Temporal variation in density dependent body growth of a large herbivore

Temporal variation both due to density dependent and density independent processes affect performance and vital rates in large herbivores. Annual fluctuations in climate affect foraging conditions and thus body growth of large herbivores during the short growing season in alpine habitats. Also, high animal densities on summer ranges may increase competition for food and reduce body mass gain. Yet, little is known about interactive effects of density and climate on alpine summer ranges, and the time scales these processes operate on. In this fully replicated landscape‐scale experiment, we kept domestic sheep at high and low densities over nine grazing seasons in an alpine habitat, and tested the relative role of density and annual variation for lamb body mass gain during summer and whether effects of density and annual variation interacted. We found that lambs at high density gained less mass over the summer season than lambs at low density. At short time scales the density effect interacted with annual fluctuations in body growth. We documented a long‐term temporal trend in body mass, consistent with the hypothesis that grazing effects affect habitat differentially at high and low density over years. At high density lamb autumn body mass declined during the first three grazing seasons and then stabilized, whereas body mass slightly increased over years at low density. This long‐term trend suggests accumulative density dependent effects from a biomass or quality reduction, and hence delayed food competition at high density and possibly facilitation at low density. Our experiment provides new insight into how density dependent effects on performance of a large herbivore depend on temporal scale of observation.     

Experimental evidence of density dependent activity pattern of a large herbivore in an alpine ecosystem

Density dependent processes affecting foraging strategies may in turn influence vital rates and population regulation in large herbivores. Increased competition may lower both forage availability and quality, but whether the main activity constraint at high density is increased searching time or increased digestion time is poorly investigated. In a fully replicated landscape‐scale experiment, we used long‐term data (2003–2009) from domestic sheep grazing at high and low density (80 and 25 sheep km^–2, respectively) on alpine summer ranges to test density dependence in allocation of time to feeding (moving) vs digestion (resting) activities and how this in turn affected body growth. Sheep at high density spent more time actively feeding than sheep at low density, but sheep moved shorter distances while foraging at high density. Increased activity levels at high density suggest that the main activity constraint at high density was availability of high‐quality food increasing searching time and possibly reducing intake rates. Increased movement distances at low density is consistent with a higher selection for more productive vegetation types since high‐quality patches are dispersed in the landscape. The alternative hypothesis, that food processing time increased at high density was not supported as it would have reduced overall activity levels. Individual activity levels increased body growth, but this was not sufficient to fully compensate for lower habitat quality leading to an overall reduced body growth at high density. Our experiment clearly documents changes in activity budgets and movement distances of a large herbivore at high population density, providing one potential behavioural mechanism of density dependent responses observed in vital rates.     

Effect of resource gradient on age and size at maturity and their influence on early‐life fecundity in the predatory Asian lady beetle,Harmonia axyridis

Variation in food availability impacts the performance of insects in terms of their size and age to maturity and fecundity. Age at maturity determines how quickly individuals in a population can start to reproduce and how much they can reproduce. Results from studies on various insect species show that food availability influences the size and fecundity of adult females. It is predicted that under poor growth conditions, variation in size is low, but variation in age at maturity is considerable. This prediction was examined in a widely distributed lady beetle species, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), a predator of aphids and coccids. Using a food gradient from low to high aphid prey density, performance of females that were reared on excess food was recorded for pre‐reproductive duration, size at reproductive maturity, number of aphids consumed, and fecundity in the first 10 days of their reproductive period. Results suggested that female H. axyridis that were reared on surplus food when kept at low prey density (poor growth condition) took, on average, three times longer to attain maturity and produced, on average, 14 times fewer eggs than females that were also reared on surplus food, but kept at high prey density (good growth condition). Females performed best at a prey density of 30 aphids per female per 150 cm². Results suggested that the current food availability significantly influenced the age and size of females at maturity and their fecundity. Age and size at maturity of female lady beetles showed non‐linear responses to prey density as well as the occurrence of a minimum size of females, below which H. axyridis females fail to mature. The steep slope recorded at lower prey densities suggests relatively high variation in age at maturity but low variation in size.     

