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.     

Balancing food and density‐dependence in the spatial distribution of an interference‐prone forager

Foraging distributions are thought to be density‐dependent, because animals not only select for a high availability and quality of resources, but also avoid conspecific interference. Since these processes are confounded, their relative importance in shaping foraging distributions remains poorly understood. Here we aimed to rank the contribution of density‐dependent and density‐independent effects on the spatio‐temporal foraging patterns of eurasian oystercatchers. In our intertidal study area, tides caused continuous variation in oystercatcher density, providing an opportunity to disentangle conspecific interference and density‐independent interactions with the food landscape. Spatial distributions were quantified using high‐resolution individual tracking of foraging activity and location. In a model environment that included a realistic reconstruction of both the tides and the benthic food, we tested a family of behaviour‐based optimality models against these tracking data. Density‐independent interactions affected spatial distributions much more strongly than conspecific interference, even in an interference‐prone species like oystercatchers. Spatial distributions were governed by avoidance of bill injury costs, selection for high interference‐free intake rates and a decreasing availability of benthic bivalve prey after their exposure. These density‐independent interactions outweighed interference competition in terms of effect size. We suggest that the bottleneck in our mechanistic understanding of foraging distributions may be primarily the role of density‐independent prey attributes unrelated to intake rates, like damage costs in the case of oystercatchers foraging on perilous prey. At a landscape scale, above the finest inter‐individual distances, effects of conspecific interaction on spatial distributions may have been overemphasised.     

The Effect of Male Incubation Feeding,Food and Temperature on the Incubation Behaviour of New Zealand Robins

Because of finite resources, organisms face conflict between their own self‐care and reproduction. This conflict is especially apparent in avian species with female‐only incubation, where females face a trade‐off between time allocated to their own self‐maintenance and the thermal requirements of developing embryos. We recorded incubation behaviour of the New Zealand robin (Petroica longipes), a species with female‐only incubation, male incubation feeding and high nest predation rates. We examined how male incubation feeding, ambient temperature and food availability (invertebrate biomass) affected the different components of females’ incubation behaviour and whether incubation behaviour explained variation in nest survival. Our results suggest that male incubation feeding rates of 2.8 per hour affect the female’s incubation rhythm by reducing both on‐ and off‐bout duration, resulting in no effect on female nest attentiveness, thus no support for the female‐nutritional hypothesis. The incubation behaviours that we measured did not explain nest survival, despite high nest predation rates. Increased ambient temperature caused an increase in off‐bout duration, whereas increased food availability increased on‐bout duration. While males play a vital role in influencing incubation behaviour, female robins attempt to resolve the trade‐off between their own foraging needs and the thermal requirements of their developing embryos via alternating their incubation rhythm in relation to both food and temperature.     

Time and travelling costs during chick‐rearing in relation to habitat quality in Little Owls Athene noctua

In many bird species, parents adjust their home‐ranges during chick‐rearing to the availability and distribution of food resources, balancing the benefits of energy intake against the costs of travelling. Over recent decades, European agricultural landscapes have changed radically, resulting in the degradation of habitats and reductions in food resources for farmland birds. Lower foraging success and longer foraging trip distances that result from these changes are often assumed to reduce the reproductive performance of parents, although the mechanisms are not well understood. We tested the behavioural response of chick‐rearing Little Owls Athene noctua to variation in habitat diversity in an agricultural landscape. We equipped females with GPS loggers and received adequate range‐use data for 19 individuals (6063–14 439 locations per bird). In habitats dominated by homogeneous cropland habitats, home‐ranges were over 12 ha in size, whereas in highly diverse habitats they were below 2 ha. Large home‐ranges were associated with increased flight activity (117% of that of birds in small home‐ranges) and distances travelled per night (152%), increased duration of foraging trips (169%) covering larger distances (246%), and reduced nest visiting rates (81%). The study therefore provides strong correlative evidence that Little Owls breeding in monotonous farmland habitats expend more time and energy for a lower benefit in terms of feeding rates than do birds in more heterogeneous landscapes. As nestling food supply is the main determinant of chick survival, these results suggest a strong impact of farmland characteristics on local demographic rates. We suggest that preserving and creating islands of high habitat diversity within uniform open agricultural landscapes should be a key target in the conservation of Little Owl populations.     

