Costs of coexistence along a gradient of competitor densities: an experiment with arvicoline rodents

1. Costs of coexistence for species with indirect resource competition usually increase monotonically with competitor numbers. Very little is known though about the shape of the cost function for species with direct interference competition. 2. Here we report the results of an experiment with two vole species in artificial coexistence in large enclosures, where density of the dominant competitor species (Microtus agrestis) was manipulated. Experimental populations of the subordinate vole species (Clethrionomys glareolus) were composed of same aged individuals to study distribution of costs of coexistence with a dominant species within an age-cohort. 3. Survival and space use decreased gradually with increasing field vole numbers. Thus, responses to interference competition in our system appeared to be similar as expected from resource competition. The total number of breeders was stable. Reproductive characteristics such as the timing of breeding, and the litter size were not affected. In the single species enclosures a proportion of surviving individuals were not able to establish a breeding territory against stronger conspecifics. Under competition with heterospecifics such nonbreeders suffered high mortality, whereas the breeders survived. 4. Combined interference of dominant conspecifics and heterospecifics probably increased the frequency of aggressive interactions, social stress and mortality for the weaker individuals within a homogeneous age cohort of the subordinate competitor population. 5. Our results suggest, that in open systems where bank voles are outcompeted over the breeding season by faster reproducing field voles, animals able to establish a territory may be able to withstand competitor pressure, while nonbreeding bank vole individuals are forced to emigrate to suboptimal forest habitats.     

Mechanisms of competition among insectivorous mammals

Summary This study investigates the mechanisms of competition between congeneric pairs of insectivorous mammals in two communities in Australia and England. Direct field observations showed that physical interactions between species do not occur, whereas conspecific encounters are frequent. In field enclosures the smaller, subordinate species in each community (Antechinus stuartii: Marsupialia: Sorex minutus: Eutheria) remained alert in the presence of the dominant species (A. swainsonii, S. araneus), and moved quickly away when the latter approached. The rate of prey capture by subordinate individuals also increased immediately after removal of the dominants. Hourly removals of some individuals of the dominant species in each community over 24 h produced hourly increases in the numbers of subordinate individuals that were captured. The rapidity of these responses suggests strongly that the dominant insectivores in each community interfered with the resource use of the subordinate species. Biomass of invertebrates increased inconsistently or slowly within 3 months of removal of the dominant insectivores; hence the rapid responses by subordinate individuals in experiments were not due to simple exploitation or tracking of resource levels. The subordinate insectivores probably detected and avoided contact with dominants instantaneously using auditory or olfactory cues; reciprocal avoidance of congeneric odours was demonstrated using odour-scented traps. Insectivorous mammals may usually compete by interference (or encounter competition, sensu Schoener 1983). For dominant species within communities the cost of interference is minimal and the benefit of gaining exclusive access to resource-rich microhabitats is high. Conversely for subordinate species the benefit of temporarily exploiting the same rich microhabitats may exceed the small costs of vigilance and movement to nearby refugia.     

Networks and dominance hierarchies: does interspecific aggression explain flower partitioning among stingless bees?

1. The distribution of consumers among resources (trophic interaction network) may be shaped by asymmetric competition. Dominance hierarchy models predict that asymmetric interference competition leads to a domination of high quality resources by hierarchically superior species. 2. In order to determine the competitive dominance hierarchy and its effect on flower partitioning in a local stingless bee community in Borneo, interspecific aggressions were tested among eight species in arena experiments. 3. All species tested were strongly mutually aggressive in the arena, and the observed interactions were often lethal for one or both opponents. Aggression significantly increased with body size differences between fighting pairs and was asymmetric: larger aggressors were superior over smaller species. Additional aggression tests involved dummies with surface extracts, and results suggest that species‐ and colony‐specific surface profiles are important in triggering the aggressive behaviour. 4. Sixteen stingless bee species were observed foraging on 41 species of flowering plants. The resulting bee–flower interaction network showed a high degree of generalisation (network‐level specialisation H₂’ = 0.11), corresponding to a random, opportunistic distribution of bee species among available flower species. 5. Aggressions on flowers were rare and only occurred at a low level. The dominance hierarchy obtained in the arena experiments did not correlate significantly with plant quality, estimated as the number of flowers per plant or as total bee visitation rate. 6. Our findings suggest that asymmetries in interference competition do not necessarily translate into actual resource partitioning in the context of complex interacting communities.     

