Manipulated Resource Availability Research Articles

Seasonal Patterns of Resource Use Within Natural Populations of Burying Beetles.

For organisms in temperate environments, seasonal variation in resource availability and weather conditions exert fluctuating selection pressures on survival and fitness, resulting in diverse adaptive responses. By manipulating resource availability on a local spatial scale, we studied seasonal patterns of resource use within natural populations of burying beetles Nicrophorus vespilloides in a Norfolk woodland. Burying beetles are necrophagous insects that breed on vertebrate carcasses. They are active in Europe between April and October, after which they burrow into the soil and overwinter. Using breeding and chemical analyses, we compared the fecundity and physiological state of beetles that differed in their seasonal resource use. We found seasonal variation in carrion use by wild burying beetles and correlated differences in their reproductive success and cuticular hydrocarbon profiles. Our results provide novel insight into the seasonal correlates of behaviour, physiology and life history in burying beetles.

The cost of adaptability: resource availability constrains functional stability under pulsed disturbances.

Understanding the communities' responses to disturbances is a prerequisite to predicting ecosystem dynamics and, thus, highly relevant considering global change. Microbial communities play key roles in numerous ecosystem functions and services, and the large diversity, rapid growth, and phenotypic plasticity of microorganisms are thought to allow high resistance and resilience. While potential metabolic costs associated with adaptations to fluctuating environments have been debated, little evidence supports trade-offs between resource availability, resistance, and resilience. Here, we experimentally assessed the compositional and functional responses of an aquatic microbial model community to disturbances and systematically manipulated resource availability. Our results demonstrate that the capacity to tolerate environmental fluctuations is constrained by resource availability and reflected in the selection of genomic traits.

Resource availability and plant age drive defense against herbivory in salt marshes

The resource‐availability hypothesis (RAH) and the intraspecific RAH (RAHintra), posit that resources, (i.e. nutrients) control plant antiherbivore defenses. Both hypotheses predict that in low‐resource environments, plant growth is slow, and constitutive defense is high. In high‐resource environments, however, the RAH predicts that plant growth is fast, and constitutive defense is low, whereas the RAHintra predicts that increased resources attract more herbivores, and this intensified grazing pressure leads to high constitutive defense. Salt marshes are nutrient‐limited ecosystems threatened by eutrophication and chronic herbivory, yet we know little about how these stressors shape saltmarsh plant antiherbivore defenses, which influence trophic interactions and ecosystem resilience. We manipulated resource availability via nutrient addition and herbivory via the marsh periwinkle Littoraria irrorata, on the saltmarsh foundation species Spartina alterniflora, in mesocosms. Because plant age can also influence trait variation, we measured traits in both original and clonally‐grown new stems. Feeding assays then evaluated how treatments and plant age affected subsequent Littoraria consumption of Spartina. Nutrient addition stimulated growth, while decreasing defensive traits (e.g. fiber and silica content), following the RAH. Herbivory enhanced belowground production and increased stem diameter, yet did not induce defensive traits, contrary to our expectations. Herbivory plus nutrients increased Spartina biomass and reduced phenolics, a defensive trait, further supporting the RAH. Regardless of treatment, clonally‐grown new stems had greater variation in measured traits. Despite altered traits, however, treatments and plant age did not affect Littoraria consumption. Our results support the RAH and part of the RAHintra and suggest: 1) nutrient availability is a primary driver of plant trait change and 2) plant age controls the magnitude of trait variation in Spartina. Further, our findings indicate that eutrophic conditions may not always increase top‒down control by herbivores, and in some instances can enhance saltmarsh resilience against sea‐level rise via stimulated Spartina biomass production.