Molecular and spatial analyses reveal links between colony‐specific foraging distance and landscape‐level resource availability in two bumblebee species

Foraging distance is a key determinant of colony survival and pollination potential in bumblebees Bombus spp. However this aspect of bumblebee ecology is poorly understood because of the difficulty in locating colonies of these central place foragers. Here, we used a combination of molecular microsatellite analyses, remote sensing and spatial analyses using kernel density estimates to estimate nest location and foraging distances for a large number of wild colonies of two species, and related these to the distribution of foraging habitats across an experimentally manipulated landscape. Mean foraging distances were 755 m for Bombus lapidarius and 775 m for B. pascuorum (using our most conservative estimation method). Colony‐specific foraging distances of both species varied with landscape structure, decreasing as the proportion of foraging habitats increased. This is the first time that foraging distance in wild bumblebees has been shown to vary with resource availability. Our method offers a means of estimating foraging distances in social insects, and informs the scale of management required to conserve bumblebee populations and enhance their pollination services across different landscapes.     

Nest and foraging‐site selection in Yellowhammers Emberiza citrinella: implications for chick provisioning

Capsule Vegetation structure and invertebrate abundance interact to influence both foraging sites and nestling provisioning rate; when invertebrate availability is low, adults may take greater risks to provide food for their young.

Aims To investigate nesting and foraging ecology in a declining farmland bird whose fledging success is influenced by the availability of invertebrate prey suitable for feeding to offspring, and where perceived predation risk during foraging can be mediated by vegetation structure.

Methods Provisioning rates of adult Yellowhammers feeding nestlings were measured at nests on arable farmland. Foraging sites were compared with control sites of both the same and different microhabitats; provisioning rate was related to habitat features of foraging‐sites.

Results Foraging sites had low vegetation density, probably enhancing detection of predators, or high invertebrate abundance at high vegetation density. Parental provisioning rate decreased with increasing vegetation cover at foraging sites with high invertebrate abundance; conversely, where invertebrate abundance was low, provisioning rate increased with increasing vegetation cover.

Conclusions Vegetation structure at foraging sites suggests that a trade‐off between predator detection and prey availability influences foraging site selection in Yellowhammers. Associations between parental provisioning rate and vegetation variables suggest that where invertebrate abundance is high birds increase time spent scanning for predators at higher vegetation densities; however, when prey are scarce, adults may take more risks to provide food for their young.     

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.     

Low larval densities in northern populations reinforce range expansion by a Mediterranean damselfly

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Environmental variation and behaviour: resource availability during ontogeny and feeding performance in salamander larvae (Ambystoma texanum)

1. The effects of resource availability during ontogeny on subsequent feeding performance were investigated in larvae of the small-mouthed salamander ( Ambystoma texanum ).
2. Salamander larvae were reared individually in either high or low prey density treatments for 7 weeks prior to intermediate prey density foraging trials. Larvae from the low prey density treatment were on average 35% smaller in body size than individuals from the high prey density treatment.
3. Resource availability during development influenced larval feeding rates and altered the relationship between body size and three feeding performance measures (attack rates, capture success and feeding rates). Feeding rates in predation trials were also positively correlated with growth rate early in the larval period (until the end of week 5).
4. These results suggest that the environment to which developing organisms are exposed can have significant effects on subsequent behaviour, and that small-mouthed salamander larvae may show state-dependent changes in feeding behaviour in response to differences in long-term feeding history. Additionally, differences in feeding performance may influence the probability of survival to the adult stage for organisms that utilize ephemeral habitats.     

