A. J. Mark Hewison

Successful sons or advantaged daughters? The Trivers-Willard model and sex-biased maternal investment in ungulates

The Trivers-Willard model predicts that in polygynous species, given certain assumptions, mothers with extra resources should bias investment towards sons by adjusting sex ratio at birth and/or by differential provisioning of sons and daughters. Evidence for facultative adjustment of the birth sex ratio from recent empirical studies on ungulates is equivocal at best. However, studies on maternal input and reproductive costs suggest that ungulate mothers do discriminate against daughters during maternal investment in the most polygynous, dimorphic species. Better data on the relationship between maternal input and fitness returns for producing sons or daughters are needed to interpret patterns of sex-biased maternal investment.

Antler size provides an honest signal of male phenotypic quality in roe deer.

Identifying factors shaping secondary sexual traits is essential in understanding how their variation may influence male fitness. Little information is available on the allocation of resources to antler growth in territorial ungulates with low sexual size dimorphism. We investigated phenotypic and environmental factors affecting both absolute and relative antler size of male roe deer in three contrasting populations in France and Sweden. In the three populations, we found marked age‐specific variation in antler size, with an increase in both absolute and relative antler size between yearling and prime‐age stages, followed by a decrease (senescence) for males older than 7 years. Antler size increased allometrically with body mass. This increase was particularly strong for senescent males, suggesting the evolution of two reproductive tactics: heavy old males invested particularly heavily in antler growth (potentially remaining competitive for territories), whereas light old males grew small antlers (potentially abandoning territory defense). Finally, environmental conditions had little effect on antler size: only population density negatively affected absolute antler size in one of the three populations. Antler size may therefore provide an honest signal of male phenotypic quality in roe deer. We discuss the implications of these results in terms of territory tenure and mating competition.

Mismatch between birth date and vegetation phenology slows the demography of roe deer.

This study of a French deer population reveals the demographic costs associated with the failure of a herbivore to modify its life cycle timing to respond to a warming world.

HETEROZYGOSITY-FITNESS CORRELATIONS REVEALED BY NEUTRAL AND CANDIDATE GENE MARKERS IN ROE DEER FROM A LONG-TERM STUDY

Heterozygosity-fitness correlations (HFCs) are increasingly reported but the underlying mechanisms causing HFCs are generally poorly understood. Here, we test for HFCs in roe deer (Capreolus capreolus) using 22 neutral microsatellites widely distributed in the genome and four microsatellites in genes that are potentially under selection. Juvenile survival was used as a proxy for individual fitness in a population that has been intensively studied for 30 years in northeastern France. For 222 juveniles, we computed two measures of genetic diversity: individual heterozygosity (H), and mean d 2 (relatedness of parental genomes). We found a relationship between genetic diversity and fitness both for the 22 neutral markers and two candidate genes: IGF1 (Insulin-like Growth Factor I) and NRAMP (natural resistance-associated macrophage protein). Statistical evidence and the size of genetic effects on juvenile survival were comparable to those reported for early development and cohort variation, suggesting a substantial influence of genetic components on fitness in this roe deer population. For the 22 neutral microsatellites, a correlation with fitness was revealed for mean d 2, but not for H, suggesting a possible outbreeding advantage. This heterosis effect could have been favored by introduction of genetically distant (Hungarian) roe deer to the population in recent times and, possibly, by the structuring of the population into distinct clans. The locus-specific correlations with fitness may be driven by growth rate advantages and resistance to diseases known to exist in the studied population. Our analyses of neutral and candidate gene markers both suggest that the observed HFCs are likely mainly due to linkage with dominant or overdominant loci that affect fitness (“local” effect) rather than to a genome-wide relationship with homozygosity due to inbreeding (“general” effect).