Prey preferences and recent changes in diet of a breeding population of the Northern Goshawk Accipiter gentilis in Southwestern Europe

Capsule: Northern Goshawk Accipiter gentilis diet has changed significantly since the 1980s, probably due to changes in populations of preferred prey species.

Aims and methods: To assess changes to the breeding season diet of the Northern Goshawk in southwest Europe over three decades. We examined prey remains at and around nests and assessed avian prey availability using point count and line transect surveys.

Results: During 2008–11, Goshawks mainly ate birds, with Feral Pigeons Columba livia f. domestica being the most important prey species. Goshawks preferred prey of 100?400?g and forest prey species to non-forest species. Goshawk diet has changed significantly over recent decades: 22% of current prey items belong to species that were not part of the diet in the 1980s. We suggest that these dietary changes reflect changes in the abundance of prey species of the preferred size caused by changes in land use leading to an increase in forest cover, new prey species colonization and changes in the abundance and management of domestic prey.

Conclusion: This study emphasizes that major transformations occurring in agroforestry systems are affecting the main preferred prey of important forest predators, which may have consequences for conservation of both the predators and their prey.     

Breeding state and season affect interspecific interaction types: indirect resource competition and direct interference

Indirect resource competition and interference are widely occurring mechanisms of interspecific interactions. We have studied the seasonal expression of these two interaction types within a two-species, boreal small mammal system. Seasons differ by resource availability, individual breeding state and intraspecific social system. Live-trapping methods were used to monitor space use and reproduction in 14 experimental populations of bank voles Myodes glareolus in large outdoor enclosures with and without a dominant competitor, the field vole Microtus agrestis. We further compared vole behaviour using staged dyadic encounters in neutral arenas in both seasons. Survival of the non-breeding overwintering bank voles was not affected by competition. In the spring, the numbers of male bank voles, but not of females, were reduced significantly in the competition populations. Bank vole home ranges expanded with vole density in the presence of competitors, indicating food limitation. A comparison of behaviour between seasons based on an analysis of similarity revealed an avoidance of costly aggression against opponents, independent of species. Interactions were more aggressive during the summer than during the winter, and heterospecific encounters were more aggressive than conspecific encounters. Based on these results, we suggest that interaction types and their respective mechanisms are not either-or categories and may change over the seasons. During the winter, energy constraints and thermoregulatory needs decrease direct aggression, but food constraints increase indirect resource competition. Direct interference appears in the summer, probably triggered by each individual's reproductive and hormonal state and the defence of offspring against conspecific and heterospecific intruders. Both interaction forms overlap in the spring, possibly contributing to spring declines in the numbers of subordinate species.     

Predicting community structure of ground-foraging ant assemblages with Markov models of behavioral dominance

Although interference competition is a conspicuous component of many animal communities, it is still uncertain whether the competitive ability of a species determines its relative abundance and patterns of association with other species. We used replicated arena tests to quantify behavioral dominance of eight common species of co-occurring ground-foraging ants in the Siskiyou Mountains of southern Oregon. We found that behavior recorded in laboratory assays was an accurate representation of a colony's ability to monopolize resources in the field. We used interaction frequencies from the behavioral tests to estimate transition probabilities in a simple Markov chain model to predict patterns of relative abundance in a metacommunity that is dominated by behavioral interactions. We also tested whether behavioral interactions between each pair of species could be used to predict patterns of species co-occurrence. We found that the Markov model did not accurately predict patterns of observed relative abundance on either the local or the regional scale. However, we did detect a significant negative correlation at the local scale in which behaviorally dominant species occupied relatively few baits. Pairwise behavioral data also did not predict species co-occurrence in any site. Although interference competition is a conspicuous process in ant communities, our results suggest that it may not contribute much to patterns of relative abundance and species co-occurrence in the system studied here. However, the negative correlation between behavioral dominance and bait occupancy at the local scale suggests that competition-colonization trade-offs may be important in resource acquisition and persistence of behaviorally subordinate species.     