Interacting water, nutrients, and shrub age control steppe grass‐on‐shrub competition: Implications for restoration

AbstractUnderstanding the factors that control grass–shrub interactions and coexistence is critical to the design of dryland management and restoration strategies. Using Stipa tenacissima tussocks as adult neighbor and Olea europaea var. sylvestris as target woody species, we performed a 5‐year‐long, large‐scale manipulative experiment to investigate the independent and combined effects of water and nutrient availability, and the modulating effects of O. europaea ontogeny on the net outcome of grass–shrub interaction. At two contrasting dryland sites, we conducted experimental plantings of Olea seedlings on two microsites: contiguous to a Stipa tussock (Stipa microsite) and on the intertussock bare‐soil areas (Open microsite), and manipulated resource availability by combining nutrient and water addition. The experiment followed a full factorial design, with all treatment combinations applied at each site. We monitored survival, height, and stem basal diameter of Olea individuals over a study period of 5 years. All across the wide range of conditions considered, negative effects of Stipa on Olea largely prevailed during the first 2 years after the plantation. We found that competition was stronger in the drier steppe, where extra inputs of water favored neutral interactions. Conversely, nutrient addition increased competition strength, particularly when combined with water inputs, pointing to the contrasting role of different resources and the importance of interactions between them in the control of plant–plant interaction outcome. The competition effects of Stipa faded with time and gradually shifted to neutral interaction as Olea individuals aged. Our results are particularly relevant to guide restoration efforts in formerly wooded xeric grasslands and rangelands.

Condition-dependent sexual reproduction is driven by benefits, not costs of sex

AbstractFacultative sexual organisms must allocate resources to both asexual and sexual reproduction. Optimal patterns of investment in sex depend on the relative costs and benefits of each reproductive mode, and may consequently be context- and condition-dependent. Two proposed explanations for the observed variation in investment in sex among facultative sexual lineages invoke alternative condition-dependent scenarios. Under the fitness-associated sex hypothesis, sex is predicted when individuals are in poor condition or experience stressful environments. Under the resource-demanding sex hypothesis, sex is only affordable to individuals in good condition experiencing favourable environments. Direct tests of these contrasting hypotheses are rare; moreover, investment in different components of sexual reproduction responds differently to cues promoting sex, and may be subject to different energetic constraints. Using genotypes of facultative sexual Daphnia carinata that differ in their level of investment in sex, we manipulated resource availability while accounting for day length (a seasonal cue for sex) to evaluate these hypotheses. The sexual response to day length depended on resource availability: increased day lengths and reduced food availability increased the production of sexual eggs, and relative investment in males, in a manner consistent with the fitness-associated sex hypothesis. The pattern of condition-dependence was specific to each component of reproductive investment – while male production covaried with asexual fecundity across genotypes, increased sexual egg production was associated with reduced asexual reproduction. Our results suggest that investment in sex is determined largely by its context-dependent advantages, and that this investment is not moderated by immediate costs to asexual reproduction.

The price of safety: food deprivation in early life influences the efficacy of chemical defence in an aposematic moth

Aposematism is the combination of a primary signal with a secondary defence that predators must learn to associate with one another. However, variation in the level of defence, both within and between species, is very common. As secondary defences influence individual fitness, this variation in quality and quantity requires an evolutionary explanation, particularly as it may or may not correlate with variation in primary signals. The costs of defence production are expected to play a considerable role in generating this variation, yet studies of the cost of chemical defence have focused on species that sequester their defences, while studies in species that produce them de novo are scarce. Here we examine effects of resource availability on the production of chemical defences in female wood tiger moths Arctia plantaginis. This species produces defensive fluids when attacked, and advertises this protection with bright colours on its hindwings. Furthermore, the chemicals in these fluids are produced de novo. In order to establish if the production of this chemical protection is costly, and thus resource‐limited, we manipulated resource availability (i.e. food) during larval development and measured its subsequent effects on adult chemical defence. We presented defensive fluids from female moths to wild blue tits, in the absence of any visual cues, to determine their effectiveness against avian predators. Our results demonstrate that the defensive fluids produced by female moths repel birds, and that these defences require resource investment to produce. We found that moths that were resource‐limited during development had less effective chemical defences, despite producing the same volume of defensive fluids as the control moths. As a reduction in available resources negatively influences the effectiveness of the chemical defence, resource availability may explain the variation seen among individuals.

Field manipulations of resources mediate the transition from intraspecific competition to facilitation.