Landscape composition influences roe deer habitat selection at both home range and landscape scales

Understanding how patterns of habitat selection vary in relation to landscape structure is essential to predict ecological responses of species to global change and inform management. We investigated behavioural plasticity in habitat selection of roe deer (Capreolus capreolus) in relation to variable habitat availability across a heterogeneous agricultural landscape at the home range and landscape scales. As expected, woodland was heavily selected, but we found no functional response for this habitat, i.e. no shift in habitat selection with changing habitat availability, possibly due to the presence of hedgerows which were increasingly selected as woodlands were less abundant. Hedgerows may thus function as a substitutable habitat for woodlands by providing roe deer with similar resources. We observed a functional response in the use of hedgerows, implying some degree of landscape complementation between hedgerows and open habitats, which may in part compensate for lower woodland availability. We also expected selection for woodland to be highest at the wider spatial scale, especially when this habitat was limiting. However, our results did not support this hypothesis, but rather indicated a marked influence of habitat composition, as both the availability and distribution of resources conditioned habitat selection. There was no marked between-sex difference in the pattern of habitat selection at either scale or between seasons at the landscape scale, however, within the home range, selection did differ between seasons. We conclude that landscape structure has a marked impact on roe deer habitat selection in agricultural landscapes through processes such as landscape complementation and supplementation.

Testing Global Positioning System Performance for Wildlife Monitoring Using Mobile Collars and Known Reference Points

Abstract To determine the spatial resolution of Global Positioning System (GPS) receiver data, rigorous testing is essential. We tested performance of the Lotek 3300 GPS collar for medium-sized mammals (Lotek Engineering, Inc., Newmarket, ON, Canada). To mimic real wildlife monitoring situations, we performed both static (stationary receiver) and mobile tests, placing the receiver collar on a dog. We compared fix locations of the mobile receiver with the actual trajectory described by a portable Trimble high-precision GPS. We determined performance in relation to habitat type and leaf cover. Location error was habitat-dependent, with the best results in open habitat and much poorer ones in forest, particularly coniferous-dominated forest. For both static and mobile tests, location accuracy was higher when the number of satellites contacted was high and when the residual positional dilution of precision (PDOP) value was low. However, location error was highly variable, even for a given PDOP value and a given number of satellites contacted. Finally, mobile collars performed less well than their static counterparts, presumably because of frequent changes of GPS position and orientation.

Habitat use under predation risk: hunting, roads and human dwellings influence the spatial behaviour of roe deer

Wildlife populations are subjected to increasing pressure linked to human activities, which introduce multiple stressors. Recently, in addition to direct effects, it has been shown that indirect (non-lethal) effects of predation risk are predominant in many populations. Predation risk is often structured in space and time, generating a heterogeneous “landscape of fear” within which animals can minimize risks by modifying their habitat use. Furthermore, for ungulates, resource quality seems to be positively correlated with human-related sources of risk. We studied the trade-off between access to resources of high-quality and risk-taking by contrasting habitat use of roe deer during daytime with that during nighttime for 94 roe deer in a hunted population. Our first hypothesis was that roe deer should avoid human disturbance by modifying their habitat use during daytime compared to nighttime. Our results supported this, as roe deer mainly used open fields during nighttime, but used more forested habitats during daytime, when human disturbance is higher. Moreover, we found that diel patterns in habitat use were influenced by hunting disturbance. Indeed, the roe deer decreased their use of high-crops during daytime, an important source of cover and food, during the hunting season. The proximity of roads and dwellings also affected habitat use, since roe deer used open fields during daytime to a greater extent when the distance to these sources of disturbance was higher. Hence, our results suggest that roe deer resolve the trade-off between the acquisition of high-quality resources and risk avoidance by modifying their habitat use between day and night.