Factors influencing home range and activity of wild boar (Sus scrofa) in a Mediterranean coastal area

This paper analyses the temporal variation in the size of home range and activity of adult wild boar ( Sus scrofa L.). Eighteen boar were radiotracked between 1991 and 1993 in the Maremma Natural Park (Central Italy). Variations of home-range size and activity were related to sex, winter food availability and population density. The hypotheses that home-range size was inversely related to food availability and population density, and that the amount of activity was directly related to population density and inversely related to food abundance, were tested. The size of the acorn crop was used as a measure of winter resource level. No differences were found in the size of ranges, core area, and activity between males and females. In 1991–92 home-range size was larger than in 1992–93; conversely, activity increased from 1991–92 to 1992–93. Winter food availability was high in 1991–92 and low in the following year. Population density increased greatly in spring 1992 but crashed in spring and summer 1993, when a high mortality of wild boar occurred due to starvation.
The decrease in home-range size during the food shortage was explained as a possible strategy adopted by wild boar to cope with starvation. The difference in home-range size and amount of activity found between the first and the second year of study were probably influenced by differences in population density, although the individual effects of food availability and population density were not clearly separable.     

Adaptive thermoregulation in golden spiny mice: the influence of season and food availability on body temperature

We studied the effect of food supplementation during summer and winter in seminatural field conditions on thermoregulation of a desert rodent, the golden spiny mouse Acomys russatus. We hypothesized that (a) under natural food availability (control conditions), mice will use less precise thermoregulation (i.e., an increase in the variance of body temperature [T(b)]) during winter because of low ambient temperatures (T(a)'s) and low food availability and during summer because of low food and water availability; (b) food supplementation will result in more precise thermoregulation during winter, but the effect will be smaller during summer because variation in T(b) in summer is also driven by water availability during that period. We found that under natural food availability, spiny mice thermoregulated more precisely during summer than during winter. They spent more time torpid during summer than during winter even when food was supplemented (although summer nights are shorter), allowing them to conserve water. Supplementing food resulted in more precise thermoregulation in both seasons, and mice spent less time torpid. In summer, thermoregulation at high T(a)'s was less precise, resulting in higher maximum T(b)'s in summer than in winter and when food was supplemented, in accord with the expected effect of water shortage on thermoregulation. Our results suggest that as expected, precise thermoregulation is beneficial when possible and is abandoned only when the costs of homeothermy outweigh the benefits.     

Influence of stock density and rate and temporal patterns of forage allocation on the diet mixing behavior of sheep grazing sagebrush steppe

An animal's ability to select a diet depends on forage availability, the density of herbivores, and the allocation of forage resources. We hypothesized that the temporal and spatial availability of forages in sagebrush steppe vegetation both influence food selection and may encourage lambs to learn to use sagebrush, a shrub that contains high levels of terpenes that limit food intake. We conducted a field study with three treatments of 20 lambs each. Forage resources were manipulated by restricting grazing areas. Lambs in treatment H grazed at a high stock density, whereas lambs in treatment L grazed at a low stock density; both groups were moved to fresh pasture daily. Lambs in treatment H3 had three times the area of treatment H, but they were moved every 3 days, making the total area grazed by H and H3 equal, but with a different temporal allocation of forage. Scan sampling was used to record the incidence of consumption among sagebrush and understory herbs. During 24 days of conditioning, stock density and temporal patterns of forage allocation both influenced the behavior of lambs. Lambs in H spent more time foraging on sagebrush (25%) than lambs in H3 (16%), and lambs in both H and H3 spent more time foraging on sagebrush than lambs in L (1%) (P < 0.05). For lambs in H3, foraging on sagebrush was cyclic and depended on the daily availability of herbs. Use of sagebrush, expressed as a percentage of total foraging time, was <1% for day 1, 13% for day 2, and 37% for day 3 (P < 0.0001). Importantly, use of sagebrush by H3 and H increased steadily as the trial progressed, whereas use of sagebrush by L was consistently low (P < 0.05). Following conditioning, when lambs were tested at low stock densities during preference test 1, use of sagebrush did not differ by groups, but at higher stock densities in preference test 2, their use of sagebrush varied by treatments: H > H3 > L (P < 0.005). Thus, the availability of alternative foods, manipulated through animal density and the temporal allocation of those resources, both affected how readily lambs learned to use sagebrush.     