Population density affects individual performance, though its effects are often mixed. For sessile species, increases in population density typically reduce performance. Still, cases of positive density-dependence do occur in sessile systems and demand explanation. The stress gradient hypothesis (SGH) predicts that under stressful conditions, positive effects of facilitation may outweigh the negative effects of competition. While some elements of the SGH are well studied, its potential to explain intraspecific facilitation has received little attention. Further, there have been questions regarding whether the SGH holds if the stressor is a resource. Most studies of interactions between the environment and intraspecific facilitation have relied on natural environmental gradients; manipulative studies are much rarer. To test the effects of intraspecific density and resources, we manipulated resource availability over natural population densities for the marine bryozoan Watersipora subtorquata. We found negative effects of density on colony performance in low resource environments, but mainly positive density-dependence in high resource environments. By adding resources, competition effects were reduced and the positive effects of facilitation were revealed. Our results suggest that resource availability mediates the relative strength of competition and facilitation in our system. We also suggest that intraspecific facilitation is more common than may be appreciated and that environmental variation may mediate the balance between negative and positive density-dependence.

Nutrient enrichment alters impacts of Hydrocotyle vulgaris invasion on native plant communities.

Nutrients may affect the invasiveness of alien plants and the invasibility of native plant communities. We performed a greenhouse experiment to investigate the interactive effect of invasion by a clonal herb Hydrocotyle vulgaris and nutrient enrichment on biomass and evenness of native plant communities. We established three types of plant communities (H. vulgaris alone, native plant communities without or with H. vulgaris) under low and high levels of nutrients. Native communities consisted of eight native, terrestrial species of three functional groups, i.e. four grasses, two legumes, and two forbs. Invasion of H. vulgaris had no effect on biomass of the native community, the functional groups, or the individual species. High nutrients increased biomass of grasses, but reduced evenness of the community. High nutrients also decreased the competitive effect, and the relative dominance index of H. vulgaris. Therefore, high nutrients reduced the competitive ability of H. vulgaris and enhanced the resistance of the native community to invasion. The results provide a basis for management strategies to control the invasion and spread of H. vulgaris by manipulating resource availability to support native communities.

Maternal manipulation of pollen provisions affects worker production in a small carpenter bee

Mothers play a key role in determining the body size, behavior, and fitness of offspring. Mothers of the small carpenter bee, Ceratina calcarata, provide smaller pollen balls to their first female offspring resulting in the development of a smaller female. This smaller female, known as the dwarf eldest daughter, is coerced to stay at the nest to forage and feed siblings as a worker. In order to better understand how this maternal manipulation leads to the physiological and behavioral differences observed in dwarf eldest daughters, we characterized and compared the quality of the pollen balls fed to theses females vs. other offspring. Our results confirm earlier studies reporting that there is a female-biased sex allocation in the first brood cell position and these daughters received mass provisions significantly smaller than other daughters. In addition to the smaller quantities of pollen provisioned, we found evidence for maternal control of the quality of pollen invested in the dwarf eldest daughters. Late brood cells receive pollen balls with significantly less floral diversity than early brood cells. This difference in floral diversity affects the protein content of the pollen balls; in that, older offspring receive less protein than their younger siblings. These results reveal that C. calcarata mothers manipulate not only the quantity but also the quality of the provision provided to her first offspring to create a small worker she is able to coerce to remain at the nest to help raise her siblings. This overlapping of generations and division of labor between mother and dwarf eldest daughter may represent the first steps in the evolution of highly social groups. One of the major transitions to the formation of highly social groups is division of labor. By manipulating resource availability to offspring, parents can force offspring to remain at the nest to serve as a worker leading to a division of labor between parent and offspring. In the small carpenter bee, C. calcarata, mothers provide their eldest daughter with less food resulting in a smaller adult body size. This dwarf eldest daughter (DED) does not have the opportunity to reproduce and serves only as a worker for the colony. In addition to overall reduced investment, we found that mothers also provide a different variety of pollen to her DED. By exploring the factors and mechanisms that influence maternal manipulation in a non-eusocial bee, we can begin to understand one of the major transitions in social group formation.

Can the Bacterial Community of a High Arctic Glacier Surface Escape Viral Control?

