Supplemental feeding reduces natural selection in juvenile red deer

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.     

Winter severity or supplementary feeding—which matters more for wild boar?

During recent decades, wild boar have successfully colonised areas previously believed to be unsuitable for the species. Estonia lies close to the northern limit of the species range, and mast foods, which are a common natural food source for many wild boar populations, are practically absent. We hypothesised that the proportion of cultivated area and especially supplementary feeding, which is widely used in Estonia to lower the winter mortality of game species, play important roles in sustaining the local wild boar population. To determine the most important factors shaping the abundance of wild boar in a northern environment, we developed mixed models to account for variation in an index of wild boar abundance (AI) based on winter track counts. The abundance of supplementary feeding sites and mean January temperature were the most important factors determining wild boar winter abundance. We conclude that the current high local population density is sustained by intensive supplementary feeding, which has exceeded the limiting effect of harsh winters.     

The effect of summer feeding on juvenile arctic fox survival - a field experiment

The arctic fox Alopex lagopus L population in Sweden is small and its numbers fluctuate widely with food availability, l e rodent populations This fluctuation is mediated through differences in recruitment rates between years The recruitment can be divided into three phases number of litters born, number of cubs per litter and cub survival rates The number of litters and their sizes have been shown to depend on food availability during winter and spring To examine cub survival during the summer and how it relates to food availability, we conducted a feeding experiment m northern Sweden during 1990, a year of low rodent density, involving six occupied arctic fox dens Feeding at dens lowered cub mortality rates However, condition and growth rates of juveniles were not influenced by supplementary feeding at dens, nor were they related to the probability of survival for an individual Thus arctic foxes seem to minimize risks rather than maximize growth The juvenile mortality from weaning and over the next 6 wk was 21%, mostly due to starvation Only 82% survived from weaning to the first breeding season Of the one-year-old foxes, 50% survived their second year Supplementary feeding of juveniles had no effect on the final survival rates over these two years However, the immediate, positive effect on cub survival could be used in a long-term, extensive management programme if combined with winter feeding     

Early growth and sucking behaviour of Soay sheep in a fluctuating population

Annual variation in growth rate and sucking behaviour of lambs was studied in a fluctuating population of Soay sheep on Hirta, St. Kilda. Growth rates during the summer declined with increasing population size in the previous autumn despite an increase in time spent sucking in the first six weeks of life. The increase in time spent sucking was due largely to an increase in sucking frequency and, since a higher proportion of sucking attempts were rejected, probably reflected that lambs obtain little milk as a result of poor maternal condition following winters at high population density. After accounting for the density dependent variation, annual differences in lamb growth rates were positively related to the increase in biomass of live grasses and forbs between spring and summer. Since food availability was influenced strongly by the numbers surviving the winter, lambs born after the very high mortality at peak population density were able to compensate to some extent for their poor initial growth. However, improved food availability was never sufficient to facilitate complete catch-up growth and may permanently influence adult size and reproduction.     

Experimental evidence for the influence of food availability on incubation attendance and hatching asynchrony in the Australian reed warbler Acrocephalus australis

The amount of time a bird allocates to incubation is likely to be limited by energetic constraints. If food is abundant, energetic constraints may be reduced and the time spent incubating (incubation attendance) may increase. Moreover, the onset of incubation in relation to clutch completion may be advanced, resulting in a higher degree of hatching asynchrony. We measured the effect of experimentally increased food availability on incubation attendance and an estimate of hatching asynchrony in the Australian reed warbler Acrocephalus australis . Supplementary food was provided every other day, from a few days before the start of egg laying until just prior to hatching. Incubation attendance was measured with temperature loggers at nests receiving supplementary food and control nests. Hatching asynchrony was inferred from mass and size differences between siblings shortly after hatching. We found that 1) food supplementation resulted in an increase in incubation attendance, when comparing both nests receiving supplementary food to control nests as well as feeding to non-feeding days in nests receiving supplementary food, and 2) food supplementation resulted in a greater hatching asynchrony, without affecting clutch size, average egg volume or the likelihood of eggs hatching. This suggests that food availability acts in a proximate way to modify the extent of incubation attendance and hatching asynchrony. We discuss the adaptive significance of increased incubation attendance and a shift in the degree of hatching asynchrony in relation to food availability.     