Determinants of territory size in the pomacentrid reef fish,Parma victoriae

Summary Factors governing the size of territories defended by the pomacentrid reef fish, Parma victoriae, were investigated, prompted by contradictory predictions in the literature concerning the effects of food supply and competitors. Observations were carried out over the non-breeding period (March–October) on a medium density population in which territories were partially contiguous. The territory size of adult fish varied between 3 and 26 m², and was inversely correlated with local densities of conspecifics. The same range in territory size was found for both males and females, which did not differ in the time they spent on territory defence and foraging activities. No correlation existed between territory size and the abundance of algal food, body size, age or time spent on territory defence. Also, there was little variation in territory size over time, despite seasonal changes in the abundance of food algae.Experimental reduction of food supplies on isolated territories of males and females had no effect on territory size. In a higher density habitat an experiment was carried out in which population density and food abundance were simultaneously manipulated. This showed that territory size was primarily determined by intraspecific interactions, as territories exhibited considerable increases in size upon removal of neighbours. No changes in the size of defended areas resulted from either artificial increases or decreases of food levels. There were also no changes in the time spent on defence of territories, foraging time or feeding rates associated with food manipulations or territory expansion, which suggested that food was not a limited resource. This conflicted with current theories proposed to explain territory defence and expansion. It is hypothesized that intraspecific interactions constrain territory size well below the optimum in terms of the abundance of preferred food algal species.     

Ancestral ecological regime shapes reaction to food limitation in the Least Killifish,Heterandria formosa

Populations with different densities often show genetically based differences in life histories. The divergent life histories could be driven by several agents of selection, one of which is variation in per‐capita food levels. Its relationship with population density is complex, as it depends on overall food availability, individual metabolic demand, and food‐independent factors potentially affecting density, such as predation intensity. Here, we present a case study of two populations of a small live‐bearing freshwater fish, one characterized by high density, low predation risk, low overall food availability, and presumably low per‐capita food levels, and the other by low density, high predation risk, high overall food availability, and presumably high per‐capita food levels. Using a laboratory experiment, we examined whether fish from these populations respond differently to food limitation, and whether size at birth, a key trait with respect to density variation in this species, is associated with any such differential responses. While at the lower food level growth was slower, body size smaller, maturation delayed, and survival reduced in both populations, these fitness costs were smaller in fish from the high‐density population. At low food, only 15% of high‐density fish died, compared to 75% of low‐density fish. This difference was much smaller at high food (0% vs. 15% mortality). The increased survival of high‐density fish may, at least partly, be due to their larger size at birth. Moreover, being larger at birth enabled fish to mature relatively early even at the lower food level. We demonstrate that sensitivities to food limitation differ between study populations, consistent with selection for a greater ability to tolerate low per‐capita food availability in the high‐density population. While we cannot preclude other agents of selection from operating in these populations simultaneously, our results suggest that variation in per‐capita food levels is one of those agents.     

Population dynamics and morphometries of Gammarus pulex L.: evidence of seasonal food limitation in a freshwater detritivore

SUMMARY. 1. Seasonal changes in population size structure of Gammarus pulex L. in a Cotswold stream appeared to indicate a growth check in late summer.
2. The relationships between dry weight and body length, and between the number of primary flagellar segments on the first antenna and either dry weight or body length provided further evidence of a reduction in growth in mid and late summer.
3. Body fat content was minimal (4% of dry weight) at the end of summer, when large particulate organic detritus was scarce or of poor quality, and maximal in late winter (17.9% dry weight in females; 9.4% in males), after a period of high food availability. In a field experiment, the fat content of animals in summer was raised to levels typical of winter by providing high quality food.
4. Field and experimental evidence together strongly infer that this population of G. pulex was subject to severe food limitation from early summer until leaf fall in autumn.     