Time budget of the green monkey,Cercopithecus sabaeus: Some optimal strategies

Members of a single group of green monkeys spent, on average, 44.8% of their waking time foraging, 46.7% resting, and 8.5% in social activities, over 1 year. There was significant variation in activity budgets over months (ranging from 35 to 55% of the time spent feeding). Diurnal rhythms of feeding and ranging were influenced by the daily cycle of temperature in predictable ways in different seasons: in the dry season, activity was reduced if it was too hot or too cold, while the temperature in the wet season did not affect activities. Feeding was also synchronized among individuals on a finer time scale, irrespective of the time of day. There was closer synchrony when feeding on less common foods. An ecological model of foraging time and energetics was tested, using estimates of the costs and benefits of foraging and predicting how these are optimally balanced in relation to the food density. Both feeding time and distance traveled increased as food availability increased. Costs and benefits were balanced over several days. Comparisons between populations of Cercopithecus aethiopswere made; differences in time budgets were compared with differences in the availability and quality of food. Insufficient comparative data are available for firm conclusions about the role of different energetic and nutritive strategies in population differences.     

Fungal chemical defence alters density‐dependent foraging behaviour and success in a fungivorous soil arthropod

1. Aggregative behaviour in fungivorous soil arthropods is widespread; its adaptive value, however, is largely unknown. In this study, the spatial foraging behaviour of a collembolan, Folsomia candida, and the fitness consequences of feeding at different densities on the filamentous fungus Aspergillus nidulans were investigated. The effect of two fungal strains were compared; a wild‐type (wt) and a transgenic strain that lacks the ability to express the global secondary metabolite regulator LaeA (ΔlaeA). 2. In laboratory foraging tests, F. candida exhibited aggregated distributions of individuals across four distinct fungal colonies that were arranged in short distances from each other. By quantifying the extent of the feeding damage at each single colony, a more evenly distributed feeding activity was found among wt colonies than among chemical‐deficient colonies. 3. In a fitness experiment, where collembolans at different densities were restricted to feed on single A. nidulans colonies, mean growth rate of F. candida was positively related to density on the wt A. nidulans strain, but negatively related to density on the chemical‐deficient strain. 4. Depending on the fungus' ability to express secondary chemicals and availability of fungal food sources, F. candida may employ different foraging strategies: (i) avoidance of prolonged feeding on single colonies in a rich habitat (travel costs low), and (ii) intensified group feeding on single colonies in a resource‐limited habitat (travel costs high). It was hypothesised that flexibility in fungivore foraging behaviour (clumping vs. spreading feeding activity) is adaptive because it allows avoidance/overcoming induced fungal chemical defence.     

Ecological correlates of space use patterns in wild western lowland gorillas

The distribution of resources is a crucial determinant of animals' space use (e.g., daily travel distance, monthly home range size, and revisitation patterns). We examined how variation in ecological parameters affected variability in space use patterns of western lowland gorillas, Gorilla gorilla gorilla. They are an interesting species for investigating this topic because key components of their diet are nonfruit items (herbaceous vegetation and tree leaves) that occur at low density and are sparsely distributed, and fruits, which show high spatiotemporal variation in availability. We estimated how availability of nonfruit foods and fruit, frugivory (proportion of feeding time consuming fruit), and swamps in areas used by the gorillas influenced daily travel distance, monthly home range size, and revisit frequency to grid cells in the home range of one habituated gorilla group in Loango National Park, Gabon. Using location data from 2015 to 2018, we found that the gorillas decreased their daily travel distance as both the density of nonfruit foods and the proportion of swamps in areas used increased. Daily travel distances were shorter when both frugivory and availability of fruit were higher, yet, daily travel distances were longer when availability of fruit was low but frugivory was still high. Furthermore, monthly home range size increased as frugivory increased and monthly revisit frequencies to an area increased as fruit availability of an area increased. In conclusion, the availability of both nonfruit and fruit influenced the gorillas' space use patterns. Gorillas decreased foraging effort when food availability was high but were willing to incur increasing foraging costs to feed on fruit when availability was low. This study highlights how animals have to adjust their space use with changing resource availability and it emphasizes the value of examining multiple parameters of space use.     