Glacial ice surfaces represent a seasonally evolving three-dimensional photic zone which accumulates microbial biomass and potentiates positive feedbacks in ice melt. Since viruses are abundant in glacial systems and may exert controls on supraglacial bacterial production, we examined whether changes in resource availability would promote changes in the bacterial community and the dynamics between viruses and bacteria of meltwater from the photic zone of a Svalbard glacier. Our results indicated that, under ambient nutrient conditions, low estimated viral decay rates account for a strong viral control of bacterial productivity, incurring a potent viral shunt of a third of bacterial carbon in the supraglacial microbial loop. Moreover, it appears that virus particles are very stable in supraglacial meltwater, raising the prospect that viruses liberated in melt are viable downstream. However, manipulating resource availability as dissolved organic carbon, nitrogen, and phosphorous in experimental microcosms demonstrates that the photic zone bacterial communities can escape viral control. This is evidenced by a marked decline in virus-to-bacterium ratio (VBR) concomitant with increased bacterial productivity and number. Pyrosequencing shows a few bacterial taxa, principally Janthinobacterium sp., dominate both the source meltwater and microcosm communities. Combined, our results suggest that viruses maintain high VBR to promote contact with low-density hosts, by the manufacture of robust particles, but that this necessitates a trade-off which limits viral production. Consequently, dominant bacterial taxa appear to access resources to evade viral control. We propose that a delicate interplay of bacterial and viral strategies affects biogeochemical cycling upon glaciers and, ultimately, downstream ecosystems.

Impacts of alien plant invasion on native plant communities are mediated by functional identity of resident species, not resource availability

Alien plant invasion and nutrient enrichment as a result of anthropogenic landscape modification seriously threaten native plant community diversity. It is poorly understood, however, whether these two disturbances interact with the functional identity of recipient native plants to drive community change. We performed a mesocosm experiment to examine whether the interactive effects of invasion by a stoloniferous turf‐grass Stenotaphrum secundatum and nutrient enrichment vary across different plant growth forms of an endangered coastal plant community. Communities contained 18 species (drawn without replacement from a pool of 31 species) with either runner, tufted or woody growth forms. Species were well‐established and reproductively mature prior to S. secundatum introduction. Species growth (% cover), reproductive output, soil temperature and light availability were monitored for two growing seasons. Invasion and nutrient enrichment (two levels: ‘natural control’ and ‘enriched’) had no effect on species richness, community composition, reproductive output, soil temperature or light penetration. There was no interactive effect of nutrients and invasion on community productivity (i.e. final biomass), such that invasion caused a reduction in community biomass at both natural and enriched nutrient levels. This was driven only by reduced biomass of functionally‐similar native runner species, which share similar root morphologies and nutrient‐acquisition strategies with S. secundatum. Our study indicates that impacts of invasion are dependent upon the functional identity of species within recipient communities, not the availability of resources. This shows that management cannot buffer invader effects by manipulating resource availability. Revegetation strategies should target functionally‐similar natives for replacement following invader control.

The relative importance of competition and predation in environment characterized by resource pulses--an experimental test with a microbial community.

BackgroundResource availability and predation are believed to affect community dynamics and composition. Although the effects of resource availability and predation on prey communities are usually studied in isolation, these factors can also have interactive effects, especially since the outcome of competition under shared predation is expected to depend on resource availability. However, there are few experimental studies that test the interactive roles of resources and predation on dynamics of more complex multispecies communities. Here, we examine the importance of competition and predation on microbial community dynamics in a resource pulse environment.ResultsWe manipulated resource availability and predation simultaneously in a microbial microcosm experiment, where a bacterial community was exposed to the protozoan predator Tetrahymena thermophila in three different resource concentrations (low, intermediate and high). The prey community consisted of three heterotrophic bacterial species: Bacillus cereus, Serratia marcescens and Novosphingobium capsulatum, all feeding on a shared plant detritus medium. In fresh culture media, all species grew in all resource concentrations used. However, during experiments without any addition of extra resources, the existing resources were soon depleted to very low levels, slowing growth of the three bacterial species. Prior to the microcosm experiment, we measured the competitive ability and grazing resistance, i.e. reduced vulnerability to predation, of each prey species. The three species differed in allocation patterns: in general, N. capsulatum had the best competitive abilities and B. cereus had good grazing resistance abilities. In the long-term microcosm experiment, N. capsulatum dominated the community without predation and, with predation, B. cereus was the dominant species in the intermediate and high resource environments.ConclusionsShort-term, single-species assays revealed significant differences in the allocation of competitive and defensive traits among the prey species. Based on these differences, we were, to some extent, able to predict how the long-term community structure, e.g. species dominance, is modified by the resource availability and predation interaction in pulsed resource environments. Our results are consistent with theoretical predictions and also highlight the importance of interactive effects of resource competition and predation, suggesting that these factors should not be studied in isolation.