Analysis of species coexistence co-mediated by resource competition and reproductive interference

Reproductive interference is any interspecific sexual interaction that reduces the reproductive success of females through promiscuous reproductive activities of heterospecific individuals. This phenomenon is ubiquitous in nature in both plants and animals, and is frequently observed in biological invasions. However, its effects on interspecific competition remain incompletely understood despite growing concern. To study the interactive effects of resource competition and reproductive interference on species coexistence and exclusion, we analyzed a unified competition model including both processes in symmetric and asymmetric scenarios. The results of our model showed that resource competition and reproductive interference act synergistically to promote competitive exclusion. We also found that when the two processes are asymmetric, the species that is superior in reproductive interference can coexist with or even exclude the species that is superior in resource competition. Therefore, coexistence is possible via an unbalanced trade-off between resource use and reproduction. Our results suggest that integration of reproductive interference and resource competition will contribute to a better understanding of interspecific competition and to more effective biodiversity conservation against management of biological invasions.     

Ants versus spiders: interference competition between two social predators

Interference competition can profoundly influence the outcome of species interactions and may lead to either coexistence or exclusion. Our understanding of how interference can lead to coexistence remains incomplete, particularly when interference fails to result in resource partitioning. I document a novel form of interference competition between an ant (Myrmecia pyriformis) and a social spider (Delena cancerides) with similar foraging patterns. Of 120 nest boxes occupied by D. cancerides in the field, 7 (6 %) were invaded by M. pyriformis ants over a 2-month period. After eliminating spiders from the nest boxes, the ants proceeded to fill the boxes with debris, rendering them useless to the spiders. The ants do not occupy the nest boxes; thus, interference occurs at a resource that is necessary to one species, but not the other. I discuss how further research into this system may improve our understanding of how interference competition can support coexistence. I also suggest modeling exploitation and interference competition on multiple resources to align with this and other empirical examples where different forms of competition occur for different resources.     

In the absence of a “landscape of fear”: How lions,hyenas, and cheetahs coexist

Aggression by top predators can create a “landscape of fear” in which subordinate predators restrict their activity to low‐risk areas or times of day. At large spatial or temporal scales, this can result in the costly loss of access to resources. However, fine‐scale reactive avoidance may minimize the risk of aggressive encounters for subordinate predators while maintaining access to resources, thereby providing a mechanism for coexistence. We investigated fine‐scale spatiotemporal avoidance in a guild of African predators characterized by intense interference competition. Vulnerable to food stealing and direct killing, cheetahs are expected to avoid both larger predators; hyenas are expected to avoid lions. We deployed a grid of 225 camera traps across 1,125 km² in Serengeti National Park, Tanzania, to evaluate concurrent patterns of habitat use by lions, hyenas, cheetahs, and their primary prey. We used hurdle models to evaluate whether smaller species avoided areas preferred by larger species, and we used time‐to‐event models to evaluate fine‐scale temporal avoidance in the hours immediately surrounding top predator activity. We found no evidence of long‐term displacement of subordinate species, even at fine spatial scales. Instead, hyenas and cheetahs were positively associated with lions except in areas with exceptionally high lion use. Hyenas and lions appeared to actively track each, while cheetahs appear to maintain long‐term access to sites with high lion use by actively avoiding those areas just in the hours immediately following lion activity. Our results suggest that cheetahs are able to use patches of preferred habitat by avoiding lions on a moment‐to‐moment basis. Such fine‐scale temporal avoidance is likely to be less costly than long‐term avoidance of preferred areas: This may help explain why cheetahs are able to coexist with lions despite high rates of lion‐inflicted mortality, and highlights reactive avoidance as a general mechanism for predator coexistence.     