Environmental variation and behaviour: resource availability during ontogeny and feeding performance in salamander larvae (Ambystoma texanum)

1. The effects of resource availability during ontogeny on subsequent feeding performance were investigated in larvae of the small-mouthed salamander ( Ambystoma texanum ).
2. Salamander larvae were reared individually in either high or low prey density treatments for 7 weeks prior to intermediate prey density foraging trials. Larvae from the low prey density treatment were on average 35% smaller in body size than individuals from the high prey density treatment.
3. Resource availability during development influenced larval feeding rates and altered the relationship between body size and three feeding performance measures (attack rates, capture success and feeding rates). Feeding rates in predation trials were also positively correlated with growth rate early in the larval period (until the end of week 5).
4. These results suggest that the environment to which developing organisms are exposed can have significant effects on subsequent behaviour, and that small-mouthed salamander larvae may show state-dependent changes in feeding behaviour in response to differences in long-term feeding history. Additionally, differences in feeding performance may influence the probability of survival to the adult stage for organisms that utilize ephemeral habitats.     

Taking a trip to the shelf: Behavioral decisions are mediated by the proximity to foraging habitats in the black‐legged kittiwake

For marine top predators like seabirds, the oceans represent a multitude of habitats regarding oceanographic conditions and food availability. Worldwide, these marine habitats are being altered by changes in climate and increased anthropogenic impact. This is causing a growing concern on how seabird populations might adapt to these changes. Understanding how seabird populations respond to fluctuating environmental conditions and to what extent behavioral flexibility can buffer variations in food availability can help predict how seabirds may cope with changes in the marine environment. Such knowledge is important to implement proper long‐term conservation measures intended to protect marine predators. We explored behavioral flexibility in choice of foraging habitat of chick‐rearing black‐legged kittiwakes Rissa tridactyla during multiple years. By comparing foraging behavior of individuals from two colonies with large differences in oceanographic conditions and distances to predictable feeding areas at the Norwegian shelf break, we investigated how foraging decisions are related to intrinsic and extrinsic factors. We found that proximity to the shelf break determined which factors drove the decision to forage there. At the colony near the shelf break, time of departure from the colony and wind speed were most important in driving the choice of habitat. At the colony farther from the shelf break, the decision to forage there was driven by adult body condition. Birds furthermore adjusted foraging behavior metrics according to time of the day, weather conditions, body condition, and the age of the chicks. The study shows that kittiwakes have high degree of flexibility in their behavioral response to a variable marine environment, which might help them buffer changes in prey distribution around the colonies. The flexibility is, however, dependent on the availability of foraging habitats near the colony.     

Density‐dependent,central‐place foraging in a grazing herbivore: competition and tradeoffs in time allocation near water

Optimal foraging theory addresses one of the core challenges of ecology: predicting the distribution and abundance of species. Tests of hypotheses of optimal foraging, however, often focus on a single conceptual model rather than drawing upon the collective body of theory, precluding generalization. Here we demonstrate links between two established theoretical frameworks predicting animal movements and resource use: central‐place foraging and density‐dependent habitat selection. Our goal is to better understand how the nature of critical, centrally placed resources like water (or minerals, breathing holes, breeding sites, etc.) might govern selection for food (energy) resources obtained elsewhere – a common situation for animals living in natural conditions. We empirically test our predictions using movement data from a large herbivore distributed along a gradient of water availability (feral horses, Sable Island, Canada, 2008–2013). Horses occupying western Sable Island obtain freshwater at ponds while in the east horses must drink at self‐excavated wells (holes). We studied the implications of differential access to water (time needed for a horse to obtain water) on selection for vegetation associations. Consistent with predictions of density‐dependent habitat selection, horses were reduced to using poorer‐quality habitat (heathland) more than expected close to water (where densities were relatively high), but were free to select for higher‐quality grasslands farther from water. Importantly, central‐place foraging was clearly influenced by the type of water‐source used (ponds vs. holes, the latter with greater time constraints on access). Horses with more freedom to travel (those using ponds) selected for grasslands at greater distances and continued to select grasslands at higher densities, whereas horses using water holes showed very strong density‐dependence in how habitat could be selected. Knowledge of more than one theoretical framework may be required to explain observed variation in foraging behavior of animals where multiple constraints simultaneously influence resource selection.     