Demography of alpine red squirrel populations in relation to fluctuations in seed crop size

Vertebrate population dynamics, social organisation and space use often are closely associated with the distribution of critical resources, such as food. Tree squirrels are ideal models to study these relationships, since both key demographic parameters (reproduction, survival and dispersal) and spatio-temporal variation in food supplies (measured as seed-crop size) can be reliably estimated. In this paper we test the following two predictions underlying the association between annual food abundance and demography in six alpine red squirrel populations, both with and without time-lag effects: 1) between-season and between-year fluctuations in survival rate, population density and increase parallel those in food availability; and 2) individuals follow a resource tracking strategy and increase in density mainly the year after a rich seed-crop. Red squirrels occurred at higher densities in Scots pine forest, characterised by stable seed-crops, than in Norway spruce with more abundant but more variable seed crops. Fluctuations in numbers were positively correlated with food availability, measured as annual conifer seed-crop sizes. Overall, adult survival rates were higher than those of subadults, and survival substantially fluctuated between seasons and years. Autumn densities and rates of population increase (summer-autumn) were strongly correlated with the same year's autumn seed-crop, while correlations with the previous year's seed-crop (time-lag models) were either weak (population density) or absent (population increase). Results of this paper show that fluctuations in red squirrel densities in habitats with strong temporal variation in seed production are more closely linked with food availability than in more stable habitats. In addition, in the Alpine conifer forests squirrel population sizes, in autumn, increase in synchrony with food resources, eliminating the population lag normally present when resources are produced in pulses.     

Density-dependent parameters and demographic equilibrium in open populations

Populations near their equilibrium are expected to show density-dependence through a negative feedback on at least one demographic parameter, e.g. survival and/or productivity. Nevertheless, it is not always clear which vital rate is affected the most, and even less whether this dependence holds in open populations in which immigration and emigration are also important. We assessed the relative importance of population density in the variation of local survival, recruitment, proportion of transients (emigrants) and productivity through the analysis of detailed life-histories of 4286  seabirds from a colony that reached an apparent demographic equilibrium after a period of exponential increase. We provide evidence that the role of population density and resource availability changes according to the demographic parameter considered. Estimates indicated that transients increased from 5% to 20% over the study period, suggesting an average turnover of about 1400 individuals per year. The parameters most influenced by population density alone were local survival and probability of transience. Recruitment was negatively associated with population density during the increasing phase but unexpected high values were also recorded at high population levels. These high values were explained by a combination of population size and food availability. Mean productivity varied with food availability, independently from population variations. The population density alone explained up to a third of the yearly variation of the vital rates considered, suggesting that open populations are equally influenced by stochastic and density-independent events (such as environmental perturbations) than by intrinsic (i.e. density-dependent) factors.     

Changes in body condition and body size affect breeding and recruitment in fluctuating house mouse populations in south‐eastern Australia

Changes in body condition and body size in field populations of house mice, Mus domesticus, were examined to investigate why mouse populations do not increase rapidly in some years when favourable environmental and demographic conditions indicate they might. Mice had repeated seasonal patterns each year in breeding, growth rates and body condition that reflected the seasonal availability of food, but mean levels for each parameter varied among years. In most years mice lost body condition during summer, breeding declined and population growth slowed. Rapid population growth occurred when body condition was generally high and was maintained throughout summer. Female mice with large body length were more likely to breed than smaller mice, at all times, but changes in body condition accounted for most of the variability in female breeding activity between years and between habitats, and for the seasonal changes in the importance of body length. During rapid population growth, the recruitment rate of juveniles relative to the number of breeding females was 150–300% higher than in other years but adult survival rates were not higher. The data indicate that the ability of mice to maintain body condition, particularly when subject to moisture stress in summer, affects the proportion of females breeding, the number of juveniles weaned and their body condition at weaning, and is promoted by foraging conditions that favour maintenance of juvenile body condition after weaning. These factors, in turn, greatly affect juvenile recruitment rates and eventual population density of mice. Low juvenile survival is suggested as a reason that numbers of house mice in southern Australian cereal‐growing areas do not increase rapidly in some years when other parameters are favourable. Similar processes are likely to play a role in regulating other rodent populations.     

Effects of a Whole-lake,Experimental Fertilization on Lake Trout in a Small Oligotrophic Arctic Lake

We tested whether increased phosphorus and nitrogen concentrations would affect a lake trout (Salvelinus namaycush) population in a small oligotrophic lake with a benthically dominated food web. From 1990 to 1994, nitrogen and phosphorus were added to Lake N1 (4.4 ha) at the arctic Long-Term Ecological Research site in Alaska. We used mark/recapture methods to determine the lake trout population size, size structure, recruitment, and individual growth from 1987 to 1999. Data were also collected on water chemistry and food availability. Fertilization resulted in increased pelagic primary productivity, chlorophyll a, turbidity, snail density, and hypoxia in summer and winter. Lake trout density was not affected by the manipulation however growth and average size increased. Recruitment was high initially, but declined throughout the fertilization. These results suggest that lake trout were affected through increased food availability and changes to the physical characteristics of the lake. During fertilization, hypoxia near the sediments may have killed over-wintering embryos and decreased habitat availability. Although lake trout responded strongly to increased nutrients, loss of recruitment might jeopardize lake trout persistence if arctic lakes undergo eutrophication.     