Interactions among salt marsh plants vary geographically but not latitudinally along the California coast

The strength of species interactions often varies geographically and locally with environmental conditions. Competitive interactions are predicted to be stronger in benign environments while facilitation is expected to be stronger in harsh ones. We tested these ideas with an aboveground neighbor removal experiment at six salt marshes along the California coast. We determined the effect of removals of either the dominant species, Salicornia pacifica, or the subordinate species on plant cover, aboveground biomass and community composition, as well as soil salinity and moisture. We found that S. pacifica consistently competed with the subordinate species and that the strength of competition varied among sites. In contrast with other studies showing that dominant species facilitate subordinates by moderating physical stress, here the subordinate species facilitated S. pacifica shortly after removal treatments were imposed, but the effect disappeared over time. Contrary to expectations based on patterns observed in east coast salt marshes, we did not see patterns in species interactions in relation to latitude, climate, or soil edaphic characteristics. Our results suggest that variation in interactions among salt marsh plants may be influenced by local‐scale site differences such as nutrients more than broad latitudinal gradients.     

Habitat Interference by Axis Deer on White-Tailed Deer

Abstract: Many studies of interactions between exotic and native ungulates have not had temporal and spatial controls nor have they considered the types of competitive interactions that would allow coexistence. For exotic axis deer (Axis axis) and native white-tailed deer (Odocoileus virginianus) to coexist one species should be superior at interference competition and the other species should be superior at exploitative competition. We generated and tested predictions, based on body size and diet breadth, about habitat selection by white-tailed deer in the presence and absence of axis deer, dominance relationships, and time at sites provisioned with high quality forage. We conducted our study in treatment (axis and white-tailed deer) and control (white-tailed deer only) areas when both species were present and after axis deer were removed. We conducted vehicle surveys to determine habitat use of both species. At provisioned feeding sites we recorded aggressive behaviors and amount of time species spent at feeding sites alone and together. In the treatment area white-tailed deer selection for wooded habitat increased 2.1 times after axis deer were removed, whereas habitat selection by white-tailed deer was constant in the control area over the same time. At feeding sites axis deer were dominant to white-tailed deer; both species spent a significantly greater amount of time alone than at feeders together, but amount of time that individuals of each species spent at feeders did not differ. Axis deer were superior at interference competition, but white-tailed deer were not superior at exploitative competition; thus, species coexistence is unlikely. Whether white-tailed deer are negatively impacted by axis deer at spatial scales larger than our experiment probably depends on abundance of axis deer at larger spatial scales. Experiments of species interactions with temporal and spatial controls that consider types of competitive interactions increase a manager's understanding of when and how native ungulates may be negatively impacted by exotic ungulates.     

The role of resource dispersion in promoting the co‐occurrence of dominant and subordinate ant species