Plant community dynamics in a calcareous grassland under climate change manipulations

This study investigates the effects of field manipulations of local climate to determine the potential impact of climate change on plant community dynamics in a calcareous grassland. The experimental site is located in a grassland at the Wytham estate, Oxfordshire, UK. The one hectare study area is within a 10 ha abandoned arable field on Jurassic corallian limestone. Two climate change scenarios were used: warmer winters with increased summer rainfall and warmer winters with summer drought. Plant cover and species richness were significantly increased in plots receiving supplemented summer rainfall, while the amount of litter was significantly reduced. Litter formation was significantly increased by winter warming and drought. The responses of the plant community to the climate manipulations were related to the life-history attributes of the dominant species. Seedling recruitment was limited by microsite availability, which also varied in the different climate manipulations. The results are discussed in terms of successional dynamics. They suggest that warmer winters may delay succession, as gap formation in the sward will provide sites for colonisation of annuals, thereby enabling their persistence in the sward. Under wetter conditions during summer, perennial grasses tend to close the sward, thereby inhibiting the establishment of later successional species.     

The effect of mollusc grazing on seedling recruitment in artificially created grassland gaps

Two experiments conducted in spring and autumn 1992 examined the effect of mollusc grazing on seedling regeneration from natural grassland seedbanks by creating artificial gaps in plots in a grassland sward. Molluscs were excluded from half the gaps by application of molluscicide. Mollusc grazing in both the spring and autumn experiment significantly reduced seedling recruitment, though the intensity of grazing was greatest in autumn. Recruitment of five species was markedly influenced by molluscicide application. In spring, plots from which molluscs were excluded contained significantly more seedlings of Chenopodium polyspermum and Ranunculus acris. In the autumn, exclusion of molluscs resulted in increased populations of R. acris, Stellaria graminea and Rumex acetosa. Cerastium holosteoides populations were greatest in autumn grazed plots. Other species, notably the grasses Holcus lanatus and Agrostis capillaris and the legume Trifolium repens were unaffected by molluscicide application. Species diversity was significantly decreased by molluscicide application in the autumn. Gap size significantly affected the recruitment of two species. Ranunculus acris populations were significantly higher in small gaps in both spring and summer, while Chenopodium recruitment in the spring was greater in small gaps. Gap size also significantly influenced the risk of mollusc attack on Ranunculus as molluscs appeared to show an aggregative feeding response in the high seedling density small gaps. Selective grazing of vulnerable seedlings by molluscs may influence the eventual relative proportions of the species present and so provide a potent mechanism in shaping community composition in grasslands.     

Experimental evidence of cost of lactation in a low risk environment for a long-lived mammal

In a previous experiment we have documented that organisms adopt a risk-sensitive reproductive allocation when summer reproductive investment competes with survival in the coming winter ( Bårdsen et al. 2008 ). This tradeoff is present through autumn female body mass, which acts as an insurance against unpredictable winter environmental conditions. We tested this hypothesis experimentally on female reindeer experiencing stable and benign winter feeding conditions. Additional supplementary feeding and removal of newborns represented two sets of experimental manipulations. Females in the supplementary feeding group increased more in winter body mass relative to control females. This manipulation, however, did not have any effect on summer body mass development for neither females nor offspring, but we found a positive effect of feeding on offspring birth mass for smaller females. In contrast, offspring removal did have a positive effect on summer body mass development as females in this group were larger in the autumn relative to control females. In essence, we documented two immediate effects as: (1) supplementary feeding did have a positive effect on spring body mass for smaller females; and (2) offspring removal did increase the female summer somatic growth as this had a positive effect on female autumn body mass. Additionally, we tested for lagged effects, but we could not document any biologically significant effects of neither manipulation in the coming spring. The fact that we only found rather weak effects of both manipulations was as expected for risk sensitive individuals experiencing benign environmental conditions over many years.     

Relative Roles of Grey Squirrels,Supplementary Feeding,and Habitat in Shaping Urban Bird Assemblages

Non-native species are frequently considered to influence urban assemblages. The grey squirrel Sciurus carolinensis is one such species that is widespread in the UK and is starting to spread across Europe; it predates birds’ nests and can compete with birds for supplementary food. Using distance sampling across the urbanisation intensity gradient in Sheffield (UK) we test whether urban grey squirrels influence avian species richness and density through nest predation and competition for supplementary food sources. We also assess how urban bird assemblages respond to supplementary feeding. We find that grey squirrels slightly reduced the abundance of breeding bird species most sensitive to squirrel nest predation by reducing the beneficial impact of woodland cover. There was no evidence that grey squirrel presence altered relationships between supplementary feeding and avian assemblage structure. This may be because, somewhat surprisingly, supplementary feeding was not associated with the richness or density of wintering bird assemblages. These associations were positive during the summer, supporting advocacy to feed birds during the breeding season and not just winter, but explanatory capacity was limited. The amount of green space and its quality, assessed as canopy cover, had a stronger influence on avian species richness and population size than the presence of grey squirrels and supplementary feeding stations. Urban bird populations are thus more likely to benefit from investment in improving the availability of high quality habitats than controlling squirrel populations or increased investment in supplementary feeding.     

