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Time- and sex-specific summer survival of roe deer fawns was estimated using capture-mark-recapture methods in two enclosed populations living in contrasting conditions. The population of Trois Fontaines (eastern France) was roughly constant in size throughout the study period, while in Chizé (western France), the population experienced frequent summer droughts and numbers decreased continuously during the study. Early survival of fawns was low and highly variable over the years at both Chizé and Trois Fontaines, and demonstrated marked variations between cohorts that need to be taken into account when modelling roe deer population dynamics. In Trois Fontaines, fawn survival was positively correlated with early body growth and total rainfall in May and June. In Chizé, fawn survival decreased with increasing density and tended to increase with increasing rainfall in May and June and adult female body mass. These factors explained more than 75% of the variability in early survival observed in both populations. Variation between cohorts had different consequences for the two populations. At Trois Fontaines, cohort variation was limited to a numerical effect on early survival. However at Chizé, cohort variation was long-lasting and affected the phenotypic quality of survivors at later ages, and thereby future survival and breeding abilities (both numerical and quality effects). Male and female fawns had similar survival over their first summer in both populations. This result contrasts with the lower survival of young males often observed in ungulates. Two ultimate causes can be proposed to account for the low and variable survival of roe deer fawns over the first summer: the high energy expenditures incurred by does during each breeding attempt and/or the low absolute body size of newborn roe deer fawns.

Summary1. High-density populations of large herbivores are now widespread. Wildlife managers commonly attempt to control large herbivores through hunting to meet specific management objectives, considering population density as the minimal key source of information. Here, we review the problems of censusing populations of large herbivores and describe an alternative approach, employing indicators of ecological change. 2. Estimating density of large herbivores with high precision and accuracy is difficult, especially over large areas, and requires considerable investment of time, people and money. Management decisions are often made on an annual basis, informed by population changes over the previous year. However, estimating year-to-year changes in density is not a realistic goal for most large herbivores. Furthermore, population density per se provides no information on the relationship between the population and its habitat. 3. For successful management of large herbivores, we need to consider not only the fate of the population, but rather changes in both population and habitat features, as well as their interaction. Managers require information on trends in both the animal population and habitat quality in order to interpret changes in the interaction between these two compartments. 4. We propose that a set of indicators of animal performance, population abundance, habitat quality and/or herbivore habitat impact provides relevant information on the populationhabitat system. Monitoring temporal changes in these indicators provides a new basis for setting hunting quotas to achieve specific management objectives. This sort of adaptive management is employed widely in France for managing roe deer Capreolus capreolus . 5. Synthesis and applications . The management of large herbivores would be improved by investing fewer resources in trying to estimate the absolute abundance of ungulates, and more resources in collecting additional data to inform understanding of the ecological status of the ungulate-habitat system being managed. This paper presents a set of indicators of ecological change for monitoring the interaction between a population and its habitat as a basis for adaptive management to attain explicit goals and to improve knowledge of the system. This approach could improve management for a variety of large herbivores, by harmonizing actions at wide spatial scales.