Dominant competitors govern resource use in many communities, leading to predictions of local exclusion and lower species diversity where dominant species are abundant. However, subordinate and dominant species frequently co‐occur. One mechanism that could facilitate resource sharing and co‐occurrence of dominant and subordinate competitors is fine‐scale resource dispersion. Here, we distributed 6 g of a food resource into 1, 2, 8, 32 or 64 units in small 0.40 m² areas centred on nests of the dominant ant Monomorium sydneyense. We tested three hypotheses. First, we hypothesized that the species richness and abundance of foraging ants would increase with increasing resource dispersion. Accordingly, species richness doubled and total ant abundance was two orders of magnitude higher in high resource dispersion treatments. Secondly, we hypothesized that increasing resource dispersion would reduce competitive interactions such as resource turnover events and lower the probability of food resources being occupied. Substantial support for this hypothesis was observed. Finally, we tested the hypothesis that the foraging time of each species would be proportional to the relative abundance of each species solely in high resource dispersion treatments. Expected and observed foraging times were statistically similar for only the dominant ant M. sydneyense. The subdominant Pheidole rugosula increased its foraging time much more than was expected, while two subordinate ants showed no relationship between observed and expected times. Thus, while increasing resource dispersion significantly increased overall species richness, this increase in co‐occurrence did not correlate with a significant increase in foraging time for the two subordinate species. Rather, changes in resource dispersion appeared to benefit only the subdominant species. Inter‐site variation appeared more important for other subordinate species indetermining co‐occurrence and foraging time. Multiple mechanisms facilitate co‐occurrence and resource sharing in this community, and probably in most other communities.     

The nested assembly of individual-resource networks

1. Much of the current understanding of ecological systems is based on theory that does not explicitly take into account individual variation within natural populations. However, individuals may show substantial variation in resource use. This variation in turn may be translated into topological properties of networks that depict interactions among individuals and the food resources they consume (individual-resource networks). 2. Different models derived from optimal diet theory (ODT) predict highly distinct patterns of trophic interactions at the individual level that should translate into distinct network topologies. As a consequence, individual-resource networks can be useful tools in revealing the incidence of different patterns of resource use by individuals and suggesting their mechanistic basis. 3. In the present study, using data from several dietary studies, we assembled individual-resource networks of 10 vertebrate species, previously reported to show interindividual diet variation, and used a network-based approach to investigate their structure. 4. We found significant nestedness, but no modularity, in all empirical networks, indicating that (i) these populations are composed of both opportunistic and selective individuals and (ii) the diets of the latter are ordered as predictable subsets of the diets of the more opportunistic individuals. 5. Nested patterns are a common feature of species networks, and our results extend its generality to trophic interactions at the individual level. This pattern is consistent with a recently proposed ODT model, in which individuals show similar rank preferences but differ in their acceptance rate for alternative resources. Our findings therefore suggest a common mechanism underlying interindividual variation in resource use in disparate taxa.     

Spatiotemporal food web dynamics along a desert riparian–upland transition

Increased awareness of spatiotemporal variation in species interactions has motivated the study of temporally-resolved food web dynamics at the landscape level. Empiricists have focused attention on cross-habitat flows of materials, nutrients, and prey, largely ignoring the movement of predators between habitats that differ in productivity (and how predators integrate pulses in resource availability over time). We set out to study seasonal variation in food web interactions between mammalian carnivores and their rodent prey along a riparian–upland gradient in semi-arid southeastern Arizona which features both spatial and temporal heterogeneity in resource availability. Specifically, we tested whether mammalian carnivores spill over from productive, near-river habitats into adjacent, desert-scrub habitats; and if they do, to document the effects of this spillover on rodent communities. Furthermore, we examined seasonal variation in top-down effects by measuring changes in carnivore diet and distribution patterns and rodent populations over time. The results indicate that carnivores track seasonally-abundant resources across the landscape, varying both their diet and movement patterns. In turn, desert-scrub rodent population dynamics track seasonal shifts in carnivore habitat use but not resource availability, suggesting that predation plays a role in structuring rodent communities along the San Pedro River. Further evidence comes from data on rodent community composition, which differs between desert-scrub habitats near and far from the river, despite similarities in resource availability. Our data also suggest that seasonal omnivory helps predators survive lean times, increasing their effects on prey populations. Taken together, these results underscore the importance of spatiotemporal variation in species interactions, highlighting the complexity of natural systems and the need for further detailed studies of food web dynamics.     