Energy or information? The role of seed availability for reproductive decisions in edible dormice

The edible dormouse is a specialized seed predator which is highly adapted to the fluctuations of food availability caused by mast seeding of beech and oak trees. Dormice produce young just in time with maximum food availability, and can completely skip reproduction in years with a lack of seeding. Because their decision to reproduce or not in any particular year is made long before the ripe seeds are available, it seems that dormice can anticipate the upcoming mast situation. We tested the hypothesis that the presence of high caloric food in spring affects their reproductive decision. Therefore, we supplementary fed dormice in a field experiment from spring to early summer with sunflower seeds, which also contain a high amount of energy. Supplemental feeding caused significant increases in the proportion of reproducing females and reproductively active males. These results suggest that edible dormice may use the occurrence of an energy rich food resource to predict the autumnal mast situation. Further, our data indicate that the decision to reproduce was not the result of an increased body mass due to the consumption of surplus food, but that sufficient seed abundance acts as an environmental signal to which dormice adjust their reproduction.     

Winter ecology of nonmigratory Alpine red deer

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.     

Overwintering survival and spring emergence inMeligethes aeneus: Effects of body weight, crowding, and soil treatment withBeauveria bassiana

High overwintering mortality was recorded forMeligethes aeneus in the field in Finland, ranging from 85 to 98% depending on the source of the overwintering population. The main factor explaining, variation in survival was the body weight of the beetles in the autumn: only the heaviest beetles survived. Body weight, on the other hand, was influenced by treatment of the rape field soil withBeauveria bassiana, resulting in a 50% decrease in winter survival compared with the reference. The soil treatment did not appear to have direct mortality effects (pathogenesis) on the beetle neither in the summer nor in the winter, and there was no apparent effect on survival of crowding of the insects at the overwintering site. Density dependence in overwintering survival ofM. aeneus may function through the availability of pollen food in the autumn, and intraspecific competition for it. Treatment of the rape field soil withB. bassiana may be a feasible way to increase, the winter mortality and to lower the pollen beetle populations, but requires further research.     

Body weight as an indication of density-dependent population regulation in badgers (Meles meles) at Woodchester Park, Gloucestershire

The body weights of badgers were examined to look for density-dependent effects of the increase in group size at Woodchester Park, Gloucestershire. The weight of badgers in 21 groups were studied from 1978–1993. A significant negative relationship was found between weight and group size in breeding adult females in autumn and winter, and adult males in summer, such that increase in group size caused a decrease in weight. However, these relationships were affected by year-to-year changes and random variation, and overall there was no downward trend in body weight. The existence of density-dependent effects on badger populations at high density was consistent with previous findings     

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.     

Variation in the size of foxes in Scotland

Foxes become larger from south to north in Scotland, independently of the climate, the prey taken or the productivity of the areas in which each population lives. In areas where fluctuating vole populations are important as food, foxes born in high vole years are no larger than those born in poor years. However foxes became smaller after a severe winter in north-east Scotland, followed by a gradual increase in the size of each year class through a series of years with mild winters and expanding rabbit populations. It is suggested that food availability is determining the average size of foxes through selection, and the north/south cline in size is the result of increased hunting hours at higher latitudes during winter. The selective advantage of different size animals under conditions of different food availability is discussed.     

Snow depth does not affect recruitment in a low-density population of boreal woodland caribou (<Emphasis Type="Italic">Rangifer tarandus caribou</Emphasis>)

Winter snow depth may be an important driver of annual variability in recruitment of ungulate calves, and low calf recruitment has been implicated as a factor in declining boreal caribou (Rangifer tarandus caribou) populations. We used 11 consecutive years (2006–2016) of aerial survey data to document calf recruitment in a low-density population of boreal woodland caribou in the Northwest Territories, Canada. We measured snow depth in winter and tested two hypotheses: (1) that calf recruitment was lower in winters with greater snow depth and (2) that calf recruitment was lower following winters with greater snow depth (1-year time lag). Recruitment, the number of calves/adult female in March, ranged twofold from 0.23 to 0.45, and snow depth also ranged twofold from 41 to 85 cm. Yet, we found no support for the hypothesis that late-winter snow depth in the current or previous year was inversely related to calf recruitment.     