Summary 1.Equids are generalist herbivores that co-exist with bovids of similar body size in many ecosystems. There are two major hypotheses to explain their co-existence, but few comparative data are available to test them. The first postulates that the very different functioning of their digestive tracts leads to fundamentally different patterns of use of grasses of different fibre contents. The second postulates resource partitioning through the use of different plant species. As domestic horses and cattle are used widely in Europe for the management of conservation areas, particularly in wetlands, a good knowledge of their foraging behaviour and comparative nutrition is necessary. 2. In this paper we describe resource-use by horses and cattle in complementary studies in two French wetlands. Horses used marshes intensively during the warmer seasons; both species used grasslands intensively throughout the year; cattle used forbs and shrubs much more than horses. Niche breadth was similar and overlap was high (Kulczinski's index 0·58-0·77). Horses spent much more time feeding on short grass than cattle. These results from the two sites indicate strong potential for competition. 3. Comparative daily food intake, measured in the field during this study for the first time, was 63% higher in horses (144 g DM kg W -0·75 day -1 ) than in cattle (88 g DM kg W -0·75 day -1 ). Digestibility of the cattle diets was a little higher, but daily intake of digestible dry matter (i.e. nutrient extraction) in all seasons was considerably higher in horses (78 g DM kg W -0·75 day -1 ) than in cattle (51 g DM kg W -0·75 day -1 ). When food is limiting, horses should outcompete cattle in habitats dominated by grasses because their functional response is steeper; under these circumstances cattle will require an ecological refuge for survival during winter, woodland or shrubland with abundant dicotyledons. 4. Horses are a good tool for plant management because they remove more vegetation per unit body weight than cattle, and use the most productive plant communities and plant species (especially graminoids) to a greater extent. They feed closer to the ground, and maintain a mosaic of patches of short and tall grass that contributes to structural diversity at this scale. Cattle use broadleaved plants to a greater extent than horses, and can reduce the rate of encroachment by certain woody species.

Summary1. Maternal care is a major component of demographic tactics in mammals. In ungulates most work has been done on capital breeders (e.g. bighorn sheep), which rely heavily on body reserves to raise their young. Roe deer, in contrast, are close to the income breeder end of the capital±income breeder continuum, and show high levels of maternal care. 2. The aim of this study was to explore the factors determining the level of maternal care in roe deer, in particular the eects of maternal body weight, mother's parity, litter size and year of birth on the amount of prenatal care (i.e. the average mass of an ospring multiplied by the number of ospring) and postnatal care (i.e. the average growth rate of the ospring multiplied by their number). The study was carried out on a captive population of roe deer fed ad libitum, and in a wild population. 3. In both populations prenatal care increased with increasing maternal body weight. In the population fed ad libitum this eect was found in light females only (< 22 kg); in the wild population the positive relationship between maternal body weight and prenatal care was entirely accounted for by variation in litter size (i.e. heaviest females produced larger litters) and density (i.e. females were lightest in years with high population density). Parity did not aect prenatal care. 4. In 14 females fed ad libitum there was no relationship between postnatal care and maternal body weight. Multiparous females had higher levels of postnatal care. In contrast, 20 wild females showed a positive relationship between postnatal care and maternal body weight, and only litter size aected the level of postnatal care. 5. Even after accounting for the confounding eects of maternal body weight, parity, litter size and population density, we found no trade-o between pre-and postnatal care in any of the two populations. The conditions under which the pattern of maternal care could impose trade-os that aect the individual ospring are discussed.

2006. Using a proxy of plant productivity (NDVI) to find key periods for animal performance: the case of roe deer. Á/ Oikos 112: 565 Á/572. Animals in seasonal environments are affected by climate in very different ways depending on season and part of the climatic effects operates indirectly through the plants. Vegetation conditions in spring and summer are regarded as decisive for the reproductive success and the offspring's condition of large herbivores, but objective ways to determine key periods during the growing season have not been done often due to limitations in plant data. Using satellite data (NDVI), we determined how plant productivity from birth to fall influences the following winter body mass of roe deer fawns. We do this in two populations, the first inhabiting the low productive Chizé reserve in south western France with an oceanic climate and the second from Trois Fontaines, a highly productive forest with continental climate in east France. The effect of plant productivity was similar for male and female fawn mass, as expected from the weak intensity of sexual selection in roe deer life history traits. We found contrasting results between sites, with a strong effect of plant productivity in spring (April-May) in the Chizé population, but no effect in the Trois Fontaines population. The relatively low variability in winter fawn body mass could account for the absence of NDVI effects at Trois Fontaines. However, such results might also point to a limitation in the use of the NDVI, since the relationship between the canopy and the plant productivity at the ground level might be weak in the highly productive forest of Trois Fontaines.The role of annual variation in climate for breeding phenology and performance of plants, amphibians and birds in specific populations has recently received much attention (reviewed by Walther et al. 2002. Clearly, the onset of breeding and the reproductive success in seasonal environments for organisms that are either dormant (such as plants and amphibians, Beebee 1995) or migratory over broad scales (for birds, Slagsvold 1976) is coupled with temperature in early summer. Large herbivores are active over the entire year and a better understanding of how animal performance (i.e. survival, reproduction or growth) are linked to plant