Mycophagous beetle females do not behave competitively during intrasexual interactions in presence of a fungal resource

Intrasexual interactions can determine which individuals within a population have access to limited resources. Despite their potential importance on fitness generally and mating success especially, female–female interactions are not often measured in the same species where male–male interactions are well‐defined. In this study, we characterized female–female interactions in Bolitotherus cornutus, a mycophagous beetle species native to Northeastern North America. We used dyadic, behavioral assays to determine whether females perform directly aggressive or indirectly exclusionary competitive behaviors. Polypore shelf fungus, an important food and egg‐laying resource for B. cornutus females, is patchily distributed and of variable quality, so we tested for competition over fungus as a resource. Behavior of females was assessed in three sets of dyadic trials with randomly paired female partners. Overall, females did not behave aggressively toward their female partner or perform exclusionary behaviors over the fungal resource. None of the behaviors performed by females were individually repeatable. Two scenarios may explain our lack of observed competition: our trial context may not induce competition, or female B. cornutus simply may not behave competitively in the wild. We compare our results to a similar study on male–male interactions in the same species and propose future studies on female–female interactions under different competitive contexts to expand the understanding of female competition.     

The Effect of Social Hierarchy on Captive Coyote (Canis latrans) Foraging Behavior

Foraging efficiency of individuals in pack forming species may be influenced by social dynamics within a pack. The effects of social hierarchy in particular may influence individual foraging behavior in canids, such as coyotes (Canis latrans). To examine the impact of social hierarchy on foraging behavior, we tested 16 captive coyotes in eight naturally established dominant–subordinate pairs, using the guesser–knower paradigm. We measured the efficiency of subordinate coyotes to relocate a food resource when alone and then allowed pairs to forage together, such that subordinates had prior knowledge of food location but dominants did not. To determine whether (1) subordinates used a direct or discursive strategy to obtain food in the presence of a dominant and (2) dominants used an exploitative or independent strategy to obtain food in the presence of a subordinate with previous knowledge, we measured their search efficiency (e.g., correct choice of area, feeder, and latency to correct feeder). Results showed subordinates learned to relocate food and increase efficiency when alone. In a social context, however, subordinate efficiency decreased. That is, subordinates approached the correct area, but searched more feeders before finding the correct one. Dominants initially used an independent search strategy but then quickly displaced the subordinate and monopolized the resource, reducing subordinate efficiency further. Despite continual displacement and reduction in efficiency, subordinates did not alter their foraging strategy over time. Our results suggest prior information can improve individual foraging advantage, but that social status strongly impacts individual foraging efficiency in social species such as coyotes.     

Connectance and parasite diet breadth in flea-mammal webs

The number of links in webs of species interactions, which lies at the heart of the biodiversity-stability debate, has given rise to controversy during the last 20 yr. Studies exploring these web properties have mainly focused on symmetric webs where each species can potentially feed on any other species; asymmetric webs such as host-parasite webs, where one set of species feed on another set of species, have been overlooked. However, food webs are incomplete without parasites and the study of parasite-host sub-web properties deserves attention. Here, using a large database involving 33 regional interaction webs between mammals and their flea parasites, we found a negative relationship between species richness and host-parasite connectance. We suggest that some phylogenetic constraints on flea diet may explain our observed patterns because we found that parasite diet breadth, measured as host taxonomic diversity, was invariant along our host richness gradient. We found that the slope of the logarithmic relationship between the number of realized links and species richness is lower than slope values reported for food webs. We suggest that connectance may not respond to increasing species richness as rapidly in host-parasite webs as in predator-prey food webs due to stronger coevolutionary requirements.     

