underpinned by consistent inter-individual variation in behavioural (Monestier et al. 2017), physiological (Carbillet et al. 2019, Monestier et al. 2016), and life history (Bonnot et al. 2018) traits, forming an individual-based “pace of life”. For example, dispersing juveniles have an inherently high energy budget (Benoit et al. 2020) and low levels of neophobia (Debeffe et al. 2014), generating alternative movement tactics during this key life history stage (Ducros et al. 2020). We recently also showed that movement tactics can be heritable (Gervais et al. 2020) and, hence, have the potential to evolve in the face of rapid global change.

Some key papers on this subject (see full list):

Benhaiem et al. (2008) Hunting increases vigilance levels in roe deer and modifies feeding site selection. Anim Behav 76, 611-618.
Benoit et al. (2020) Accelerating across the landscape: the energetic costs of natal dispersal in a large herbivore. J Anim Ecol 89, 173-185.
Bonnot et al. (2015) Interindividual variability in habitat use: evidence for a risk management syndrome in roe deer? Behav Ecol 26, 105-114.
Bonnot et al. (2018) Boldness-mediated habitat use tactics and reproductive success in a wild large herbivore. Anim Behav 145, 107-115.
Bonnot et al. (2017) Stick or twist: roe deer adjust their flight behaviour to the perceived trade-off between risk and reward. Anim Behav 124, 35-46.
Carbillet et al. (2019) Using the N:L ratio to highlight individual coping styles across variable environments in the wild. Behav Ecol Sociobiol 73, 144.
Debeffe et al. (2014) The link between behavioural type and natal dispersal propensity reveals a dispersal syndrome in a large herbivore. ProcRoy Soc B 281, doi: 10.1098/rspb.2014.0873.
Ducros et al. (2020) Beyond dispersal versus philopatry? Alternative behavioural tactics of juvenile roe deer in a heterogeneous landscape. Oikos 129, 81-92.
Gervais et al. (2020) Pedigree-free quantitative genetic approach provides evidence for heritability of movement tactics in wild roe deer. J Evol Biol 33, 595-607.
Monestier et al. (2015) Is a proactive mum a good mum? A mother’s coping style influences early fawn survival in roe deer. Behav Ecol 26, 1395-1403.
Monestier et al.  (2016) Individual variation in an acute stress response reflects divergent coping strategies in a large herbivore. Behav Proc 132, 22-28.
Monestier et al. (2017) Neophobia is linked to behavioural and haematological indicators of stress in captive roe deer. Anim Behav 126, 135-143.

 3.Mating systems and reproductive allocation

I was first interested in how individuals allocate their effort to reproduction during my PhD (Southampton Unversity 1993) when I studied the link between sexual dimorphism, degree of polygyny and sex allocation tactics in large wild herbivores (Hewison & Gaillard 1999). Most of the work was based on observation of birth sex ratios (Hewison & Gaillard 1996, Hewison et al. 1999) or post-natal growth (Hewison et al. 2005) in wild populations, but I did start some experimental work on captive roe deer, with promising results, which I would like to explore again sometime in the future in our dedicated roe deer enclosures (see http://www.toulouse.inra.fr/Outils-et-Ressources/Unites-Experimentales/IE-Gardouch/%28key%29/4).
Since then, I have supervised or co-supervised PhD projects on the role of sexual selection in determining both female (Debeffe et al. 2014, Vanpé et al. 2009a, 2019) and male (Vanpé et al 2007, 2008, 2009b,c, Lemaitre et al. 2018) mating tactics and allocation to reproduction (Hewison et al. 2011). For example, from long-term monitoring, we were able to identify a lack of plasticity in roe deer birth date (Plard et al. 2013), leading to an ever more marked mismatch between resource availability and energy requirements due to earlier and earlier springs (Plard et al. 2014). This mismatch has driven a marked decrease over time in individual fitness (Plard et al. 2015) and population growth in forest habitat (Gaillard, Hewison et al. 2013), We have suggested that this might be the driving mechanism behind the rapid colonisation of more open landscapes by roe deer over the last half century, as they seek to escape increasingly unfavourable conditions during spring by intensifying their use of the fertilised agricultural matrix (Hewison et al. 2009).

Some key papers on this subject (see full list):

Debeffe et al. (2014) A one night stand? Reproductive excursions of female roe deer as a breeding dispersal tactic. Oecologia 176, 431-443.
Gaillard et al. (2013) How does climate change influence demographic processes of widespread species? Lessons from the comparative analysis of contrasted populations of roe deer. Ecol Lett 16, 48-57.
Hewison, A.J.M. & Gaillard, J.M. (1996) Birth sex-ratios and local resource competition in roe deer. Behav Ecol, 7, 461-464.
Hewison, A.J.M. & Gaillard, J.M. (1999) Successful sons or advantaged daughters? The Trivers-Willard model and sex-biased maternal investment in ungulates Trends Ecol Evol, 14, 229-234.
Hewison et al. (1999) Contradictory findings in studies of sex-ratio variation in roe deer. Behav Ecol Sociobiol, 45, 339-348.
Hewison et al. (2002) Maternal age is not a predominant determinant of progeny sex ratio variation in ungulates. Oikos, 98, 334-339.
Hewison et al. (2005) Big mothers invest more in daughters - reversed sex allocation in a weakly polygynous mammal. Ecol Lett, 8, 430-437.
Hewison et al.  (2011) Reproductive constraints, not environmental conditions, shape the ontogeny of sex-specific mass-size allometry in roe deer. Oikos 120, 1217-1226.
Lemaître et al. (2018) The influence of early-life allocation to antlers on male performance during adulthood: Evidence from contrasted populations of a large herbivore. J Anim Ecol 87, 921-932.
Plard et al. (2013) Parturition date for a given female is highly repeatable in five roe deer populations. Biol Lett 9, article number 20120841.
Plard et al. (2014) Mismatch between birth date and vegetation phenology slows the demography of roe deer. PLoS Biology 12, e1001828. doi:10.1371/journal.pbio.1001828.
Plard et al. (2015) The influence of birth date via body mass on individual fitness in a long-lived mammal. Ecology 96, 1516-1528.
Vanpé et al. (2007) Antler size provides a honest signal of male phenotypic quality in roe deer. Am Nat 169, 481-493.
Vanpé et al. (2008) Mating system, sexual dimorphism and the opportunity for sexual selection in a territorial ungulate. Behav Ecol 19, 309-316.
Vanpé et al. (2009a) Multiple paternity occurs with low frequency in the territorial roe deer, Capreolus capreolus. Biol J Linn Soc 97, 128-139.
Vanpé et al. (2009b) Age-specific variation in male breeding success of a territorial ungulate species, the European roe deer. J Mammal 90, 661-665.
Vanpé et al. (2009c) Access to mates in a territorial ungulate is determined by the size of a male’s territory, but not by its habitat quality. J Anim Ecol 78, 42-51.
Vanpé et al. (2019) Old females rarely mate with old males in roe deer, Capreolus capreolus. Biol J Linn Soc 128, 515–525.