In red deer, yearling antler length is a largely nutrition-mediated phenotypic character, and is therefore sometimes used as an indirect estimate of range quality and population condition. However, the parameters affecting yearling antler length have been little studied. We analyse the contributions of density, weather and maternal effects on yearling antler length of 581 individual stags born 1970-1996 on the Isle of Rum (Scotland). We show that antler length is a good measure of yearling condition: the probability of overwinter survival in yearlings that developed antlers was 3 times higher than for yearlings that did not develop antlers, and yearling antler length was correlated with the number of antler points the following year. Between years, variation in yearling antler length was best explained by variation in red deer density and June temperature at 12 months of age. Both of these variables were negatively correlated with antler length, and most likely this effect is due to changes in nutrient availability. Population density affects biomass availability for the individual, while low temperatures in early summer prolong the availability of high forage quality. At the individual level, antler length increased with birth weight and decreased with birth date, reflecting the persistent and pervasive influence of conditions in early life.

Storaas, T. (2014). To feed or not to feed? Evidence of the intended and unintended effects of feeding wild ungulates.Journal of Wildlife Management, 78(8) Large ungulates are keystone species in many ecosystems and are a highly valued natural 47 resource for social, cultural, and economic reasons (Gordon et al. 2004). Many populations have grown over recent decades, causing environmental and socio-economic 49 impacts on wider communities and conservation concerns, as well as density-dependent 50 changes in performance (Côté et al. 2004, Putman et al. 2011. Consequently, conflicts 51 arise between stakeholders who benefit economically from high ungulate numbers (e.g., 52hunters, outfitters, tourism operators) and those faced with the costs of their presence 53(e.g., land managers, conservationists, transport users and operators; Austin et al. 2010). 54Manipulating forage availability through the provision of additional food could 60Wildlife can be fed by humans in many different ways, intentionally (e.g., at 61 feeding stations) and unintentionally (e.g., unprotected agricultural crops and rubbish 62 dumps; Oro et al. 2013, Sorensen et al. 2014. In this review, we focus on the effects of 63 intentional feeding of ungulates, specifically considering supplementary feeding (often 64 called winter feeding) and diversionary, or intercept, feeding used to attract animals away 65 from vulnerable vegetation, livestock, or major traffic arteries (defined fully in 66 Supplemental Material). However, many of the issues raised also apply across a broad 67 range of unintentional but predictable anthropogenic food subsidies (Oro et al. 2013). 68Earlier reviews of the consequences of feeding large herbivores, published a decade or 77 METHODS 78We reviewed articles in the peer-reviewed and, to a lesser extent, grey literature that our search terms resulted in 232 articles. 86We identified 5 major management goals of supplementary and diversionary 87 feeding and considered the intended effects of feeding that would allow these goals to be 88 achieved (Table 1). We also identified commonly perceived unintended or secondary 89 consequences of feeding ( Table S1). 152Among adult females, an effect of feeding is complicated by the strong impact of 153 reproductive status on autumn mass (Cook et al. 2013 and higher neonatal growth rates (Jacobsen et al. 1981, Smith et al. 1997). 160The reproductive rate (number of offspring per adult female in summer or 161 autumn) of supplementally fed herbivores increased in 5/7 studies ( were used (Gundersen et al. 2004, Sahlsten et al. 2010 feeding site users and non-users (van Beest et al. 2010b). 282Of 16 controlled studies, 6 showed diversionary feeding to be effective in 283 reducing damage in the targeted areas, whereas 4 showed a significantly increased 284 impact, opposing the management goal ( we found no studies that formally addressed this ( 379In studies addressing the increased probability of browsing or grazing impacts in 380 response to feeding (4/4 studies; Table 2 and Supplem...

In winter, red deer (Cervus elaphus hippelaphus) that are not receiving supplementary feeding emphasize and rely on energy-saving strategies. In severe winters with deep snow, they select a home-range habitat-windblown alpine pastures above the timberline-that meets the requirements of this strategy: a much lower sinking depth and good forage availability are due to patchy snow distribution, while the open area provides higher radiation, permits group formation and thus sociobiological wellbeing. The energy costs of thermoregulation are minimized by selecting slopes with south to south-west exposures, seeking thermal shelter from strong winds, while high flight distances and horizontal flight routes reduce the energy costs of fleeing. In years with little winter snowfall non-supplemented red deer prefer the greater security of forested habitats in lower areas and obtain higher food intake from slightly more nutritious meadows in the valley bottom. The food-supplemented herd has much more constant and predictable habitat use, combining an energy-saving strategy-choosing alpine pastures as low-cost winter home range-with high energy income at the feeding station.

In red deer, variation in winter and spring weather conditions encountered by the mothers during pregnancy and during the first year of life are a main determinant for individual life‐history as well as population dynamics. We tested the hypothesis that supplementary feeding which provides constant food supply throughout winter removes the selective pressure of winter harshness on nutrition‐mediated phenotypic traits. We analysed cohort variation in body weight in calves in October, before their first winter, and in yearlings in June, after their first winter, in a food‐supplemented population in the Eastern Austrian Alps. Over eleven years, cohort body weight varied between years in calves and yearlings. Contrary to studies on non‐supplemented red deer populations we found neither short‐ nor long‐term effects of winter weather on body weight. In calves, autumn body weight was negatively related to April–May and June temperatures, suggesting that cool weather during the main growth period retarded plant senescence and thereby prolonged the period of high protein content of summer forage. In yearlings, variation in June body weight, shortly after the end of the feeding period, was lower after a wet April–May, suggesting a negative effect of a prolonged period of supplemental feeding. For both calves and yearlings intra‐cohort variation in body weight was higher, inter‐cohort variation was lower as compared to non‐supplemented red deer, suggesting that in their first year of life supplemented red deer are under reduced natural selection pressure.