Determinants of the detrital arthropod community structure: the effects of temperature and resources along an environmental gradient

Understanding the factors that shape community structure, and whether those factors vary geographically, has a long history in ecology. Because the abiotic environment often varies in predictable ways along elevational gradients, montane systems are ideal to study geographic variation in the determinants of community structure. In this study, we first examined the relative importance of environmental gradients, microclimate, and food resources in driving spatial variation in the structure of detrital communities in forests of the southeastern USA. Then, in order to assess whether the determinants of detrital community structure varied along a climatic gradient, we manipulated resource availability and microclimatic conditions at 15 sites along a well‐studied elevational gradient. We found that arthropod abundance and richness generally declined with increasing elevation, though the shape of the relationship varied among taxa. Overall community composition and species evenness also varied systematically along the climatic gradient, suggesting that broad‐scale variation in the abiotic environment drives geographic variation in both patterns of diversity and community composition. After controlling for the effect of climatic variation along the elevational gradient, food resource addition and microclimate alteration influenced the richness and abundance of some taxa. However, the effect of food resource addition and microclimate alteration on the richness and abundance of arthropods did not vary with elevation. In addition, the degree of community similarity between control plots and either resource‐added or microclimate‐altered plots did not vary with elevation suggesting a consistent influence of microclimate and food addition on detrital arthropod community structure. We conclude that using manipulative experiments along environmental gradients can help tease apart the relative importance and detect the interactive effects of local‐scale factors and broad‐scale climatic variation in shaping communities.

Higher parasite resistance in Daphnia populations with recent epidemics

Natural populations often show genetic variation in parasite resistance, forming the basis for evolutionary response to selection imposed by parasitism. We investigated whether previous epidemics selected for higher resistance to novel parasite isolates in a Daphnia galeata-microparasite system by comparing susceptibility of host clones from populations with varying epidemic history. We manipulated resource availability to evaluate whether diet influences Daphnia susceptibility as epidemics are common in nutrient-rich lakes. Exposing clones from 10 lakes under two food treatments to an allopatric protozoan parasite, we found that Daphnia originating from lakes (mainly nutrient rich) with previous epidemics better resist infection. Despite this result, there was a tendency of higher susceptibility in the low food treatment, suggesting that higher resistance of clones from populations with epidemic background is not directly caused by lake nutrient level. Rather, our results imply that host populations respond to parasite-mediated selection by evolving higher parasite resistance.

Intensive, site-specific silviculture: Manipulating resource availability at establishment for improved stand productivity. A review of South African research

An important window of opportunity to increase and sustain productivity in short-rotation plantations is the period from felling through re-establishment to canopy closure. This paper explores the effects, interactions and response mechanisms of intensive silvicultural practices on plantation productivity and sustainability, using five South African case studies (a–e). (a) Land preparation trials showed that complete surface cultivation by ploughing had a significant beneficial effect when afforestation is done for the first time in grasslands, improving basal area growth by 11–52% over pitting only. However, similar treatments have not resulted in significant growth responses under re-establishment conditions. (b) Stand growth suppression resulting chiefly from soil compaction during mechanised harvesting operations is strongly related to soil type, soil textural class and residue management options. Volume growth reduction in short-rotation eucalypt crops ranged from 25% on compaction sensitive loamy soils to less than 2% in resistant sandy soils. (c) The response mechanism whereby vegetation management improves stand productivity is a reduction in both inter-specific and intra-genotypic competition for resources, as well as a decrease in stand variability. Operationally, the most important criteria in a vegetation management programme relate to the timing of control operations across diverse site conditions. In local trials, the primary factors controlling the time taken for competition-induced tree growth suppression to occur were related to altitude, slash burning and the interaction between these factors, which facilitated the development of regional vegetation management strategies. (d) Empirical fertilizer trials in short-rotation hardwood stands have shown significant improvements in final productivity (commonly 20–90 m 3 ha −1 in eucalypts and 30–50 m 3 ha −1 in Acacia), as well as wood density (15–30 kg m −3 for eucalypts) following improvements in early nutrition. Improved nutrition was achieved through fertilization at planting or indirectly through residue management. The response mechanism is primarily due to early canopy development and associated increases in light capture, coupled with a more modest increase in canopy quantum efficiency and above-ground carbon allocation on a dry site. On sites with abundant water supply, increased quantum efficiency is likely to be the dominant response mechanism. (e) A series of operational gains trials tested the interactive effect of genetic tree improvement, site–genotype interaction, stand density and vegetation management + fertilization on eucalypt stand growth across five sites. There were no significant interactions between factors, but importantly, the results were additive, emphasizing the need to optimise each practice in the value chain to achieve maximum productivity.