Foraging in precocial chicks of the black-tailed godwit Limosa limosa: vulnerability to weather and prey size

Self-feeding precocial development is associated with high energy requirements and potentially vulnerable to short-term reductions in food availability, yet few studies have investigated development of foraging in precocial chicks and its sensitivity to environmental conditions. We studied time budgets and foraging behaviour during the 25-d prefledging period in the insectivorous chicks of a grassland shorebird, the black-tailed godwit Limosa limosa . Until 8–10  d old, parental brooding was the main determinant of chicks' daily foraging time. Brooding decreased with age and temperature and increased during rainfall. Foraging time increased to 70–90% of the daylight period in chicks older than a week, during which distances of 3–12  km  d⁻¹ were covered. Chicks took 98% of their arthropod prey from the grassland vegetation. Prey ingestion rates increased in the first week and slowly declined thereafter, modified by wind speed, temperature and time of day. Chicks in poor body condition were brooded more than chicks growing normally and hence had less feeding time, potentially leading to a negative condition spiral under adverse conditions. However, we found no effect of condition on prey ingestion rate that would preclude recovery when conditions improve. Combining behavioural observations with data on energy expenditure revealed that mean prey size was small (1–4.5  mg), necessitating a high feeding rate, but increased notably after 7–10  d of age. This coincided with a decrease in walking speed, suggesting that chicks fed more selectively. Prey of older chicks approached the upper limit of sizes available in exploitable densities in the grassland vegetation, and this enhances the chicks' sensitivity to variation in prey availability due to weather and agricultural practice.     

Sex-Related Differences in the Trade-Off between Foraging and Vigilance in a Granivorous Forager

The relationship between intake rate and food density can provide the foundation for models that predict the spatiotemporal distribution of organisms across a range of resource densities. The functional response, describing the relationship between resource density and intake rate is often interpreted mechanistically as the relationships between times spend searching and handling. While several functional response models incorporate anti-predator vigilance (defined here as an interruption of feeding or some other activity to visually scan the environment, directed mainly towards detecting potential predators), the impacts of environmental factors influencing directly anti-predator vigilance remains unclear. We examined the combined effects of different scenarios of predation risk and food density on time allocation between foraging and anti-predator vigilance in a granivorous species. We experimentally exposed Skylarks to various cover heights and seed densities, and measured individual time budget and pecking and intake rates. Our results indicated that time devoted to different activities varied as a function of both seed density and cover height. Foraging time increased with seed density for all cover heights. Conversely, an increased cover height resulted in a decreased foraging time. Contrary to males, the decreased proportion of time spent foraging did not translate into a foraging disadvantage for females. When vegetation height was higher, females maintained similar pecking and intake rates compared to intermediate levels, while males consistently decreased their energy gain. This difference in anti-predator responses suggests a sexually mediated strategy in the food-safety trade-off: when resource density is high a females would adopt a camouflage strategy while an escape strategy would be adopted by males. In other words, males would leave risky-areas, whereas females would stay when resource density is high. Our results suggest that increased predation risk might generate sexually mediated behavioural responses that functional response models should perhaps better consider in the future.     