Contradictory findings in studies of sex ratio variation in roe deer (Capreolus capreolus)

Abstract Patterns of sex ratio variation and maternal investment reported in the literature are often inconsistent. This could be due to intra- and inter-specific variation in social systems, but may also be a result of the a posteriori nature of much of this type of analysis or the testing of models which are inappropriate. Two recent papers reported directly opposed results concerning variation in offspring sex ratios in relation to maternal condition in roe deer, interpreting the results as support for the Trivers and Willard model and for the local resource competition hypothesis, respectively. In this paper, we present data on offspring sex ratios and early juvenile body weight from two long-term studies of this species to test predictions arising from these two models concerning sex biases in litter composition and maternal care. First, we observed no consistent pattern of sex differences in an index of weaning weight or body weight at 1 month old in either population, indicating a lack of sex bias in maternal care. However, in one population, higher maternal body weight was associated with higher juvenile body weight of daughters, but not of sons. Secondly, we found a negative, but not statistically significant, relationship between maternal body weight and litter sex ratio such that heavier females tended to produce more daughters and lighter females to produce more sons. These results indicate that roe females which have additional investment potential available do not invest it in sons, as predicted by the Trivers and Willard model. Our results may provide some support that roe deer are subject to local resource competition acting at the level of the individual mother; however, the fact that particular trends in sex ratio data can be explained in functional terms provides no indication that they are actually adaptive.

Landscape fragmentation generates spatial variation of diet composition and quality in a generalist herbivore

Forest fragmentation may benefit generalist herbivores by increasing access to various substitutable food resources, with potential consequences for their population dynamics. We studied a European roe deer (Capreolus capreolus) population living in an agricultural mosaic of forest, woodlots, meadows and cultivated crops. We tested whether diet composition and quality varied spatially across the landscape using botanical analyses of rumen contents and chemical analyses of the plants consumed in relation to landscape metrics. In summer and non-mast winters, roe deer ate more cultivated seeds and less native forest browse with increasing availability of crops in the local landscape. This spatial variation resulted in contrasting diet quality, with more cell content and lower lignin and hemicellulose content (high quality) for individuals living in more open habitats. The pattern was less marked in the other seasons when diet composition, but not diet quality, was only weakly related to landscape structure. In mast autumns and winters, the consumption of acorns across the entire landscape resulted in a low level of differentiation in diet composition and quality. Our results reflect the ability of generalist species, such as roe deer, to adapt to the fragmentation of their forest habitat by exhibiting a plastic feeding behavior, enabling them to use supplementary resources available in the agricultural matrix. This flexibility confers nutritional advantages to individuals with access to cultivated fields when their native food resources are depleted or decline in quality (e.g. during non-mast years) and may explain local heterogeneities in individual phenotypic quality.

Condition‐dependent natal dispersal in a large herbivore: heavier animals show a greater propensity to disperse and travel further

Natal dispersal is defined as the movement between the natal range and the site of first breeding and is one of the most important processes in population dynamics. The choice an individual makes between dispersal and philopatry may be condition dependent, influenced by either phenotypic attributes and/or environmental factors. Interindividual variability in dispersal tactics has profound consequences for population dynamics, particularly with respect to metapopulation maintenance. A better understanding of the mechanisms underlying this variability is thus of primary interest. We investigated the ranging behaviour of 60 juvenile European roe deer, Capreolus capreolus, monitored with GPS collars for 1 year prior to their first reproduction, from 2003 to 2010 in South-West France. Dispersal occurs across a spatial continuum so that dividing individuals into two categories (dispersers vs. philopatric) may lead to information loss. Therefore, to investigate condition-dependent dispersal more accurately, we developed an individual-based measure of dispersal distance, which took into account interindividual variation in ranging behaviour. We assessed the influence of body mass, the degree of habitat heterogeneity and sex on dispersal initiation date, dispersal propensity and distance. The overall population dispersal rate was 0·34, with a mean ± SD linear distance between natal and post-dispersal home ranges of 12·3 ± 10·5 km. Dispersal distances followed a classical leptokurtic distribution. We found no sex bias in either dispersal rate or distance. Forest animals dispersed less than those living in more heterogeneous habitats. Heavier individuals dispersed with a higher probability, earlier and further than lighter individuals. Our individual-based standardised dispersal distance increased linearly with body mass, with some suggestion of a body mass threshold of 14 kg under which no individual dispersed. Natal dispersal in roe deer was thus dependent on both phenotypic attributes and environmental context. Our results suggest that population connectivity can be altered by a change in average body condition and is likely higher in the rich and heterogeneous habitats typical of modern day agricultural landscapes.