AN EXPERIMENTAL TEST OF THE EFFECTS OF RESOURCES AND SEX RATIO ON MATERNAL FITNESS AND PHENOTYPIC SELECTION IN GYNODIOECIOUSFRAGARIA VIRGINIANA

Resources, sex ratio, and seed production by hermaphrodites covary among natural populations of many gynodioecious plant species, such that they are functionally "more dioecious" as resources become more limiting. Strong correlations among these three factors confound our understanding of their relative roles in maintaining polymorphic sexual systems. We manipulated resource availability and sex ratio and measured their effects on relative fertility and phenotypic selection through the maternal fitness of females and hermaphrodites of Fragaria virginiana. Two results were particularly surprising. First, hermaphrodites showed little variability in fecundity across resource treatments and showed strong positive and context-dependent selection for fruit set. This suggests that variation in hermaphrodite seed production along resource gradients in nature may result from adaptation rather than plasticity. Second, although females increased their fecundity with higher resources, their fertility was unaffected by sex ratio, which is predicted to mediate pollen limitation of females in natural populations where they are common. Selection on petal size of females was also weak, indicating a minimal effect of pollinator attraction on variation in the fertility of female plants. Hence, we found no mechanistic explanation for the complete absence of high-resource high female populations in nature. Despite strong selection for increased fruit set of hermaphrodites, both the strength of selection and its contribution to the maintenance of gynodioecy are severely reduced under conditions where females have high relative fecundity (i.e., low resources and high-female sex ratios). High relative fertility plus high female frequency means that the evolution of phenotypic traits in hermaphrodites (i.e., response to selection via seed function) should be manifested through females because most hermaphrodites will have female mothers. Fruit set was never under strong selection in females; hence, selection to increase fruit set hermaphrodites will be less effective in maintaining their fruiting ability in natural populations with low resources and high female frequency. In sum, both sex ratio and resource availability influence trait evolution indirectly-through their effects on relative fertility of the sexes and patterns of selection. Sex ratio did not impose strong pollen limitation on females but did directly moderate the outcome of natural selection by biasing the maternal sex of the next generation. This direct effect of sex ratio on the manifestation of natural selection is expected to have far greater impact on the evolution of traits, such as seed-producing ability in hermaphrodites and the maintenance of sexual polymorphisms in nature, compared to indirect effects of sex ratio on relative fertility of the sexes.

Environmental maternal effects on seed morphology and germination inSinapis arvensis(Cruciferae)

SummaryPioneer plants are adapted to grow in unpredictable environments. These plants have evolved several traits related to seed morphology and germination to cope with this unpredictability. The main aim of this study was to investigate the effect of plant identity and maternal environment on seed mass and germination ofSinapis arvensis. This study was undertaken withS. arvensisplants grown in four experimentally manipulated resource availability scenarios: (i) nitrogen addition, (ii) water addition, (iii) nitrogen and water simultaneously added, and (iv) control. Plants grew under field conditions in an emerging plant community. Germination response was mainly affected by maternal plant identity. This reflects not only genotype quality but also phenotypic plasticity, which allows adaptation to habitat conditions. Seed coat colour together with mean seed mass per plant explained significant fractions of the variability in germination rate. Changes in the germination response because of maternal environmental effects were also detected. Germination rate decreased in seeds from nitrogen‐enriched conditions; this may be due to the induction of dormancy caused by high concentration of nitrogen in the seeds. Addition of water to maternal environment caused a decrease in total germination percentage and germination rate of seeds. Thus, our results provide evidence for maternal environmental effects on germination under field conditions. Furthermore, changes in germination timing, as detected in our study, may determine plant development and fitness in unpredictable conditions.