Body mass is a key determinant of fitness components in many organisms, and adult mass varies considerably among individuals within populations. These variations have several causes, involve temporal and spatial factors, and are not yet well understood. We use long-term data from 20 roe deer cohorts in a 2600 ha study area (Chizé, western France) with two habitats contrasting in quality (rich oak forest in the North versus poor beech forest in the South) to analyse the effects of both cohort and habitat quality on adult mass (i.e. median body mass between 4 and 10 years of age) of roe deer (Capreolus capreolus). Cohort strongly influenced the adult body mass of roe deer in both sexes: males born in 1994 were 5.2 kg heavier when aged between 4 and 10 years old than males born in 1986, while females born in 1995 were 4.7 kg heavier between 4 and 10 years old than females born in 1982. For a given cohort, adult males were, on average, 0.9 kg heavier in the rich oak forest than in the poor beech forest. A similar trend occurred for adult females (0.5 kg heavier in the oak forest). The effects of cohort and habitat were additive and accounted for ca. 40% of the variation observed in the adult mass of roe deer at Chizé (males: 41.2%; females: 40.2%). Population density during the spring of the birth accounted for about 35% of cohort variation, whereas rainfall in May-June had no effect. Such delayed effects of density at birth on adult body mass probably affect population dynamics, and might constitute a mechanism by which delayed density-dependence occurs in ungulate populations.

We used a data set of ungulate censuses from 31 natural ecosystems from East and Southern Africa to test two hypotheses: (1) megaherbivores should dominate ungulate communities in ecosystems with high rainfall and low soil nutrient status because of their ability to survive on poor quality food resources, and (2) the abundance of megaherbivores affects the abundance of the mesoherbivores, distinguishing the different feeding guilds: mesograzers, mesobrowsers and mesomixed feeders. Two axes of a multivariate analysis (77% of the variance) discriminated the sites well, the first separating sites dominated by megaherbivores from those dominated by mesoherbivores, and the second representing a gradient between mesograzers and mesobrowsers. Our analysis shows (1) that megaherbivores can be considered to be a separate trophic guild and (2) that mesograzers and mesobrowsers respond differently to variation in their trophic environments. The metabolic biomass density of megaherbivores increased with annual rainfall, but was not related to soil nutrient status, and as predicted, megaherbivores comprised a larger proportion of the biomass of ungulate communities in ecosystems with high rainfall and low nutrient soils. The metabolic biomass density of mesoherbivores increased with rainfall and soil nutrient status. Within the mesoherbivores, the metabolic biomass density of mesograzers showed the same trend, and seemed unaffected by megaherbivores. Conversely, mesobrowsers and mesomixed feeders appeared to be unaffected by rainfall or soil nutrient status, but mesomixed feeders declined when megaherbivores were abundant. This suggests that megaherbivores may compete with the mesomixed-feeder species for food or they may alter the vegetation communities unfavourably. A similar analysis using elephants alone instead of megaherbivores as a group showed that both mesobrowsers and mesomixed feeders were affected significantly by elephant, which is consistent with the fact that most of the effect of megaherbivores on browse resources or woodland habitat is due to elephants. This study shows that the different trophic guilds within African ungulate communities react differently to environmental factors (rain and soil), and that megaherbivores, and particularly elephants, appear to compete with mesomixed feeders and mesobrowsers. These results are relevant for the understanding of the functioning of African ungulate communities and call for further testing with longitudinal data.