Responses by Central‐Place Foragers to Manipulations of Brood Size: Parent Flickers Respond to Proximate Cues but do not Increase Work Rate

Manipulations of brood size measure the willingness or ability of parents to invest in offspring and different reproductive roles may lead to differences in feeding effort between the sexes. Parental investment in birds is usually assessed by quantifying feeding rates, but this provides an incomplete picture of parental effort because it fails to account for how parents collect food on the landscape. We studied northern flickers (Colaptes auratus), a woodpecker in which males provide the majority of parental care and used a repeated measures design and short‐term (24 h) brood enlargements (N = 35) and reductions (N = 27) to assess effects of treatment on feeding rates to nestlings and parental foraging behaviour. Parents of enlarged broods did not significantly increase feeding rate, resulting in a decline in nestling mass. Parents of reduced broods decreased their feeding rates by 84%, but increased per capita feeding rates, resulting in nestling mass gain. The variation in feeding rates to enlarged broods was not influenced by feather corticosterone, body condition, feather re‐growth rate or mass change between the incubation and nestling periods. Foraging pattern on the landscape remained the same during the enlarged treatment for both sexes. We conclude that flickers respond to proximate cues in brood demands, but do not increase feeding rates to enlarged broods, at least in the short term. A literature review suggested that this lack of response is atypical for short‐lived species. We hypothesize that parents in species with large home ranges and long nestling periods face energetic limitations that constrain their ability to respond to enlarged broods. We encourage future studies to assess foraging behaviour on the landscape to document important trade‐offs for parents such as predation risk and energy expenditure while feeding offspring.     

Patch Use in Free‐Ranging Goats: Does a Large Mammalian Herbivore Forage like Other Central Place Foragers?

Classic central place foraging theory does not focus on the foraging of central place herbivores. This is especially true with regard to large mammalian herbivores. To understand the foraging dynamics of these neglected foragers, we measured giving‐up densities (GUDs) in artificial food patches. We did this at different distances away from the central point (i.e. corral) for a herd of free‐ranging domestic goats. To determine temporal changes, we conducted the study over a 3‐mo period during an extended dry season. Throughout our study, goats foraged across a gradient of food availability where forage was more available farther away from the central point. In contrast to the prediction that predation risk and/or increased travel costs were the main drivers of foraging decisions, we found that the goats increased their feeding effort (i.e. achieved lower GUDs) the farther away they moved from the central point. This suggests that either metabolic or missed opportunity costs were the main factors that influenced foraging decisions. In addition, we suggest that social foraging may have also played a role. With increases in foraging opportunities away from the central point, a herd will likely move slowly while foraging. As a result, individuals can feed intensively from patches but remain part of the group. Ironically, owing to the sustained close proximity of other group members, individuals may perceive patches farther from the central point as being safer. Temporally, the goats increased their feeding effort throughout the dry season. This suggests there was a decline in food quality and/or availability across the environment as the study progressed. Despite this increase in feeding effort, the negative relationship with distance did not change. Ultimately, our results provide key insight into how metabolic, missed opportunity and perceived predation costs influence the feeding decisions of large central place herbivores.     

Effects of seasonal variation in prey abundance on field metabolism, water flux, and activity of a tropical ambush foraging snake

The responses of animals to seasonal food shortages can have important consequences for population dynamics and the structure and function of food webs. We investigated how an ambush foraging snake, the northern death adder Acanthophis praelongus, responds to seasonal fluctuations in prey availability in its tropical environment. In the dry season, field metabolic rates and water flux, as measured by doubly labeled water, were significantly lower than in the wet season. Unlike some other reptiles of the wet-dry tropics, death adders showed no seasonal difference in their resting metabolism. About 94% of the decrease in energy expended in the dry season was due to a decrease in activity and digestion, with lower body temperatures accounting for the remainder. In the dry season, death adders were less active and moved shorter distances between foraging sites than in the wet season. Analysis of energy expenditure suggested that adders fed no more than every 2-3 wk in the dry season but fed more frequently during the wet season. Unlike many lizards that cease feeding during the dry season, death adders remain active and attempt to maximize their energy intake year-round.