Altered parasite assemblages in raccoons in response to manipulated resource availability

The role that host aggregation plays in structuring parasite assemblages was examined by experimentally increasing the contact rates of raccoons, Procyon lotor. Two populations of raccoons in southern New York were monitored for 2 years to determine baseline levels of host interaction and to identify the parasite assemblage. In the third year of the study, one population was provisioned with the addition of clumped food resources, while the other was provisioned with equal quantities of dispersed food resources. Remote photography showed that raccoons aggregated at clumped resources but not at dispersed resources, and therefore contact rates between individuals were higher in the site with clumped resources. There were no differences in parasitism between the sites prior to resource augmentation. Among ectoparasites, there were no significant changes in the prevalence or abundance of any species in response to the perturbation. In contrast, across the endoparasite assemblage within and across hosts, the prevalence of infection increased as a result of increased host contact. Strong increases in the prevalence of a few directly transmitted species and slight increases among most species lead to increased evenness in parasite prevalence, suggesting that parasites in this system are transmission limited. In addition, the number of parasite species per host (the parasite infracommunity) was higher in the clumped-resource population. These endoparasite results suggest that intraspecific variation in the species richness of parasite communities of individual hosts, and the prevalence of parasitic species in host populations as assessed across entire parasitic assemblages, is robustly influenced by intraspecific variation in the degree of host contact. Further, these results suggest that anthropogenic changes which alter resource availability may have important consequences for disease transmission in wildlife.

Sources and consequences of seed size variation inLupinus perennis(Fabaceae): adaptive and non‐adaptive hypotheses

Seed size variation within species and individuals is common. This variation may be adaptive in heterogeneous landscapes if the fitness consequences of seed size differ among environments or through time. Variation may also arise from constraints that limit control of seed size. I manipulated resource availability in both maternal and offspring environments to test conditions underlying these explanations for seed size variation in the herbaceous perennial Lupinus perennis. A fivefold variation in seed size arose primarily from differences among individuals and within-plant variability rather than from environmental conditions manipulated in the experiment. Environmental conditions had little effect on mean seed size; in contrast, within-plant variation in seed size increased with reduced resources. Fitness benefits from large seed size were similar across offspring environments, suggesting that environmental heterogeneity alone may not maintain seed size variation in this species. Surprisingly, seed size affected long-term fitness measures, including a plant's size and probability of flowering through its second year. These results are consistent with non-adaptive but not adaptive explanations for seed size variation. They also suggest that offspring size variation per se may contribute to variation in maternal fitness.

Frequent nest relocation in the antAphaenogaster araneoides: resources, competition, and natural enemies

Sessile and vagile organisms differ from one another in some fundamental ways, including methods of resource acquisition and competition. Ant colonies are typically studied as sessile entities, even though a large fraction of ant species frequently relocate their nests in the course of their life history. Little is known about the causes and consequences of nest relocation, but it is likely that the costs and benefits of relocation are driven by nest quality, neighborhood competition, or resource availability. In this paper, we document several cycles of nest relocation in a population of the Central American antAphaenogaster araneoides. In our first experiment, we tracked the pattern of relocation, testing whether environmental characteristics and colony demography were associated with relocation behavior. In our second experiment, we manipulated resource availability by adding or subtracting leaf litter, which is known to predict colony growth. We found that colonies relocated their nests once per week on average and colonies often reoccupied nests from which they had once emigrated. Larger colonies relocated more frequently than smaller colonies, and quickly growing colonies utilized a greater number of nests within their home range compared to slowly growing colonies. Relocation events were most likely to occur in periods when vapor pressure deficits were greatest. Nearest neighbor distance and other measures of environmental conditions were not associated with relocation behavior and there was no significant effect of litter removal or supplementation. We found evidence that multiple natural enemies attackedA. araneoidescolonies. Based on the demographic correlates of relocation and our rejection of other plausible hypotheses, we propose that nest relocation is driven by the escape from natural enemies.