Growth in field and laboratory populations of Wood mice (Apodemus sylvaticus)

Intrinsic growth rate and asymptotic body weight parameters were estimated for two laboratory and three field populations of the Wood mouse, Apodemus sylvaticus , using a logistic model fitted by an ordinary least squares method. The data upon which the growth equations were based were differences in body weight for individual animals between sampling occasions. Suckling mice in one of the laboratory studies suffered high mortalities and poor body growth which was related to disturbance to the mothers and their young during examination. Growth in weaned mice appeared normal and was compared with growth in autumn and winter populations of wild mice with poor and good natural seed supplies. Males tended to have higher growth rates, and to a lesser extent, higher body weight asymptotes than females in all groups. Growth rates were highest in the autumn field population with a good seed supply; differences between the other groups were less marked. These results are discussed and it is suggested that, despite the approximations inherent in the method, the method will be a useful tool for studying the the productivity of wild populations of small mammals.     

Diet of the mara (Dolichotis patagonum), food availability and effects of an extended drought in Northern Patagonia (Mendoza,Argentina)

The mara is a near threatened Caviomorph, endemic to Argentina. Studies on feeding ecology allow assessment of the dietary adaptability of maras to habitat changes. The mara diet and food availability on two sampling sites, belonging to distinct landscapes of Northern Patagonia, were estimated using microhistological analysis and point-quadrat transects, over four seasons, and besides during an extended drought. Significant differences were detected by Kruskall-Wallis ANOVA, feeding selection by the χ² test, and dietary preferences by Bailey's confidence interval. Grasses dominated food availability, with Panicum and Poa as major species, followed by shrubs and scarce forbs. Plant cover and forbs increased in spring and summer. The drought caused a strong decrease in plant cover and proportion of grasses. Maras ate all grass species, most forbs and several shrubs. Grasses dominated the diet, with Poa and Panicum being the major species, supplemented by the shrubs Lycium and Prosopis. Maras ate more grasses and forbs in spring and summer, and shrubs in autumn and winter. More shrubs and forbs, and less grasses, were eaten during the drought. Plant categories were used selectively only in autumn and winter, and in the drought period, with preference for shrubs and avoidance of grasses. Bromus, Poa, Plantago and Prosopis were preferred, and Panicum avoided. The mara qualified as a grazer but shifted to a mixed feeder during the drought. Dolichotis patagonum shared habitats with several big and medium-sized herbivores and showed the highest dietary similarities with plain vizcachas, brown hares and horses. Protective measures for natural habitats are needed, given that increasing impacts on food resources and habitat quality could be threats to the survival of maras and other wild vertebrates in Northern Patagonia.     

Reproductive success and failure: the role of winter body mass in reproductive allocation in Norwegian moose

A life history strategy that favours somatic growth over reproduction is well known for long-lived iteroparous species, especially in unpredictable environments. Risk-sensitive female reproductive allocation can be achieved by a reduced reproductive effort at conception, or the subsequent adjustment of investment during gestation or lactation in response to unexpected environmental conditions or resource availability. We investigated the relative importance of reduced investment at conception compared with later in the reproductive cycle (i.e. prenatal, perinatal or neonatal mortality) in explaining reproductive failure in two high-density moose (Alces alces) populations in southern Norway. We followed 65 multiparous, global positioning system (GPS)-collared females throughout the reproductive cycle and focused on the role of maternal nutrition during gestation in determining reproductive success using a quasi-experimental approach to manipulate winter forage availability. Pregnancy rates in early winter were normal (≥0.8) in all years while spring calving rates ranged from 0.4 to 0.83, with prenatal mortality accounting for most of the difference. Further losses over summer reduced autumn recruitment rates to 0.23–0.69, despite negligible predation. Over-winter mass loss explained variation in both spring calving and autumn recruitment success better than absolute body mass in early or late winter. Although pregnancy was related to body mass in early winter, overall reproductive success was unrelated to pre-winter body condition. We therefore concluded that reproductive success was limited by winter nutritional conditions. However, we could not determine whether the observed reproductive allocation adjustment was a bet-hedging strategy to maximise reproduction without compromising survival or whether females were simply unable to invest more resources in their offspring.