The Role of Individual Traits and Environmental Factors for Diet Composition of Sheep

Large herbivore consumption of forage is known to affect vegetation composition and thereby ecosystem functions. It is thus important to understand how diet composition arises as a mixture of individual variation in preferences and environmental drivers of availability, but few studies have quantified both. Based on 10 years of data on diet composition by aid of microhistological analysis for sheep kept at high and low population density, we analysed how both individual traits (sex, age, body mass, litter size) linked to preference and environmental variation (density, climate proxies) linked to forage availability affected proportional intake of herbs (high quality/low availability) and Avenella flexuosa (lower quality/high availability). Environmental factors affecting current forage availability such as population density and seasonal and annual variation in diet had the most marked impact on diet composition. Previous environment of sheep (switch between high and low population density) had no impact on diet, suggesting a comparably minor role of learning for density dependent diet selection. For individual traits, only the difference between lambs and ewes affected proportion of A. flexuosa, while body mass better predicted proportion of herbs in diet. Neither sex, body mass, litter size, ewe age nor mass of ewe affected diet composition of lambs, and there was no effect of age, body mass or litter size on diet composition of ewes. Our study highlights that diet composition arises from a combination of preferences being predicted by lamb and ewes’ age and/or body mass differences, and the immediate environment in terms of population density and proxies for vegetation development.     

Test of the ecological-constraints model on ursine colobus monkeys (Colobus vellerosus) in Ghana

For group-living mammals, the ecological-constraints model predicts that within-group feeding competition will increase as group size increases, necessitating more daily travel to find food and thereby constraining group size. It provides a useful tool for detecting scramble competition any time it is difficult to determine whether or not food is limiting. We tested the ecological-constraints model on highly folivorous ursine colobus monkeys (Colobus vellerosus) at the Boabeng-Fiema Monkey Sanctuary in Ghana. Three differently sized groups were followed for 13 months and two others were followed for 6 months each in 2004-2005 using focal-animal sampling and ranging scans; ecological plots and phenology surveys were used to determine home-range quality and food availability. There was relatively little difference in home-range quality, monthly food availability, diet, adult female ingestion rates, and rate of travel within food patches between the groups. However, home-range size, day-range length, and percent of time spent feeding all increased with group size. We performed a single large test of the ecological-constraints model by combining several separate Spearman correlations, each testing different predictions under the model, using Fisher's log-likelihood method. It showed that the ecological-constraints model was supported in this study; scramble competition in this population is manifesting in increased ranging and time spent feeding. How costly this increased energy expenditure is for individuals in larger groups remains to be determined.     

Food limitation of population density in the orb-wed spider,Nephila clavata

Field studies were conducted to clarify whether variation in food availability among habitats influences population density, and whether population density has a negative effect on foraging success in the orb-web spider, Nephila clavata. Lifetime food consumption per individual (i.e., foraging success) strongly correlated with mean body size of adult females and mean fecundity in populations. Also, there was a positive correlation between foraging success and population density. Since foraging success reflected potential prey availability in the habitat, food resource appeared to be a limiting factor for populations in this spider. Mean fecundity per individual correlated with population density of the following year, suggesting that decreased reproduction is a major component of food limitation on population density. Consistent defferences in mean body size between particular sites were observed over years, while such difference was less obvious in density. Thus, ranking of food abundance among habitats seems to be predictable between years. A field experiment revealed that an artificial increase in population density had no negative effect on the feeding rate of individuals, suggesting that intraspecific competition for food is not important in this species.     