Spatial relationships and mechanisms of coexistence between dominant and subordinate top predators

Most forest ecosystems contain a diverse community of top‐level predators. How these predator species interact, and how their interactions influence their spatial distribution is still poorly understood. Here we studied interactions among top predators in a guild of diurnal forest raptors in order to test the hypothesis that predation among competing predators (intraguild predation) significantly affects the spatial distribution of predator species, causing subordinate species to nest farther away from the dominant ones. The study analyzed a guild in southwestern Europe comprising three raptor species. For 8 years we studied the spatial distribution of used nests, breeding phenology, intraguild predation, territory occupancy, and nest‐builder species and subsequent nest‐user species. The subordinate species (sparrowhawk Accipiter nisus) nested farther away from the dominant species (goshawk A. gentilis), which preyed on sparrowhawks but not on buzzards Buteo buteo, and closer to buzzards, with which sparrowhawks do not share many common prey. This presumably reflects an effort to seek protection from goshawks. This potential positive effect of buzzards on sparrowhawks may be reciprocal, because buzzards benefit from old sparrowhawk nests, which buzzards used as a base for their nests, and from used sparrowhawk nests, from which buzzards stole prey. Buzzards occasionally occupied old goshawk nests. These results support our initial hypothesis that interspecific interactions within the raptor guild influence the spatial distribution of predator species in forest ecosystems, with intraguild predation as a key driver. We discuss several mechanisms that may promote the coexistence of subordinate and dominant predators and the spatial assembly of this raptor guild: spatial refuges, different breeding phenology, spatial avoidance, low territory occupancy between neighboring nesting territories, nest concealment and protection, and diet segregation.     

The role of competition by dominants and temperature in the foraging of subordinate species in Mediterranean ant communities

In this paper we test the influence of temperature and interference competition by dominant species on the foraging of subordinate species in Mediterranean ant communities. We have analyzed the changes in resource use by subordinate species in plots with different abundances of dominant ants, and in different periods of the day and the year, i.e., at different temperatures. The expected effects of competition by dominant species on foraging of subordinates were only detected for two species in the number of baits occupied per day, and for one species in the number of foragers at pitfall traps. In all three cases, subordinate species were less represented at baits or in traps in plots with a high density of dominants than in plots with a medium or low density of dominants. The number of workers per bait, and the foraging efficiency of subordinate species did not differ in plots differing in dominant abundance. Daily activity rhythms and curves of temperature versus foraging activity of subordinate species were also similar in plots with different abundance of dominant species, indicating no effect of dominants on the foraging times of subordinates. Instead, temperature had a considerable effect on the foraging of subordinate species. A significant relationship was found between maximum daily temperature and several variables related to foraging (the number of foragers at pitfall traps, the number of baits occupied per day, and the number of workers per bait) of a number subordinate species, both in summer and autumn. These results suggest that the foraging of subordinate ant species in open Mediterranean habitats is influenced more by temperature than by competition of dominants, although an effect of dominants on subordinates has been shown in a few cases. In ant communities living in these severe and variable environments, thermal tolerance reduces the importance of competition, and the mutual exclusion usually found between dominant and subordinate species appears to be the result of physiological specialization to different temperature ranges. Received: 8 May 1998 / Accepted: 30 July 1998     

Seasonal variation in diet breadth of folivorous Lepidoptera in the Brazilian cerrado

Documented patterns of specialization and species interactions often omit plasticity in resource use across space and time, yet such variation is an important part of species interactions. To examine temporal variation in resource use, we compared species‐ and community‐level patterns of host plant use by folivorous caterpillars between the dry and rainy seasons in four preserved areas of cerrado vegetation in the Distrito Federal, Brazil. We sampled plants and caterpillars in 10 m circular plots monthly from March 2010 to March 2011. At the community level, we found a significant increase in the mean diet breadth of dry season caterpillars relative to those collected in the rainy season. Families and species of moths varied in diet breadth, but most exhibited a seasonal expansion. In particular, intraspecific comparisons showed a 30% increase in the number of host plants used in the dry season compared to the rainy season. These results provide a clear example of how temporal variation in resource use is heterogeneous, and more generalized patterns of resource use can emerge from studies at large temporal scales. Thus, seasonal or annual heterogeneity may obscure ecologically relevant specialized interactions that occur at smaller scales.