Effects of food abundance on genetic and maternal variation in the growth rate of juvenile red squirrels

Sources of variation in growth in body mass were assessed in natural and experimental conditions of high and low food abundance using reciprocal cross-fostering techniques and long-term data (1987-2002) for a population of North American red squirrels (Tamiasciurus hudsonicus). Growth rates were significantly higher in naturally good and food supplemented conditions, than in poor conditions. Mother-offspring resemblance was higher in poor conditions as a result of large increases in both the direct genetic variance and direct-maternal genetic covariance and a smaller increase in the coefficient of maternal variation. Furthermore, the genetic correlation across environments was significantly less than one indicating that sources of heritable variation differed between the two environments. These results are consistent with the hypothesis that selection has eroded heritable variation for growth more in good conditions and indicate the potential for independent adaptation of growth rates in good and poor conditions.     

The effects of food abundance on breeding performance and adult time budgets of Guillemots Uria aalge

The breeding performance, food fed to chicks and adult time budgets of Guillemots Uria aalge were examined in a year of high and a year of low food availabiIity. There was no difference between the 2 years in reproductive success, although the rate of chick feeding, chick weight and fledging success were greater in the year of high food availability. On average, chick prey items were larger in the poor food year, but this was insufficient to compensate for the lower feeding frequency. Chick feeding frequency did not differ between days in the good year but did increase later in the season in the poor food year. Compared with the high food availability year, adult Guillemots in the year of low food availability spent much less time resting at the breeding colony. and their foraging trips were twice as long. Foraging birds tended to make several successive trips before resuming brooding duties from their mates when food supplies were good, but in the low food availability year single trips were the norm. These results demonstrate that predators experiencing reduced food supply may mitigate the effects on their reproductive output by shifting their time allocation such that more time is available for foraging.     

Plastic reproductive allocation as a buffer against environmental stochasticity – linking life history and population dynamics to climate

Empirical work suggest that long‐lived organisms have adopted risk sensitive reproductive strategies where individuals trade the amount of resources spent on reproduction versus survival according to expected future environmental conditions. Earlier studies also suggest that climate affects population dynamics both directly by affecting population vital rates and indirectly through long‐term changes in individual life histories. Using a seasonal and state‐dependent individual‐based model we investigated how environmental variability affects the selection of reproductive strategies and their effect on population dynamics. We found that: (1) dynamic, i.e. plastic, reproductive strategies were optimal in a variable climate. (2) Females in poor and unpredictable climatic regimes allocated fewer available resources in reproduction and more in own somatic growth. This resulted in populations with low population densities, and a high average female age and body mass. (3) Strong negative density dependence on offspring body mass and survival, along with co‐variation between climatic severity and population density, resulted in no clear negative climatic effects on reproductive success and offspring body mass. (4) Time series analyses of population growth rates revealed that populations inhabiting benign environments showed the clearest response to climatic perturbations as high population density prohibited an effective buffering of adverse climatic effects as individuals were not able to gain sufficient body reserves during summer. Regularly occurring harsh winters ‘harvested’ populations, resulting in persistent low densities, and released them from negative density dependent effects, resulting in high rewards for a given resource allocation.     

Density dependent and temporal variability in habitat selection by a large herbivore; an experimental approach

Both density dependent and density independent processes such as climate affect population dynamics in large herbivores. Understanding herbivore foraging patterns is essential to identify the underlying mechanisms behind variation in vital rates. However, very little is known about how animals vary their selection of habitat temporally, alone or in interaction with density during summer. At the foraging scale, we tested using a fully replicated experiment whether domestic sheep Ovis aries stocked at high (80  per  km²) and low (25  per  km²) densities (spatial contrasts) varied their habitat selection temporally over a four year period. We predicted reduced selection of high productivity vegetation types with increasing density, and that seasonal and annual variation in climate would affect this density dependent selection pattern by increasing competition for high quality habitats in late grazing season and in years with poor vegetation development and over time related to vegetation responses to grazing. As predicted from the Ideal free distribution model, selection of high productivity habitat decreased at high density. There was also a marked temporal variation in habitat selection. Selection of the most productive vegetation types declined towards the end of each grazing season, but increased over years both at low and high sheep density. There was only weak evidence for interactions, as selection ratio of highly productive habitats tended to increase more over years at low density as compared to high density. Limited interactive effects of density and annual variation on habitat selection during summer may explain why similar interactions in vital rates have rarely been reported for summer seasons. Our results are consistent with the view that variation in habitat selection is a central mechanism for climate and density related variation in vital rates.